Epinephelus Drummondhayi Goode and Bean, 1878 EED Frequent Synonyms / Misidentifications: None / None

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1340 Bony Fishes

Epinephelus drummondhayi Goode and Bean, 1878 EED Frequent synonyms / misidentifications: None / None. FAO names: En - Speckled hind; Fr - Mérou grivelé; Sp - Mero pintaroja.

Diagnostic characters: Body depth subequal to head length, 2.4 to 2.6 times in standard length (for fish 20 to 43 cm standard length). Nostrils subequal; preopercle rounded, evenly serrate. Gill rakers on first arch 9 or 10 on upper limb, 17 or 18 on lower limb, total 26 to 28. Dorsal fin with 11 spines and 15 or 16 soft rays, the membrane incised between the anterior spines; anal fin with 3 spines and 9 soft rays; caudal fin truncate or slightly emarginate, the corners acute; pectoral-fin rays 18. Scales strongly ctenoid, about 125 lateral-scale series; lateral-line scales 72 to 76. Colour: adults (larger than 33 cm) dark reddish brown, densely covered with small pearly white spots; juveniles (less than 20 cm) bright yellow, covered with small bluish white spots. Size: Maximum about 110 cm; maximum weight 30 kg. Habitat, biology, and fisheries: Adults inhabit offshore rocky bottoms in depths of 25 to 183 m but are most common between 60 and 120 m.Females mature at 4 or 5 years of age (total length 45 to 60 cm).Spawning occurs from July to September, and a large female may produce up to 2 million eggs at 1 spawning. Back-calculated total lengths for fish aged 1 to 15 years are 19, 32, 41, 48, 53, 57, 61, 65, 68, 71, 74, 77, 81, 84, and 86 cm; the maximum age attained is at least 25 years, and the largest specimen measured was 110 cm. The weight/length relationship isW=1.1X10-8L3.073 for W in kg and L (total length) in mm.The von Bertalanffy growth equation is L t = 967(1-e- 0.13 ( t+1.01)). Adults feed on a variety of fishes and invertebrates, including shrimps, crabs, squid, and octopus. An important species in the recreational and commercial fisheries of the southeastern USA and also in the grouper fishery of the Campeche Bank off Yucatán. Mainly caught with hook-and-line, but some might also be taken in traps, trawls, and on bottom-set longlines. Distribution: Bermuda, North Carolina to Florida Keys, Gulf of Mexico. Reports of speckled hind from Cuba and the Bahamas are unsub- stantiated. Remarks: The speckled hind is also known as ‘Kitty Mitchell’ in the Gulf of Mexico region; and the Mexican name is ‘lenteja’. Perciformes: Percoidei: Serranidae 1341

Epinephelus flavolimbatus Poey, 1865 EEL Frequent synonyms / misidentifications: None / None. FAO names: En - Yellowedge grouper; Fr - Mérou aile jaune; Sp - Mero aleta amarilla.

Diagnostic characters: Body depth distinctly less than head length, 2.7 to 2.9 in standard length (for fish 13 to 64 cm standard length).Eye diameter equal to or greater than interorbital width in fish less than 45 cm standard length. Nostrils subequal. Gill rakers on first arch 8 or 9 on upper limb, 15 to 17 on lower limb, total 23 to 25. Dorsal fin with 11 spines and 13 to 15 soft rays, third or fourth spine longest, and fin membrane slightlly incised between anterior spines; anal fin with 3 spines and 9 soft rays; rear margin of caudal fin convex in fish less than 30 cm standard length, truncate or even slightly concave in larger fish; pectoral-fin rays 18. Preopercle angular, with serrae at angle distinctly enlarged and, in large fish, co- alesced into a flat serrate lobe. Lateral body scales ctenoid (rough); lateral scale series 82 to 99; lateral-line scales about 65. Colour: head and body buff or greyish brown, whitish ventrally; margins of dorsal and pectoral fins, and sometimes anal and caudal fins yellow; a prominent blue line from eye to corner of preopercle. Ju- veniles less than 20 cm standard length, with pearly spots arranged in a grid of 4 longitudinal rows and 7 vertical columns and a dark brown saddle blotch on caudal peduncle; dorsal fin with broad yellow margin; caudal fin white; anal and pelvic fins blackish; black saddle on peduncle (if present) ending abruptly at lateral line; this spotted juvenile pattern gradually disappears with growth and is faint to absent in fish larger than 30 cm standard length. Adults generally immaculate, but sometimes (momentarily) they display the white-spotted grid pattern. Size: Maximum about 115 cm; maximum weight 20 kg. Habitat, biology, and fisheries: A deep-water species occurring in rocky areas and on sand/mud bottoms in depths of 64 to 370 m. On soft bottoms, often seen in or near trenches or burrow-like excavations. Females attain maturity at 53 to 60 cm total length and are thought to change sex at about 75 cm. Spawning occurs in the Gulf of Mexico from May to September. The maximum age is at least 20 years. The relationship of weight to length for yellowedge grouper in the eastern Gulf of Mexico is W = 5.37 x 10-8 standard length2.872 (n = 472, r2 = 0.99) where W is whole weight in kg and standard length is in millimetres. This species feeds on a wide variety of invertebrates (mainly brachyuran crabs) and fishes. One of the 2 most important species of groupers in the deep-water longline fishery in the eastern Gulf of Mexico; also of some importance in sport and commercial fisheries off the southeastern coast of the USA and along the coast of Central and South America. Distribution: North Carolina to southern Brazil, including Gulf of Mexico and Caribbean; not reported from Bermuda. Remarks: The local name in Mexico is “’cherna del alto’. 1342 Bony Fishes

Epinephelus guttatus (Linnaeus, 1758) EEU Frequent synonyms / misidentifications: None / None. FAO Names: En - Red hind; Fr - Mérou couronné; Sp - Mero colorado.

Diagnostic characters: Body depth distinctly less than head length, 2.7 to 3.1 in standard length (for fish 17 to 38 cm standard length). Gill rakers on first arch 8 or 9 on upper limb and 16 to 18 on lower limb, total 24 to 26. Dorsal fin with 11 spines and 15 or 16 soft rays, the third or fourth spine longest, the interspinous membranes incised and produced into a short flag behind tip of each spine; anal fin with 3 spines and 8 soft rays; pectoral-fin rays 16 to 18; rear edge of caudal fin convex. Preopercle rounded, evenly serrate; rear nostril larger than front nostril. Scales ctenoid, lateral scale series 92 to 104. Colour: ground colour buff, greenish white, or pale reddish brown, the head and body covered with bright red spots which become reddish brown dorsally; spinous dorsal fin olive, with yellow flags at tips of spines; soft dorsal, caudal, and anal fins olivaceous, with a broad blackish submarginal band and narrow pale edge; pectoral fins pale orange-red with darker red spots on base; pelvic fins coloured like body but darker distally and along leading edge. Size: Maximum 76 cm; maximum weight 8.3 kg. Habitat, biology, and fisheries: Shallow reefs and rocky bottoms in depths of 2 to 100 m. Red hind feed mainly on crabs (40%) and fishes (21%). Females mature at 22 to 24 cm total length, and change sex at 28 to 38 cm total length.On the outer reef in 20 m off the south coast of Puerto Rico, spawning of fish in aggregations occurred during the full moon in January and February. Ripe females were recognized by their swollen abdo- mens and colour pattern of dark spots on a white background; males displayed a darker mottled pattern, with an area of dark vertical bars or squares on the body above the anal fin. Females rested on or close to the bottom, while males patrolled an area that included 1 to 5 females and defended this territory from other males. On 2 occasions, spawning was initiated by a female swimming about 0.5 m up off the bottom and being joined by a male; gametes were released without any upward rush or rapid movement. In one case, another female joined the pair above the bottom and spawned with them. The transparent eggs are buoyant and usually con- tain a single oil globule. For larvae reared in the laboratory, hatching occurred 27 h after fertilization at 26.5° C; mortality increased greatly after 6 to 7 days, and no larva survived through metamorphosis. Fecundity varies from 90 000 eggs for a 26 cm (total length) fish to over 3 million eggs in a 45 cm (total length) female. Although not as large as some other groupers, it is the most important species in the Caribbean grouper fishery. Caught with hook-and-line, traps, and spears. Distribution: Bermuda, North Carolina to Brazil, including Gulf of Mexico and the West In- dies. Remarks: In the West Indies, the red hind is also called ‘cabrilla morja’ or ‘strawberry grouper’; in Venezuela, it is known as ‘tofia’. Perciformes: Percoidei: Serranidae 1343

Epinephelus itajara (Lichtenstein, 1822) EET Frequent synonyms / misidentifications: Promicrops itajara (Lichtenstein, 1822) / None. FAO names: En - Goliath grouper (AFS: Jewfish); Fr - Mérou géant; Sp - Mero guasa.

Diagnostic characters: Body robust, oblong, the greatest width equals half or more of body depth, which is less than head length (in fish 15 to 160 cm). Head extremely broad; interorbital flat and very wide, its width equals eye diameter in fish 10 to 15 cm standard length, is distinctly greater than eye diameter in fish 18 to 30 cm standard length, and 2 to 5 times greater than eye diameter in fish 30 to 160 cm standard length. Eye small, its diameter contained 5 to 9 times in head length for fish 10 to 30 cm standard length and 12 times in head of 160 cm standard length fish.Preopercle subangular, finely serrate;nostrils subequal.Maxilla reaching well past eye. Gill rakers 8 or 9 on upper limb, 13 to 15 on lower limb, total 21 to 24. Dorsal fin with 11 spines and 15 or 16 soft rays, third to eleventh spines subequal and distinctly shorter than longest dorsal rays; anal fin with 3 spines and 8 soft rays; caudal fin rounded; pectoral-fin rays 18 or 19. Body scales strongly ctenoid, 89 to 110 lateral-scale series; lateral-line scales 61 to 64. Colour: generally brownish yellow, grey or greenish; dorsal part of head, body, and fins with small black spots. Fish less than about1mhave5ir- regular oblique bars on body; large adults darker and more uniformly coloured than juveniles. Size: The largest Atlantic grouper, attaining 250 cm and a weight of at least 400 kg. Habitat, biology,and fisheries: Large adults often found in shallow water, and also offshore on wrecks and in areas of high relief; juveniles common in mangrove swamps and both juveniles and adults occur in bays and harbours.In the eastern Gulf of Mexico, spawning occurred from July through September.Males mature at 110 to 115 cm, at age 4 to 6 years and live at least 26 years; females mature at 120 to 130 cm, at age 6 or 7 and live at least 37 years. The weight/length relationshipW=1.31x10-8 total length3.056 where W (whole weight) is in kilograms and length is in millimetres. Unlike most other groupers, there is no evidence for hermaphroditism. Adults appear to occupy limited home ranges with little inter-reef movement, and the same individuals were seen at specific reef sites for more than a year. Adults and juveniles feed heavily on crustaceans (shrimps, crabs, and lobsters), and in the Caribbean region it is an important predator of lobsters, also fishes and young sea turtles. With their slow growth, longevity, site specific spawning aggregations, and vulnerability to spearfishing, jewfish are very susceptible to overfishing. Consequently, catching of jewfish in USA Exclusive Economic Zone waters is now prohibited. Reported as very common in Vene- zuela. Caught with hook-and-line, spears, traps, and trawls. Distribution: Tropical and subtropical waters of Atlantic and eastern Pacific oceans. In the western Atlantic, from Florida and Gulf of Mexico to southern Brazil, including most of the West In- dies and probably Bermuda; in the eastern At- lantic, reported (as Epinephelus esonue) from Senegal to the Congo; in the eastern Pacific, from Gulf of California to Peru. 1344 Bony Fishes

Epinephelus morio (Valenciennes 1828) GPR Frequent synonyms / misidentifications: None / None. FAO names: En - Red grouper; Fr - Mérou rouge; Sp - Mero americano.

Diagnostic characters: Body depth less than head length, 2.6 to 3.0 times in standard length (for fish 13 to 26 cm standard length). Gill rakers on first arch 8 or 9 on upper limb, 15 or 16 on lower limb, total 23 to 25. Dor- sal fin with 11 spines and 16 or 17 soft rays, the interspinous membranes not incised, the second or third spine longest, giving the fin a triangular sail-like aspect. Anal fin with 3 spines and 8 to 10 soft rays; pectoral-fin rays 16 to 18. Caudal fin convex in fish less than 15 cm standard length, truncate, or slightly concave in larger fish. Preopercle subangular, the serrae at angle slightly enlarged; upper edge of operculum straight. Nostrils subequal. Scales ctenoid (rough), about 60 lateral-line scales and 112 to 128 lateral-scale series. Colour: head and body dark reddish brown, shading to pink or reddish below; soft dorsal, caudal, and anal fins dark distally, with a narrow white edge; a few dark dots on snout and/or cheeks; body often with irregular white spots and/or large pale blotches; inside of mouth bright reddish orange. Size: Maximum at least 90 cm and 20 kg. Habitat, biology, and fisheries: Juveniles (20 to 40 cm standard length) commonly found in crevices and un- der ledges on rocky reefs in 5 to 25 m. At 40 to 50 cm standard length (4 to 6 years of age), become mature females and begin to migrate to deeper water (50 to 300 m) where they also occur over sandy or mud bottoms. Most females transform to males between ages 7 and 14, and maximum age is at least 25 years. The von Bertalanffy growth equation is L t = 672(1-e-0.179 ( t+0.449)). The length-weight function for Gulf of Mexico fish is W=5.42x10-8 L2.897 where W is whole weight in kilograms and L is standard length in millimetres.The eastern Gulf of Mexico population spawns during April and May, and fecundity ranged from 312 000 to 5 735 700 eggs per female. After 30 to 40 days (20 to 25 mm standard length), the pelagic postlarvae transform to the benthic juvenile stage. Juveniles of 3 to 20 cm standard length are occasionally found on shallow grass beds and inshore reefs. Adults feed on a wide variety of fishes and invertebrates. Red groupers are particularly susceptible to the toxin of red tide (Ptychodiscus brevi) blooms, and in 1971 the species was exterminated on reefs in 12 to 15 m off Sarasota, Florida.The red grouper is the most important of the commercial reef fishes caught off the coast of Florida. On the Campeche Bank off the Yucatán Peninsula, red grouper made up 90% of the total catch of Cuban otter trawl fisheries. FAO statis- tics report landings of 86 to 276 t between 1995 and 1999. Also of some commercial importance on the coast of Venezuela, where it is known as ‘mero paracamo’. The species is caught with hook-and-line, bottom set longlines, traps, and trawls. Distribution: Bermuda, North Carolina to southern Brazil, including the Gulf of Mexico and Ca- ribbean; stray juveniles occur north to Massachusetts. Remarks: In Mexico, the red grouper is known as ‘cherna americana’ or ‘cherna de vivero’. Perciformes: Percoidei: Serranidae 1345

Epinephelus mystacinus (Poey, 1852) EEY Frequent synonyms / misidentifications: None / Epinephelus octofasciatus Griffin, 1926. FAO names: En - Misty grouper; Fr - Mérou brouillard; Sp - Mero listado.

Diagnostic characters:Body depth contained 2.4 to 2.9 times, head length 2.3 to 2.5 times in standard length (for fish 14 to 40 cm standard length).Eye diameter greater than interorbital width for fish less than 30 cm standard length, but less than interorbital in fish larger than 40 cm standard length. Posterior nostrils greatly enlarged, their diameter 4 or more times that of anterior nostrils. Preopercle angular, but the corner rounded. Dorsal fin with 11 spines and 14 or 15 soft rays, third spine longest and fin membrane distinctly incised between spines; anal fin with 3 spines and 9 soft rays; rear margin of caudal fin convex or truncate with rounded corners; pectoral-fin rays 18 or 19; pelvic fins shorter than pectorals and inserted anterior to lower end of pectoral-fin base. Body scales distinctly ctenoid (rough), without auxiliary scales (minute scales atttached to base of main body scales); lateral-line scales 58 to 69; lateral-scale series 99 to 112. Py- loric caeca very numerous. Gill rakers on first arch 8 or 9 on upper limb, 14 to 16 on lower limb, total 22 to 25. Colour: head and body buff, with 8 or 9 dark, subvertical bars on body, the first on nape, the last 2 (which are darker than other bars) may be fused to form a wide dark band around caudal peduncle. Prominent blackish brown moustache mark on cheeks at upper edge of maxilla. Size: Maximum at least 115 cm; and over 54 kg. Habitat, biology, and fisheries: A deep-water species reported from depths of 100 to 500 m; juveniles occasionally found in water as shallow as 30 m. Generalized carnivore feeding on fishes, crustaceans, and squid. Virtually nothing is known of the growth or repro- duction of this species; females attain at least 100 cm.E. mystacinus is the dominant grouper in the deep-water sport and commercial fisheries of the Virgin Islands. Distribution: Bermuda, North Carolina to Florida, Gulf of Mexico, the West Indies to Trini- dad; probably northern coast of South America. Remarks: Reports of ‘Epinephelus mystacinus’ from Japan were based on the Indo-Pacific species Epinephelus octofasciatus. Remarks: In Puerto Rico this grouper is called ‘cherna del alto’. 1346 Bony Fishes

Epinephelus nigritus (Holbrook, 1855) ELG Frequent synonyms / misidentifications: None / None. FAO names: En - Warsaw grouper; Fr - Mérou varsovie (= Mérou polonais); Sp - Mero negro.

Diagnostic characters: Body depth less than head length, 2.4 to 2.6 times in standard length (for fish 11 to 58 cm standard length); head length 2.1 to 2.5 times in standard length. Nostrils subequal, or rear nostril slightly larger; maxilla reaches well past eye; adults with 4 or 5 rows of teeth at midside of lower jaw; juveniles with 2 or 3 rows. Interorbital area distinctly convex, wider than eye diameter for fish larger than 15 cm standard length; preopercle corner rounded, with slightly enlarged serrae and 1 or 2 small spines on lower edge just in front of corner; interopercle and subopercle smooth. Gill rakers on first arch 9 to 11 on upper limb, 14 to 16 on lower limb; total 23 to 25. Dorsal fin with 10 spines and 13 to 15 soft rays, the second spine distinctly elongated (in fish greater than 40 cm standard length) and the interspinous membranes deeply incised; anal fin with 3 spines and 9 soft rays; rear margin of caudal fin convex or truncate with rounded corners;pectoral-fin rays 18 or 19; pelvic fins longer than pectoral fins in fish 7 to 58 cm standard length and inserted in front of vertical from lower end of pectoral-fin base. Midlateral body scales distinctly ctenoid (rough), with auxiliary scales in adults; lateral-line scales 62 to 71; lateral scale series 99 to 107. Pyloric caeca very numerous, in a large dendritic mass. Colour: dark reddish brown or brownish grey to almost black dorsally, dull reddish grey below. Juveniles with yellow caudal fin and a few randomly scattered whitish spots on body; no dark saddle blotch on peduncle. Size: Maximum total length about 235 cm; maximum weight 200 kg. Habitat, biology, and fisheries: Adults usually on rough, rocky bottom in depths of 55 to 525 m; juveniles occasionally seen on jetties and shallow-water reefs. Adults feed on a variety of crabs, shrimps, lobsters, and fishes. A long-lived, slow-growing species that may reach an age of 41 years. Average total lengths for fish aged 1, 5, 10, 25 and 41 years are 30 cm, 92 cm, 119 cm, 188 cm, and 233 cm respectively.The von Bertalanffy growth equation is L t = 2394(1-e-0,0544 (t+3616)) where L t is total length in mm at age t.Because of its large size, of considerable importance in the sport fishery for bottom fishes, but it contributes less than 1% by weight of commercial grouper landings on the southeast USA coast. In the 1983 and 1984 Florida west coast commercial fishery, warsaw grouper landings were about 55 t. The species is also common in the western Gulf of Mex- ico and southern Caribbean (from Venezuela to French Guiana). Caught mainly with hook-and-line and bottom longlines. Distribution: Massachusetts to Gulf of Mexico, Cuba, Haiti, Trinidad, Venezuela to Rio de Ja- neiro; also known (but rare) on the coast of France. Reports of E. nigritus from the eastern Pacific are apparently based on misidentifications of Epinephelus exsul (Fowler, 1944). Remarks: Epinephelus exsul differs from E. nigritus in having 8 anal-fin rays, 87 to 92 lateral- scale series, body depth 2.3 times in standard length, and pelvic fins equal to or shorter than pectoral fins. Perciformes: Percoidei: Serranidae 1347

Epinephelus niveatus Valenciennes, 1828 EFV Frequent synonyms / misidentifications: Alphestes scholanderi Walters, 1957 / None. FAO names: En - Snowy grouper; Fr - Mérou neige; Sp - Cherna pintada.

Diagnostic characters: Body depth 2.5 to 2.8, head length 2.2 to 2.4 times in standard length (for fish 15 to 27 cm standard length). Preopercle with enlarged serrae at the angle; upper edge of operculum very convex; posterior nostril 2 to 5 times larger than anterior nostril. Dorsal fin with 11 spines and 13 to 15 soft rays, the membrane distinctly incised between spines; anal fin with 3 spines and 9 soft rays. Rear margin of caudal fin convex in juveniles (5 to 30 cm standard length), straight or concave in adults; pectoral-fin rays 18 or 19; juveniles with pelvic fins longer than pectoral fins. Midlateral body scales rough; lateral-line scales about 65. Gill rakers on first arch 7 to 10 on upper limb, 15 to 17 on lower limb, total 22 to 26. Pyloric caeca numerous. Colour: adults dark brown, with margin of spinous dorsal fin black; juveniles dark brown with conspicuous white spots in vertical series usually extending onto head and dorsal fin, the caudal and pectoral fins pale yellow, and a black saddle-blotch on caudal peduncle reaching below lateral line. Size: Maximum total length about 120 cm; maximum weight 30 kg. Habitat, biology, and fisheries: Adults occur on rocky bottoms at depths of 30 to 400 m, but are most common between 100 and 200 m.Juveniles occur inshore and often reported from northeast coast of USA.Off the southeast coast of the USA, most females are mature by age 4 or 5 (40 to 50 cm); as expected for a protogynous species, males less than 6 years old (55 to 60 cm) were not found, and 40% of all fish 8 years or older (70 cm) were males. Spawning occurs off the Florida Keys from April to July.Age and growth parameters of this relatively unexploited population in the lower Florida Keys in 1983 were L t = 1320(1-e-0.087 (t-1.013)) where L t is total length at age t; the maximum age attained was at least 27 years. The weight/length relationship isW=2.45X10-8L2.93 where weight (W) is in kilograms and total length (L) is in millimetres. Adults feed mainly on fishes, gastropods, cephalopods, and brachyuran crustaceans. Commercially important stocks occur from North Carolina to Georgia, off Florida west coast, lower Florida Keys, Honduras/Nicaraguan shelf, Panama, and Colombia. Caught with hook-and-line, bottom longlines, and traps. Distribution: Mainly continental:Massachusetts to Gulf of Mexico, Bermuda, Caribbean (main- land coast, but rare in Venezuela) and southern Brazil; not known from West Indies, except Cuba and Bimini. Remarks: The eastern Pacific species Epinephelus niphobles is very similar to E. niveatus; juvenile niphobles (5 to 10 cm standard length) have smaller eyes, pelvic fins not longer than pectoral fins, and more pectoral-fin rays (modally 19). 1348 Bony Fishes

Epinephelus striatus (Bloch, 1792) GPN Frequent synonyms / misidentifications: None / None. FAO names: En - Nassau grouper; Fr - Mérou rayé; Sp - Cherna criolla.

Diagnostic characters: Body depth less than head length, 2.6 to 2.9 times in standard length (for fish 16 to 33 cm standard length).Preopercle rounded and evenly serrate.Gill rakers on first arch 8 or 9 on upper limb, 15 to 17 on lower limb, total 23 to 26.Dorsal fin with 11 spines and 16 to 18 soft rays, third or fourth spine longest, interspinous membranes distinctly incised; anal fin with 3 spines and 8 soft rays; caudal fin rounded in juveniles, convex (corners angular) in adults; pectoral-fin rays 17 to 19. Scales ctenoid, about 50 lateral-line scales and 98 to 106 lateral-scale series. Colour: body generally buff, with 5 dark brown vertical bars; large black saddle-blotch on caudal peduncle; a row of black dots below and behind eye. Dis- tinctive dark tuning fork beginning at front of upper jaw, extending dorsally along interorbital region, and bifurcating on top of head; another dark band from tip of snout through eye and then curving upward to meet its fellow just before dorsal-fin origin. Some fish have irregular pale spots and blotches all over head and body; fish from deep water usually pinkish or reddish ventrally. As in many groupers, the colour pattern can change in less than a minute from almost white to uniformly dark brown depending on the mood of the fish. Size: Maximum total length about 100 cm and 25 kg. Habitat, biology, and fisheries: Coral reefs from 5 to 100 m; juveniles occur on seagrass beds. Feeds mainly on fishes (54%) and crabs (23%), with lesser amounts of other crustaceans and molluscs.Spawning aggregations of 50 to perhaps 100 000 fish have been reported from the Bahamas, Belize, Jamaica, Cayman, and Vir- gin Islands. These aggregations occur in depths of 20 to 40 m at the outer reef shelf edge in January and February at or near the time of the new moon. During spawning, most fish (males and females) display the bicoloured (dark dorsally, pale ventrally ‘non-aggressive’) pattern and circle above the bottom. Some females remained in the barred pattern, becoming very dark as mating approached and were closely followed by bicoloured fish during courtship.Spawning occurs at sunset, in groups of 3 to about 25 fish and is preceded by vari- ous movements of the courting group: vertical spirals, vertical runs followed by rapidly crowding together then rapidly dispersing, and horizontal runs near the bottom. Mating is initiated by a dark phase fish (presumed female) dashing forward and upward, followed closely by bicoloured males releasing a white cloud of sperm, and other bicoloured fish (presumed females) apparently shedding eggs. The larvae metamorphose to pelagic juveniles after 37 to 45 days. In Bermuda, spawning lasts from early May to August. Based on studies of settlement of pelagic juveniles, spawning in Exuma Sound (Bahamas) occurs near the full moon of December. An important foodfish throughout the Caribbean. Caught with hook-and-line and in traps. Distribution: Bermuda, Florida (Tortugas and off Key West), northwestern Gulf of Mexico, Campeche Bank throughout Caribbean and southwards to Bahia, Brazil.

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