click for previous page 82 FAO Species Catalogue Vol. 16 13a. Juveniles brown, with 7 dark brown bars on body, the last covering most of caudal peduncle, its upper half black; distance between fourth and fifth dark bars greater than that between third and fourth and fifth and sixth dark bars; adults pinkish grey without bars, the fins darker except for broad pale margins on caudal fin and soft dorsal fin; skin of body wrinkled in a narrow zone along base of dorsal fin; no auxiliary scales on body: no scales on maxilla (Fig. 167, Plate XII ) (eastern Australia from 18º to 36ºS)... E. ergastularius 13b. Juveniles greyish brown, with 3 faint dark bars on dorsal part of body and a dark brown saddle spot on peduncle; large adults grey-brown; no wrinkles on body at base of dorsal fin; adults with auxiliary scales; small embedded scales on maxilla (Fig. 168, Plate X) (Kenya to South Africa)... E. chabaudi skin along dorsal-fin base wrinkled soft dorsal-fin and caudal-fin margin pale no scales on maxilla Fig. 167 Epinephelus ergastularius small embedded scales on maxilla Fig. 168 Epinephelus chabaudi auxiliary scales 14a. Dorsal-fin rays 14; lateral-line scales 56 to 66; body and rear of head with well-separated dark brown spots; prominent dark brown maxillary streak; soft dorsal and anal fins dusky distally, with prominent white edge; margin of interspinous dorsal-fin membranes usually orange-yellow (Fig. 169, Plate VII) (southern Mozambique to South Africa)... E. albomarginatus 14b. Dorsal-fin rays 16 or 17; lateral-line scales 64 to 76; colour not as in 14a (brown spots, if present, close-set)... 15 well separated dark brown spotsllll ll14 dorsal-fin rays dark brown maxillary streak 56-66 lateral-line scales Fig. 169 Epinephelus albomarginatus 15a. Head, body and fins with numerous close-set small dark brown spots; rear margin of caudal fin with white edge; 2 rows of teeth on midside of lower jaw (Fig. 170, Plate XX) (Gulf of Aden and Persian Gulf to India)... E. polylepis 15b. No dark spots on head, body or fins; no white margin posteriorly on caudal fin; 3 or 4 rows of teeth on midside of lower jaw.... 2 rows of teeth on midside close-set small dark spots white of lower jaw caudal-fin margin 16 Fig: 170 Epinephelus polylepis
Groupers of the World 83 16a. Caudal fin truncate to slightly convex; nostrils subequal; pectoral-fin rays 19 to 20; caudal peduncle depth 3.2 to 3.8 times in head length; no scales on maxilla; margin of interspinous dorsal-fin membranes dark red to black (Fig. 171, Plate XXI) (Indo-Pacific)... E. retouti 16b. Caudal fin slightly emarginate; posterior nostrils of adults more than twice diameter of anteriors; pectoral-fin rays 18; caudal-peduncle depth 3.9 times in head length; maxilla with small scales; no conspicuous markings on head, body or spinous dorsal fin (Fig. 172, Plate XI) (1 specimen, 53 cm standard length, Timor Sea)... E. darwinensis 19-20 pectoral-fin rays posterior nostrils more than twice 18 pectoral-fin rays nostrils subequal diameter of anteriors caudal-peduncle depth 3.2-3.8 times in head length Fig. 171 Epinephelus retouti caudal-peduncle depth 3.9 times in head length Fig. 172 Epinephelus darwinensis 17a. 17b. Anal-fin rays 9 (rarely 10).... -> 18 Anal-fin rays 8 (rarely 7 or 9)... 21 18a. Dorsal-fin rays 16; scales smooth and mostly embedded; lateral-scale series 87 to 108; body depth 3.15 to 3.6 times in standard length; nostrils subequal; interorbital space flat; pale with reddish brown spots on head and body; interspinous dorsal-fin membranes with alternating dark brown and white stripes (Fig. 173, Plate XXI) (Madagascar, Mozambique, and South Africa)... E. posteli 18b. Dorsal-fin rays 14 to 15; scales on body ctenoid (rough) and mostly exposed; lateral-scale series 103 to 139; body depth 2.3 to 3.0 times in standard length; rear nostrils of adults vertically ovate, their diameter 2 or more times larger than anterior nostrils; interorbital space convex; no small dark spots on head or body; no dark stripes in dorsal fin... 19 19a. 19b. nostrils subequal No small spinules on ventral edge of preopercle; lateral-scale series 122 to 139; juveniles dark greyish brown, with 8 vertical series of white spots on body (Fig. 174, Plate XXI) (Hawaiian Islands)... E. quernus Lower edge of preopercle with 1 to 4 serrae anterior to serrae at corner; juveniles with dark bars on body; lateral-scale series 103 to 128... 20 lateral-scale series 87-108 16 dorsal-fin rays lateral scale series 122-139 body depth 3.15-3.6 times in standard length Fig. 173 Epinephelus posteli no small spinulus on ventral edge of preopercle Fig. 173 Epinephelus quernus
84 FAO Species Catalogue Vol. 16 20a. Body with 6 dark bars below dorsal fin, the last 2 bars close set and much narrower than preceding bar; 3 pale interspaces below soft dorsal fin; greatest diameter of rear nostrils more than distance from nostril to eye; ratio of caudal-peduncle length/depth = 1.2-1.5 (Fig. 175, Plate XXII) (Japan, Korea, and China)... E. septemfasciatus 20b. Body with 5 dark bars below dorsal fin, the last 2 bars as broad as preceding bars; 2 pale interspaces below soft dorsal fin; rear nostril diameter not more than distance from nostril to eye: ratio of peduncle length/depth = 0.9-1.2 (Fig. 176, Plate XX) (Indo-Pacific but not in Red Sea or Persian Gulf, in deep water)... E. octofasciatus greatest diameter of rear nostril more than distance from nostril to eye 6 bars below dorsal fin 5 bars below dorsal fin ratio of caudal-peduncle length/depth = 1.2-1.5 Fig. 175 Epinephelus septemfasciatus ratio of caudal-peduncle Fig. 176 Epinephelus octofasciatus 21a. Dorsal-fin rays 12 or 13 (rarely 13).... 22 21b. Dorsal-fin rays 14 to 18 (rarely 13 in E. bruneus)... 23 22a. Lateral-line scales 52; lateral-body scales ctenoid (rough); maxilla naked; rear nostrils twice as large as anterior nostrils; no obvious dark or light markings (Fig. 177) (1 specimen, 465 mm standard length, from Queensland, Australia)... E. perplexus 22b. Lateral-line scales 56 to 65; lateral-body scales smooth; small scales usually present on maxilla; nostrils subequal; juveniles (<20 cm standard length) with 2 broad, longitudinal, black-edged whitish bands that disappear in adults, the dark edges breaking into dashes and spots (may be lost in large adults) (Fig. 178, Plate XVI) (Red Sea to southern Japan)... E. latifasciatus rear nostrils twice as large as anterior ones 52 lateral-line scales nostrils subequal 56-65 lateral-line scales maxilla without scales lateral-body scales ctenoid Fig. 177 Epinephelus perplexus lateral-body; scales smooth adults with dark lines, breaking into dashes and spots juveniles with 2 black-edged bands Fig. 178 Epinephelus latifasciatus
llllllllllll Groupers of the World 85 23a. 23b. Lateral-line scales with branched tubules; eye diameter contained about 8 times in head length at 20 cm standard length, about 9 times in head length at 35 cm, and 13 times in head length for 145 cm fish; maximum total length about 2.7 m; juveniles yellow, with 3 broad black bars on body and irregular broad black bands on head (Fig. 179, Plate XVI) (Indo-Pacific)... E. lanceolatus Lateral-line scales with a single tubule (except anterior scales of large E. brunneus, E. coioides and E. malabaricus); eye diameter contained less than 7 times in head length of 20 cm fish and less than 8 times in head length for 35 cm fish; maximum total length less than 1.5 m (except E. tukula, which may reach 2 m); juveniles not coloured as in 23a... 24 eye diameter contained 13 times in head length (at 145 cm standard length) Fig. 179 Epinephelus lanceolatus lateral-line scales with branched tubules 24a. Numerous distinct dark spots (not dots) over most of head and body (spots brownish red to black in life and distinct in preservative) (see Fig. 180)... 25 24b. No distinct dark spots over most of head and body (there may be yellow or orange spots in life, but these usually do not persist in alcohol; or there may be dark dots or scattered small dark spots, but not over most of head and body) (see Figs 209,210).... 49 25a Lateral-line scales 46 to 53... 26 25b. Lateral-line scales 53 to 74... 37 26a. Midlateral-body scales smooth, at least on adults (may be ctenoid in area covered by pectoral 26b. Midlateral-body scales rough (mainly smooth on large adults of E. andersoni and E. socialis)... 30 27a. Dorsal profile of head nearly straight; orange-red to brownish red spots on head, body, and median and pelvic fins; 3 or 4 prominent black spots or blotches dorsally on body, the first (and largest, greater than orbit diameter) at base of middle of dorsal fin, the last blotch dorsally on caudal peduncle; no blackish maxillary streak (Fig. 180, Plate XXIV) (Korea and southern Japan to Taiwan and China).... E. trimaculatus 27b. Dorsal profile of head convex; spots on head, body, and fins dark brown to black; black blotches on body at base of dorsal fin present or absent; blackish maxillary streak usually present... 28 dorsal profile of head nearly straight 3 or 4 black blotches dorsally no blackish maxillary streak Fig. 180 Epinephelus trimaculatus
86 FAO Species Catalogue Vol. 16 28a. 28b. Dorsal-fin rays 16 to 18; dark spots on pectoral fins progressively smaller distally; pelvic-fin length 2.2 to 2.6 times in head length (Fig. 181, Plate XIII) (India, Sri Lanka and Indonesia)... E. faveatus Dorsal-fin rays 15 to 17; no dark spots on distal part of pectoral fins; pelvic-fin length 1.9 to 2.3 times in head length (at standard length less than 25 cm).... 29 16 18 dorsal-fin rays pelvic-fin length 2.2-2.6 times in head length lllllll dark spots on pectoral fin smaller distally 29a. 29b. Large blackish blotch (containing 4 to 7 black spots) on body at base of last 3 dorsal-fin spines; dark spots on head, body, and fins numerous, 17 on soft dorsal fin of a 10 cm standard length fish and 28 dark spots on the fin of a 28 cm fish; pectoral-fin rays 17 to 19 (rarely 19); total gill rakers 23 to 26 (modally 25); body width 1.7 to 2.2 times in body depth; lower jaw not strongly projecting (Fig. 182, Plate XV) (Oceania and western Pacific)... E. howlandi No large blackish blotch at base of posterior dorsal-fin spines (though 1 to 3 spots at this location are usually darker than adjacent spots); dark spots on head, body, and fins less numerous, 5 to 7 on soft dorsal fin of a 11 cm fish and 20 to 21 dark spots on the fin of a 29 cm fish; pectoral-fin rays 18 or 19 (often 19); total gill rakers 21 to 24 (modally 23); body width 1.4 to 1.8 times in body depth; lower jaw strongly projecting (Fig. 183, Plate XVII) (Indo-Pacific but not in Red Sea or Persian Gulf)... E. macrospilos numerous spots on head, body, and fins large blotch at base of last 3 dorsal-fin spines lower jaw strongly projecting spots on head, body, and fin less numerous body width 1.7-2.2 times in body depth 17-19 pectoral-fin rays Fig. 182 Epinephelus howlandi body width 1.4-1.8 times in body depth 18-19 pectoral-fin rays Fig. 183 Epinephelus macrospilos 30a. 30b. Dark spots on body elongate, oblique, and more numerous posteriorly; dark spots on head separated by more than 2 spot diameters (Fig. 184); ventral edge of maxilla of subadults (33 to 40 cm standard length) a distinct step-like shape that develops into a bony knob in adults (Fig. 185, Plate XVII) (Indian Ocean and Indonesia)... E. longispinis Dark spots on body not elongate and not more numerous posteriorly; dark spots on head separated by less than a spot diameter (except E. bontoides); no distinct step-like shape to ventral edge of maxilla (Fig. 185)... 31 spots separated by more than 2 spots diameters dark spots elongate and oblique ventral edge of maxilla with a step ventral edge of maxilla with "step" Fig. 184 Epinephelus longispinis no step on ventral edge of maxilla Fig. 185
Groupers of the World 87 31a. Pectoral-fin length 1.2 to 1.6 times in head length (except fish from northwest Australia); dorsal-fin rays 16 to 18; serrae at corner of preopercle distinctly enlarged (Fig. 186); 2 oblique dark brown bands (or elongate dark blotches linked by narrow bands) on chest (Fig. 187, Plate XXI) (Andaman Islands and western Pacific).... E. quoyanus 31b. Pectoral-fin length 1.5 to 2.0 times in head length; dorsal-fin rays 14 to 17; serrae at corner of preopercle slightly enlarged (Fig. 186); no oblique dark bands on chest... 32 pectoral-fin length 1.2-l.6 times in head length 16-18 dorsal-fin rays serraeat corner of serrae at corner of pre- opercle distinctly enlarged serrae at corner of pre- preopercle opercle slightly enlarged distinctly enlarged 2 oblique dark bands on chest Fig. 186 Fig. 187 Epinephelus quoyanus 32a. 32b. Lateral-scale series 82 to 86; pectoral-fin rays 18 or 19; total gill rakers 20 to 22; head and body with small dark spots separated by spaces greater than 2 spot diameters; no dark spots on ventral parts of head and body; caudal, soft dorsal, and pectoral fins blackish brown with a pale yellow to white margin (Fig. 188, Plate IX) (Taiwan to Indonesia and New Britain).... E. bontoides Lateral-scale series 92 to 120; pectoral-fin rays 16 to 19; total gill rakers 23 to 27 (except E. merra with 21 to 25); dark spots on head and body extending ventrally, and most within a spot diameter of adjacent spots; fins not dark brown to black with a distinct pale margin... 33 33a. 33b. Dorsal-fin rays 14 or 15; longest dorsal-fin spine 2.7 to 3.3 times in head length: a black saddle-blotch on caudal peduncle; head and body covered by small orange-brown to dark brown spots; head and dorsal part of body with irregular dark brown blotches superimposed over the dark spots (Fig. 189, Plate XX) (Indo-Pacific).... E. polyphekadion Dorsal-fin rays 15 to 18; longest dorsal-fin spines 2.1 to 2.7 times in head length (except E. merra with 2.6 to 2.9 times in head length); colour not as in 33a (especially, no black saddle on caudal peduncle).... lateral-scale series 82-86 caudal, soft dorsal and pectoral fins with palemargin longest dorsal-fin spine 2.7-3.3 times in head length 34 14-15 dorsal-fin rays black saddle blotch on caudal peduncle 20-22 gill rakers in total 18-19 pectoral-fin rays Fig. 188 Epinephelus bontoides Fig. 189 Epinephelus polyphekadion
88 FAO Species Catalogue Vol. 16 34a. Dark spots on head and body less than half size of pupil; dark brown to black spots on fins (except spinous dorsal fin) much larger than those on body (Fig. 190, Plate XVIII); maxilla reaching a vertical at rear edge of orbit (Fig. 191) (Indo-Pacific but not in Red Sea)... E. miliaris 34b. Dark spots on body larger than pupil; dark spots on fins smaller than those on body; maxilla usually reaching to or past a vertical at rear edge of orbit (Fig. 191).... 35 dark spots on head spots on fins much larger maxilla reaching to or past a and body small than those on body vertical at rear edge of orbit 35a. 35b. Fig. 190 Epinephelus miliaris Fig. 191 Pectoral-fin rays 16 to 18; longest dorsal-fin spine 2.6 to 2.9 times in head length; some dark brown spots on body often joined to form short horizontal or oblique bands; black spots on pectoral fins very small and largely confined to rays (Fig. 192, Plate XVIII) (Indo-Pacific but not in Red Sea).... E. merra Pectoral-fin rays 17 to 19; longest dorsal-fin spines 2.1 to 2.6 times in head length; no confluent dark spots on body; dark spots on pectoral fins not uniformly small and not confined to rays... 36 longest dorsal-fin spine 2.6-2.9 times in head length 16-18 pectoral-fin rays Fig. 192 Epinephelus merra 36a. Dorsal-fin rays 17 or 18; lateral-scale series 94 to 102; 3 close-set pairs of dark brown to black spots on body at base of dorsal fin (Fig. 193) (Western Australia)... E. bilobatus 36b. Dorsal-fin rays 15 to 17; lateral-scale series 102 to 120; no pairs of dark brown to black spots along base of dorsal fin; 2 large dusky to blackish areas on body and dorsal fin, these separated by a broad whitish (though still faintly spotted) area (Fig. 194, Plate XVII) (central and western Pacific)... E. maculatus 3 close-set pairs of dark spots along dorsal-fin base 17-18 dorsal-fin rays 2 large dusky areas on 15-17 body and dorsal fin dorsal-fin rays lateral-scale series 94-102 Fig. 193 Epinephelus bilobatus Fig. 194 Epinephelus maculatus click for next page