Lutjanidae 1479

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LUTJANIDAE Snappers by W.D. Anderson, Jr., Grice Marine Biological Laboratory, Charleston, South Carolina, USA iagnostic characters: Small to medium-sized (to about 160 cm) perch-like fishes, oblong in shape, mod- Derately compressed laterally.Two nostrils on each side of snout. No enlarged pores on chin. Mouth ter- minal and fairly large. Maxilla slipping for most or all of its length under lachrymal when mouth closed. Supramaxilla absent. Jaws with distinct canines or canine-like teeth; no incisiform or molariform teeth. Vomer and palatines with teeth. Ectopterygoid teeth present only in Ocyurus and Rhomboplites. Cheek and operculum scaly; maxilla with or without scales; snout, lachrymal, and lower jaw naked. Preopercle typically serrate, often finely. Premaxillae moderately protrusible. Opercular spines 2. Branchiostegal rays 7. Gill membranes separate, free from isthmus. Dorsal fin single; in Etelis spinous portion deeply incised posteriorly where it joins soft portion. Caudal fin truncate, or nearly so, to deeply forked. Dorsal fin with 10 or 12 (rarely 9, 11, or 13) spines and 10 to 14 (rarely 9 or 15) soft rays. Anal fin with 3 spines and 8 or 9 (rarely 7) soft rays. Caudal fin with 17 principal rays (9 in upper lobe+8inlowerlobe). Pelvic fin thoracic, in- serted beneath pectoral fin, with 1 spine and 5 soft rays.Scales moderate in size, ctenoid.Pelvic axillary scales usually well developed. Lateral line complete. Vertebrae 24 (10 precaudal and 14 caudal). Colour: highly variable;many species mainly red or reddish, others with violet, brown, or grey prominent;often with spots or lines.

snout and lacrymal scaleless

vomer and palatines with teeth

maxilla mostly under lachrymal

cheek and preopercle operculum serrate scaly

Habitat, biology, and fisheries: Occur worldwide in warm seas; juveniles of some species enter estuaries and the lower reaches of rivers; a few western Pacific species of Lutjanus are inhabitants of fresh waters; on occasion some species are found in hypersaline lagoons. Mostly bottom-associated fishes, occurring from shallow inshore areas to depths of about 550 m, mainly over reefs or rocky outcrops. Active, mostly nocturnal predators feeding on fishes, crustaceans (especially crabs, shrimps, stomatopods, lobsters), molluscs (gas- tropods, cephalopods), and pelagic urochordates; plankton is particularly important in the diets of those species with reduced dentition and numerous well-developed gill rakers. Gonochoristic (sexes separate), reaching sexual maturity at about 40 to 50% of maximum length, with big females producing large numbers of eggs. Populations in continental waters have extended spawning throughout the summer, whereas those occurring around islands spawn throughout the year with peaks in spring and autumn;lutjanids are batch spawn- ers, with individual females usually spawning several times in a reproductive season. Spawning is apparently at night, on some occasions coinciding with spring tides.In those species in which it has been observed, court- ship terminates in a spiral swim upward, with gametes released just below the surface.Eggs and larvae identi- fied as lutjanid are pelagic; the larvae avoid surface waters during the day, but display a more even vertical distribution at night. Long-lived, slow-growing fishes with relatively low rates of natural mortality and with con- siderable vulnerability to overfishing.Snappers are important to artisanal fisheries, but seldom the prime inter- est of major commercial fishing activities; many are fine foodfishes, frequently found in markets. The species that reach large sizes are important recreational fishes in some areas.Some species have been reported to be occasionally ciguatoxic in certain areas. They are caught with bottom longlines, handlines, traps, a variety of nets, and trawls. The total commercial catch of Lutjanidae reported from the Western Central Atlantic from 1995 to 1999 ranged from 10 588 to 16 413 t. 1480 Bony Fishes

Similar families occurring in the area Haemulidae: scales present on snout and lachrymal, those on lachrymal often embedded; chin with 2 enlarged pores anteriorly; no teeth on vomer or palatines; vertebrae 26 or 27. Sparidae: teeth in jaws variable, conical, incisiform, or molariform; vomer and palatines usually without teeth; preopercular margin smooth; branchiostegal rays 6.

Haemulidae Sparidae

Key to the genera and species of Lutjanidae occurring in the area Notes: Counts of gill rakers are of those on the first arch, including rudiments, except where noted. Counts of lateral-line scales are of tubed scales. Counts of rows of lateral scales are of the number of anteriorly inclined oblique rows above the lateral line between the upper edge of the opercle and the middle of the caudal-fin base. Counts of scales above the lateral line are made in a posteroventral direction from origin of dorsal fin to, but not including, a lateral-line scale.Counts of scales below the lateral line are made in an anterodorsal direction from origin of anal fin to, but not including, a lateral-line scale. 1a. Dorsal and anal fins without scales (Fig. 1); dorsal fin with 10 spines and usually 10 or 11, rarely 9, soft ® rays ...... 2 no scales scales on soft 1b. Soft dorsal and anal fins dorsal and with scales (Fig. 2); dorsal anal fins fin with 10 or 12, rarely, 9, 11, or 13 spines, and 11 to 14, rarely 10 or 15, soft rays . . . ® 4 Fig. 1 dorsal and anal fins Fig. 2 dorsal and anal fins

2a. Maxilla with scales; spinous portion of dorsal fin deeply notched at its junction with soft portion (Fig 3) ...... Etelis oculatus 2b. Maxilla without scales; spinous portion of dorsal fin not deeply notched at its junction with soft portion (Fig. 4)...... ® 3

deeply notched not deeply notched

Fig. 3 Etelis Fig. 4 Pristipomoides and Apsilus Perciformes: Percoidei: Lutjanidae 1481

3a. Interorbital region flattened, not convex (Fig. 5); last soft ray of both dorsal and anal fins longer than next to last soft ray (Fig. 7); dorsal fin with 11 (rarely 10) soft rays ...... Pristipomoides 3b. Interorbital region convex, not flattened (Fig. 6); last soft ray of both dorsal and anal fins a little shorter than next to last soft ray (Fig. 8); dorsal fin with 10 (occasionally 9) soft rays ...... Apsilus dentatus

interorbital interorbital last soft ray next to last region flat region longer soft ray longer convex

Fig. 5 Pristipomoides Fig. 6 Apsilus Fig. 7 Pristipomoides Fig. 8 Apsilus

vomerine teeth palatine teeth

4a. Ectopterygoid teeth absent; gill rakers, ex- ectopterygoid cluding rudiments, 16 or fewer, rarely 17, on teeth lower limb of first gill arch; caudal fin truncate, or nearly so, to moderately forked, lobes of fin not elongated...... Lutjanus 4b. Ectopterygoid teeth present (Fig. 9); gill rakers, excluding rudiments, 17 to 22 on lower limb of first gill arch; caudal-fin forked, lobes of fin moderately to well elongated ...... ® 5 Fig. 9 teeth on roof of mouth

5a. Dorsal fin with 12 (very rarely 13) spines and 11 (rarely 10 or 12) soft rays (Fig. 10); vermil- ion in life, no yellow stripe along side of body, colour fading in preservative . Rhomboplites aurorubens 5b. Dorsal fin with 10, rarely 9 or 11, spines and 12 or 13, rarely 14, soft rays (Fig. 11); yellow stripe from tip of snout (passing under eye) to caudal peduncle, widening to cover much of caudal peduncle and caudal fin, colour fading in preservative ...... Ocyurus chrysurus

Fig. 10 Rhomboplites aurorubens Fig. 11 Ocyurus chrysurus 1482 Bony Fishes

Key to the species of Lutjanus occurring in the area 1a. Dorsal fin with 10 spines and usually 12 (rarely 11 or 13) soft rays; a dark spot below anterior part of soft dorsal fin, usually persisting throughout life (occasionally absent in Lutjanus synagris) ...... ® 2 1b. Dorsal fin usually with 10 spines and 14 soft rays, rarely 9 or 11 spines and 13 or 15 soft rays; dark spot below anterior part of soft dorsal fin present or absent ...... ® 3

2a. About 1/4 to 1/2 of dark lateral spot extending below lateral line (Fig. 12); angle of preopercle with prominent, well-serrated posterior projection; gill rakers on first arch 7 or 8 on upper limb and 15 to 17 on lower limb ...... Lutjanus mahogoni 2b. Less than 1/4 to none of dark lateral spot extending below lateral line in specimens larger than about 6 cm standard length (Fig. 13); angle of preopercle without prominent posterior projection; gill rakers on first arch 6 or 7 on upper limb and 13 or 14 on lower limb, rarely 12 or 15 on lower limb ...... Lutjanus synagris

1/4-1/2 of spot 1/4orlessofspot extending below below lateral line lateral line

Fig. 12 Lutjanus mahogoni Fig. 13 Lutjanus synagris

3a. A large, pronounced black spot at base and in axil of pectoral fin; no dark spot below anterior part of soft dorsal fin; anal fin rounded; a dark area on scales at base of soft dorsal fin (not always obvious on preserved specimens); iris of eye golden yellow to orange in life (Fig. 14) ...... Lutjanus buccanella 3b. No large, pronounced black spot at base and in axil of pectoral fin; dark spot below anterior part of soft dor- Fig. 14 Lutjanus buccanella sal fin present or absent; anal fin rounded or angulated ...... ® 4 Perciformes: Percoidei: Lutjanidae 1483

4a. Anal fin rounded at all sizes, the middle rays less than half length of head (Fig. 15); no dark spot below anterior part of soft dorsal fin ...... ® 5 4b. Anal fin angulated in larger individuals, the middle rays elongated, the longest almost half to greater than half length of head (anal fin rounded in L. analis less than about 4 cm standard length, in L. campechanus and L. purpureus less than about 5 cm standard length, and in L. vivanus less than about 6 cm standard length) (Fig. 16); a dark spot below anterior part of soft dorsal fin, at least in young (this spot present in L. analis to at least 46 cm standard length, but disappearing by about 20 to 30 cm standard length in L. campechanus, L. purpureus, and L. vivanus...... ® 8

anal fin angular

anal fin rounded Fig. 15 anal fin Fig. 16 anal fin

5a. Vomerine tooth patch without a distinct posterior extension on median line (Fig. 17a); upper and lower canines very strong, about equally developed; cheek scales in 8 to 10, usually 9, rows (Fig. 18) . . . . . Lutjanus cyanopterus 5b. Vomerine tooth patch anchor-shaped, with a median posterior extension (Fig. 17b); up- a) Lutjanus cyanopterus b) per canines much larger than lower; cheek scales in 6 to 9 (usually 7 or 8) rows ...... ® 6 Fig. 17

upper and lower canines about equally strong Fig. 18 Lutjanus cyanopterus Fig. 19 Lutjanus griseus

6a. Pectoral-fin length about equal to distance from tip of snout to posterior edge of preopercle, 3.7 to 4.2 times in standard length; body comparatively slender, greatest depth 2.6 to 3.2, usually 2.7 to 3.1, times in standard length (Fig. 19) ...... Lutjanus griseus 6b. Pectoral-fin longer than distance from tip of snout to posterior edge of preopercle, 3.0 to 3.5 times in standard length (in L. apodus of 7 to 10 cm standard length pectoral-fin length approximately equal to that of L. griseus of similar size); body comparatively deep, greatest depth 2.3 to 2.8, usually 2.4 to 2.7, times in standard length ...... ® 7 1484 Bony Fishes

7a. Scales relatively large, transverse rows between upper edge of opercle and caudal-fin base 39 to 44, usually 40 to 43; lateral-line scales 40 to 45; scales above lateral line 5 to 7; no whitish bar below eye (Fig. 20)...... Lutjanus apodus 7b. Scales of moderate size, transverse rows between upper edge of opercle and caudal-fin base 45 to 49, usually 46 to 48; lateral-line scales 46 to 49; scales above lateral line 8 to 11; a rather diffuse whitish bar below eye, not obvious in all preserved specimens (Fig. 21) . Lutjanus jocu

no whitish bar whitish bar below eye below eye

Fig. 20 Lutjanus apodus Fig. 21 Lutjanus jocu

8a. Vomerine tooth patch without a distinct posterior extension on median line (Fig. 22); soft rays in anal fin usually 8, rarely 7; spot below anterior part of soft dorsal fin relatively large in small individuals, small but distinct in large ones; iris of eye red in life ...... Lutjanus analis 8b. Vomerine tooth patch triangular or anchor- shaped, with a median posterior extension (Fig. 23); soft rays in anal fin 7 to 9, usually 8 or 9; spot below anterior part of soft dorsal fin present in young, diffuse or absent in Fig. 22 Fig. 23 adults...... ® 9 Lutjanus analis

9a. Soft rays in anal fin 9, rarely 8; rows of lateral scales 46 to 50, usually 47 to 49; scales above lateral line 7 to 10, usually 8 or 9; scales below lateral line 15 to 19, usually 16 or 17; sum of rows of lateral scales and scales above and below lateral line 69 to 75; iris of eye red in life (Fig. 24) . . . . Lutjanus campechanus 9b. Soft rays in anal fin 8, rarely 7 or 9; rows of lateral scales 49 to 53, usually 50 or 51; scales above lateral-line 9 to 12, usually 10 to 12; scales below lateral line 16 to 24, usually 17 to 23; Fig. 24 Lutjanus campechanus sum of rows of lateral scales and scales above and below lateral line 76 to 88 ...... ® 10 Perciformes: Percoidei: Lutjanidae 1485

10a. Scales below lateral line 16 to 19; scales above lateral line 9 to 11, usually 10; sum of rows of lateral scales and scales above and below lateral line 76 to 82; iris of eye red in life (Fig. 25) ...... Lutjanus purpureus 10b. Scales below lateral line 20 to 24; scales above lateral line 10 to 12, usually 11 or 12; sum of rows of lateral scales and scales above and below lateral line 81 to 88; iris of eye bright yellow in life (Fig. 26) ...... Lutjanus vivanus

Fig. 25 Lutjanus purpureus Fig. 26 Lutjanus vivanus

Key to the species of Pristipomoides occurring in the area

1a. Depth of body at origin of dorsal fin 3.5 to 4.2 times in standard length (24 to 28% standard length); total gill rakers on first arch 28 to 32; lateral-line scales 49 to 51 (Fig. 27) . . . . . Pristipomoides freemani 1b. Depth of body at origin of dorsal fin 2.5 to 3.2 times in standard length (31 to 41% standard length); total gill rakers on first arch 19 to 28; lateral-line scales 48 to 57 ...... ® 2 Fig. 27 Pristipomoides freemani

2a. Lateral-line scales 48 to 52; total gill rakers on first arch 24 to 28 (Fig. 28) . Pristipomoides aquilonaris 2b. Lateral-line scales 54 to 57; total gill rakers on first arch 19 to 25 (Fig. 29) ...... Pristipomoides macrophthalmus

Fig. 28 Pristipomoides aquilonaris Fig. 29 Pristipomoides macrophthalmus 1486 Bony Fishes

List of species occurring in the area The symbol 0 is given when species accounts are included. 0 Apsilus dentatus Guichenot, 1853. 0 Etelis oculatus (Valenciennes, 1828). 0 Lutjanus analis (Cuvier, 1828). 0 Lutjanus apodus (Walbaum, 1792). 0 Lutjanus buccanella (Cuvier, 1828). 0 Lutjanus campechanus (Poey, 1860). 0 Lutjanus cyanopterus (Cuvier, 1828). 0 Lutjanus griseus (Linnaeus, 1758). 0 Lutjanus jocu (Bloch and Schneider, 1801). 0 Lutjanus mahogoni (Cuvier, 1828). 0 Lutjanus purpureus (Poey, 1866). 0 Lutjanus synagris (Linnaeus, 1758). 0 Lutjanus vivanus (Cuvier, 1828). 0 Ocyurus chrysurus (Bloch, 1791). 0 Pristipomoides aquilonaris (Goode and Bean, 1896). 0 Pristipomoides freemani Anderson, 1966. 0 Pristipomoides macrophthalmus (Müller and Troschel, 1848). 0 Rhomboplites aurorubens (Cuvier, 1829).

References Allen, G.R. 1985. FAO species catalogue. Vol. 6. Snappers of the world. An annotated and illustrated catalogue of lutjanid species known to date. FAO Fish. Synop., (125)Vol.6:208 p. Anderson, W.D., Jr. 1966. A new species of Pristipomoides (Pisces: Lutjanidae) from the tropical western Atlantic. Bull. Mar. Sci., 16:814-826. Anderson, W.D., Jr. 1967. Field guide to the snappers (Lutjanidae) of the western Atlantic. U. S. Dept. Inter., Fish and Wildl. Serv., Bur. Comm. Fish., Circular 252:1-14. Anderson, W.D., Jr. 1987. Systematics of the fishes of the family Lutjanidae (Perciformes: Percoidei), the snappers. In Tropical snappers and groupers: Biology and fisheries management, edited by J. J. Polovina and S. Ralston. Boulder, Colorado, Westview Press, pp 1-31. Rivas, L.R. 1966. Review of the Lutjanus campechanus complex of red snappers. Quart. Journ. Florida Acad. Sci., 29:117-136. Perciformes: Percoidei: Lutjanidae 1487

Apsilus dentatus Guichenot, 1853 ASX Frequent synonyms / misidentifications: None / None. FAO names: En - Black snapper; Fr - Vivaneau noir; Sp - Pargo mulato.

Diagnostic characters: Upper and lower jaws each with inner band of villiform to small conical teeth and outer series of conical teeth; canine or canine-like teeth present anteriorly in both jaws; teeth on roof of mouth in a triangular or chevron-shaped patch on vomer and in elongate band on each palatine; no teeth on ectopterygoids. Maxilla without scales. Interorbital region convex. Gill rakers on first arch 7 or 8 on upper limb and 15 or 16 on lower limb, total 22 to 24.Dorsal fin single, spinous portion of fin not deeply incised at its junction with soft portion. Last soft ray of both dorsal and anal fins a little shorter than next to last soft ray. Caudal fin forked to emarginate. Dorsal fin with 10 spines and 10, occasionally 9, soft rays. Anal fin with 3 spines and 8 soft rays. Pectoral fin with 15 or 16 rays. Membranes of dorsal and anal fins without scales. Tubed scales in lateral line 58 to 63. Colour: body violet to brownish black, more intense on head; lower sides and belly paler; small juveniles bright blue; iris of eye almost black peripherally, surrounding bronze central area; fins mostly brown to black, caudal fin with pale distal margin, some individuals with con- siderable blue on fins. Size: Maximum standard length to at least 55 cm, commonly to 40 cm standard length. Habitat, biology, and fisheries: Mainly found over rocky bottoms in depths between 12 and 240 m; very com- mon in the Bahamas along steep drop offs. The young sometimes found near the surface. Feeds on fishes, cephalopods, and tunicates. Apparently spawns during most of the year. Juveniles have been ovserved to mimic blue chromis (Chromis cyanea; family Pomacentridae) in waters off the Cayman Islands. Caught mostly with handlines. Marketed mainly fresh, sometimes frozen. Distribution: Known from the West Indies, Florida Keys, the northwestern Gulf of Mexico (near the West Flower Garden Bank, southeast of Galveston, Texas), and from the Caribbean off Belize and Venezuela; probably more wide- spread.

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