LUTJANIDAESnappers

by W.D. Anderson, Jr., Grice Marine Biological Laboratory, Charleston, South Carolina, USA

Diagnostic characters: Small to medium-sized (to about 160 cm) perch-like fishes, oblong in shape, mod-erately 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 molariformteeth. 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 portiondeeply incised posteriorly where it joins soft portion. Caudal fin truncate, or nearly so, to deeply forked. Dorsalfin 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 or9 (rarely 7) soft rays. Caudal fin with 17 principal rays (9 in upper lobe + 8 in lower lobe). Pelvic fin thoracic, in-serted beneath pectoral fin, with 1 spine and 5 soft rays.Scales moderate in size, ctenoid.Pelvic axillary scalesusually well developed. Lateral line complete. Vertebrae 24 (10 precaudal and 14 caudal). Colour: highly vari-able;many species mainly red or reddish, others with violet, brown, or grey prominent;often with spots or lines.

Habitat, biology, and fisheries: Occur worldwide in warm seas; juveniles of some species enter estuariesand the lower reaches of rivers; a few western Pacific species of Lutjanus are inhabitants of fresh waters; onoccasion some species are found in hypersaline lagoons. Mostly bottom-associated fishes, occurring fromshallow inshore areas to depths of about 550 m, mainly over reefs or rocky outcrops. Active, mostly nocturnalpredators 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 spe-cies 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 ofeggs. Populations in continental waters have extended spawning throughout the summer, whereas those oc-curring 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 apparentlyat 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 verticaldistribution 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 speciesthat reach large sizes are important recreational fishes in some areas.Some species have been reported to beoccasionally ciguatoxic in certain areas. They are caught with bottom longlines, handlines, traps, a variety ofnets, and trawls. The total commercial catch of Lutjanidae reported from the Western Central Atlantic from1995 to 1999 ranged from 10 588 to 16 413 t.

Perciformes: Percoidei: Lutjanidae 1479

snout and lacrymalscaleless

vomer andpalatineswith teeth

maxillamostly under

lachrymal

cheek andoperculum

scaly

preopercleserrate

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Similar families occurring in the areaHaemulidae: scales present on snout and lachrymal, those on lachrymal often embedded; chin with 2 en-larged 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.

Key to the genera and species of Lutjanidae occurring in the areaNotes: Counts of gill rakers are of those on the first arch, including rudiments, except where noted. Counts oflateral-line scales are of tubed scales. Counts of rows of lateral scales are of the number of anteriorly inclinedoblique rows above the lateral line between the upper edge of the opercle and the middle of the caudal-finbase. 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 direc-tion from origin of anal fin to, but not including, a lateral-line scale.

2a. Maxilla with scales; spinous portion of dorsal fin deeply notched at its junction with soft por-tion (Fig 3) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Etelis oculatus

2b. Maxilla without scales; spinous portion of dorsal fin not deeply notched at its junction withsoft portion (Fig. 4). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . � 3

1480 Bony Fishes

1a. Dorsal and anal fins with-out scales (Fig. 1); dorsalfin with 10 spines and usu-ally 10 or 11, rarely 9, softrays . . . . . . . . . . . . . � 2

1b. Soft dorsal and anal finswith scales (Fig. 2); dorsalfin with 10 or 12, rarely, 9,11, or 13 spines, and 11 to14, rarely 10 or 15, soft rays . . . � 4

scales on softdorsal andanal fins

Fig. 1 dorsal and anal fins Fig. 2 dorsal and anal fins

no scales

Haemulidae Sparidae

Fig. 3 Etelis Fig. 4 Pristipomoides and Apsilus

deeply notched not deeply notched

3a. Interorbital region flattened, not convex (Fig.5); last soft ray of both dorsal and anal fins lon-ger 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 alittle shorter than next to last soft ray (Fig. 8); dorsal fin with 10 (occasionally 9) soft rays. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Apsilus dentatus

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); yellowstripe from tip of snout (passing under eye) to caudal peduncle, widening to cover much ofcaudal peduncle and caudal fin, colour fading in preservative . . . . . . . . . . Ocyurus chrysurus

Perciformes: Percoidei: Lutjanidae 1481

Fig. 10 Rhomboplites aurorubens Fig. 11 Ocyurus chrysurus

Fig. 9 teeth on roof of mouth

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

interorbitalregion flat

interorbitalregionconvex

last soft raylonger

next to lastsoft ray longer

4a. Ectopterygoid teeth absent; gill rakers, ex-cluding rudiments, 16 or fewer, rarely 17, onlower limb of first gill arch; caudal fin trun-cate, or nearly so, to moderately forked,lobes of fin not elongated . . . . . . . . . . Lutjanus

4b. Ectopterygoid teeth present (Fig. 9); gill rak-ers, excluding rudiments, 17 to 22 on lowerlimb of first gill arch; caudal-fin forked, lobesof fin moderately to well elongated . . . . . . . . � 5

ectopterygoidteeth

vomerine teeth palatine teeth

Key to the species of Lutjanus occurring in the area1a. Dorsal fin with 10 spines and usually 12 (rarely 11 or 13) soft rays; a dark spot below ante-

rior part of soft dorsal fin, usually persisting throughout life (occasionally absent inLutjanus synagris) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . � 2

1b. Dorsal fin usually with 10 spines and 14 soft rays, rarely 9 or 11 spines and 13 or 15 softrays; 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 ofpreopercle with prominent, well-serrated posterior projection; gill rakers on first arch 7 or 8on 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 largerthan about 6 cm standard length (Fig. 13); angle of preopercle without prominent posteriorprojection; gill rakers on first arch 6 or 7 on upper limb and 13 or 14 on lower limb, rarely 12or 15 on lower limb . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lutjanus synagris

1482 Bony Fishes

Fig. 13 Lutjanus synagrisFig. 12 Lutjanus mahogoni

1/4-1/2 of spotextending below

lateral line

1/4 or less of spotbelow lateral line

3a. A large, pronounced black spot atbase and in axil of pectoral fin; nodark spot below anterior part ofsoft dorsal fin; anal fin rounded; adark area on scales at base of softdorsal fin (not always obvious onpreserved specimens); iris of eyegolden yellow to orange in life (Fig.14) . . . . . . . . . . Lutjanus buccanella

3b. No large, pronounced black spot atbase and in axil of pectoral fin; darkspot below anterior part of soft dor-sal fin present or absent; anal finrounded or angulated . . . . . . . . . . � 4

Fig. 14 Lutjanus buccanella

4a. Anal fin rounded at all sizes, the middle rays less than half length of head (Fig. 15); no darkspot below anterior part of soft dorsal fin . . . . . . . . . . . . . . . . . . . . . . . . . � 5

4b. Anal fin angulated in larger individuals, the middle rays elongated, the longest almost halfto greater than half length of head (anal fin rounded in L. analis less than about 4 cm stan-dard 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 ante-rior part of soft dorsal fin, at least in young (this spot present in L. analis to at least 46 cmstandard length, but disappearing by about 20 to 30 cm standard length in L.campechanus, L. purpureus, and L. vivanus . . . . . . . . . . . . . . . . . . . . . . . . � 8

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 to3.5 times in standard length (in L. apodus of 7 to 10 cm standard length pectoral-fin lengthapproximately equal to that of L. griseus of similar size); body comparatively deep, greatestdepth 2.3 to 2.8, usually 2.4 to 2.7, times in standard length . . . . . . . . . . . . . . . . . � 7

Perciformes: Percoidei: Lutjanidae 1483

a) Lutjanus cyanopterus

Fig. 17

5a. Vomerine tooth patch without a distinct pos-terior extension on median line (Fig. 17a);upper and lower canines very strong, aboutequally developed; cheek scales in 8 to 10,usually 9, rows (Fig. 18) . . . . . Lutjanus cyanopterus

5b. Vomerine tooth patch anchor-shaped, witha median posterior extension (Fig. 17b); up-per canines much larger than lower; cheekscales in 6 to 9 (usually 7 or 8) rows . . . . . . . � 6

Fig. 15 anal fin Fig. 16 anal fin

anal finrounded

anal finangular

Fig. 18 Lutjanus cyanopterus Fig. 19 Lutjanus griseus

b)

upper and lowercanines aboutequally strong

7a. Scales relatively large, transverse rows between upper edge of opercle and caudal-finbase 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-finbase 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

1484 Bony Fishes

Fig. 22Lutjanus analis

Fig. 23

8a. Vomerine tooth patch without a distinct pos-terior extension on median line (Fig. 22); softrays in anal fin usually 8, rarely 7; spot belowanterior part of soft dorsal fin relatively largein small individuals, small but distinct in largeones; 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 8or 9; spot below anterior part of soft dorsalfin present in young, diffuse or absent inadults. . . . . . . . . . . . . . . . . . . . . � 9

Fig. 20 Lutjanus apodus Fig. 21 Lutjanus jocu

no whitish barbelow eye

whitish barbelow eye

9a. Soft rays in anal fin 9, rarely 8; rows oflateral scales 46 to 50, usually 47 to49; scales above lateral line 7 to 10,usually 8 or 9; scales below lateral line15 to 19, usually 16 or 17; sum of rowsof lateral scales and scales above andbelow lateral line 69 to 75; iris of eyered in life (Fig. 24) . . . . Lutjanus campechanus

9b. Soft rays in anal fin 8, rarely 7 or 9;rows of lateral scales 49 to 53, usually50 or 51; scales above lateral-line 9 to12, usually 10 to 12; scales below lat-eral line 16 to 24, usually 17 to 23;sum of rows of lateral scales andscales above and below lateral line 76to 88 . . . . . . . . . . . . . . . . . . � 10

Fig. 24 Lutjanus campechanus

10a. Scales below lateral line 16 to 19; scales above lateral line 9 to 11, usually 10; sum of rowsof 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; sumof rows of lateral scales and scales above and below lateral line 81 to 88; iris of eye brightyellow in life (Fig. 26) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lutjanus vivanus

Key to the species of Pristipomoides occurring in the area

2a. Lateral-line scales 48 to 52; total gill rakers on first arch 24 to 28 (Fig. 28) . Pristipomoides aquilonaris2b. Lateral-line scales 54 to 57; total gill rakers on first arch 19 to 25 (Fig. 29)

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pristipomoides macrophthalmus

Perciformes: Percoidei: Lutjanidae 1485

Fig. 25 Lutjanus purpureus Fig. 26 Lutjanus vivanus

Fig. 27 Pristipomoides freemani

1a. Depth of body at origin of dorsal fin 3.5to 4.2 times in standard length (24 to28% standard length); total gill rakerson first arch 28 to 32; lateral-line scales49 to 51 (Fig. 27) . . . . . Pristipomoides freemani

1b. Depth of body at origin of dorsal fin 2.5to 3.2 times in standard length (31 to41% standard length); total gill rakerson first arch 19 to 28; lateral-line scales48 to 57 . . . . . . . . . . . . . . . . . . � 2

Fig. 28 Pristipomoides aquilonaris Fig. 29 Pristipomoides macrophthalmus

List of species occurring in the areaThe symbol� is given when species accounts are included.� Apsilus dentatus Guichenot, 1853.

� Etelis oculatus (Valenciennes, 1828).

� Lutjanus analis (Cuvier, 1828).� Lutjanus apodus (Walbaum, 1792).� Lutjanus buccanella (Cuvier, 1828).� Lutjanus campechanus (Poey, 1860).� Lutjanus cyanopterus (Cuvier, 1828).� Lutjanus griseus (Linnaeus, 1758).� Lutjanus jocu (Bloch and Schneider, 1801).� Lutjanus mahogoni (Cuvier, 1828).� Lutjanus purpureus (Poey, 1866).� Lutjanus synagris (Linnaeus, 1758).� Lutjanus vivanus (Cuvier, 1828).

� Ocyurus chrysurus (Bloch, 1791).

� Pristipomoides aquilonaris (Goode and Bean, 1896).� Pristipomoides freemani Anderson, 1966.� Pristipomoides macrophthalmus (Müller and Troschel, 1848).

� Rhomboplites aurorubens (Cuvier, 1829).

ReferencesAllen, 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.

1486 Bony Fishes

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 andouter series of conical teeth; canine or canine-like teeth present anteriorly in both jaws; teeth on roof of mouthin a triangular or chevron-shaped patch on vomer and in elongate band on each palatine; no teeth onectopterygoids. Maxilla without scales. Interorbital region convex. Gill rakers on first arch 7 or 8 on upperlimb and 15 or 16 on lower limb, total 22 to 24.Dorsal fin single, spinous portion of fin not deeply incised atits junction with soft portion. Last soft ray of both dorsal and anal fins a little shorter than next to lastsoft ray. Caudal fin forked to emarginate. Dorsal fin with 10 spines and 10, occasionally 9, soft rays. Analfin with 3 spines and 8 soft rays. Pectoral fin with 15 or 16 rays. Membranes of dorsal and anal fins withoutscales. 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, surroundingbronze 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 spawnsduring most of the year. Juveniles have beenovserved to mimic blue chromis (Chromiscyanea; family Pomacentridae) in waters off theCayman 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, southeastof Galveston, Texas), and from the Caribbean offBelize and Venezuela; probably more wide-spread.

Perciformes: Percoidei: Lutjanidae 1487

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