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3890 Bony

CYNOGLOSSIDAE Tonguesoles by T.A. Munroe iagnostic characters: Lance- or tongue-shaped with eyes on left side of body (size to D40 cm); body highly compressed and tapering to a point posteriorly. Eyes small and usually close together. Mouth small, subterminal, asymmetrical; reaching posteriorly to point between verticals at anterior and posterior margins of lower eye or slightly posterior to lower eye; jaws moderately curved on eyed side and notably on blind side; teeth minute and usually only on blind-side jaws; lips fringed with labial papillae in Paraplagusia. Rostral hook usually present below mouth. Posterior margin of preopercle strongly attached to opercle, without free margin and covered with skin and scales. No spiny rays in dorsal, anal, and pelvic fins; reaching far forward onto head; dorsal and anal fins confluent with caudal fin; pectoral fins absent; usually only right pelvic fin present. Scales small, ctenoid or cycloid (smooth). Lateral lines variable; none present in Symphurus; 1 to 3 on eyed side and 0 to 2 on blind side in and Paraplagusia. of Symphurus with only a single proximal dorsal-fin pterygiophore inserted into first interneural space; species of Cynoglossus and Paraplagusia with several pterygiophores inserted into first interneural space. Colour: colour pattern may vary within a species; eyed side usually uniformly brownish or greyish, often variably marked with patches, spots, or cross bands on body, and some species with blotches and spots on fins; most species uniformly whitish or yellowish on blind side, others with small indistinct cloudy patches or darker spots.

eyes on left scales between middle and side of head upper lateral lines lateral lines no fin spines origin of dorsal fin anterior to eyes

dorsal, anal, and caudal fins confluent rostral hook usually present below mouth pectoral fins preopercular margin always absent embedded in skin

Habitat, biology, and fisheries: Small to medium-sized benthic fishes, commonly found on muddy bottoms and other substrates, from tidepools to deep waters on outer continental shelf and upper continental slope (to about 1 500 m). Tonguesoles are of limited economic importance with catch statistics for individual species not usually available. From 1990 to 1995, the FAO Yearbook of Fishery Statistics reports a yearly catch of around 22 700 to 28 600 t of Cynoglossidae from the Western Central Pacific. Marketed fresh, frozen, and also dried-salted. dorsal and anal fins origin of dorsal fin attached or not attached anterior to eyes Similar families occurring in the area to caudal fin : also with small mouth, preopercular margin embedded in skin, no fin spines, dorsal-fin orgin far forward on head, and some species with dorsal and anal fins joined to caudal fin, but eyes on right side of body (on left side in Cynoglossidae). Also, always with 2 pelvic fins, and pectoral fins sometimes present.

no fin spines eyes on right pectoral fins sometimes absent side of head Soleidae Pleuronectiformes: Cynoglossidae 3891 anterior rays of dorsal dorsal-fin origin and anal fins spinous posterior to eyes Psettodidae: mouth large, with large teeth; dorsal and mouth anal fins separate from caudal fin; dorsal fin not large, with extending forward on head; anterior dorsal- and anal-fin large teeth rays spinous; margin of preopercle free and distinct, not embedded in skin. , , : also with dorsal-fin origin far forward on head and no spines in fins, but dorsal and anal fins separate from caudal fin, and margin of preopercle free and distinct, not eyes on left or embedded in skin. right side of head Psettodidae margin of preopercle free eyes on left side

eyes on right side dorsal and anal fins not attached margin of to caudal fin preopercle free Bothidae

eyes on left side

margin of Paralichtyidae Pleuronectidae margin of preopercle preopercle free distinct, not covered with skin and scales Identification note 1. Midlateral-line scales are counted from the vertical lateral-line branch on the head (i.e. branch between midlateral and dorsolateral line) to the base of the caudal fin. 2. Since the caudal fin is confluent with both the dorsal and anal fins, caudal-fin rays are best counted from the blind side of the using transmitted light. In order to observe the point where the caudal-fin rays articulate to the epural and hypural bones, it may be necessary to remove some scales and skin at the caudal-fin base.

Key to the species of Cynoglossidae occurring in the area 1a. No lateral line on eyed side of body; mouth terminal, snout not hooked over mouth (Fig. 1) ...... Symphurus (primarily small-sized deep-water species with no commercial importance) 1b. Two or 3 lateral lines on eyed side of body; mouth inferior, front part of snout produced into rostral hook and covering mouth (Fig. 2) ...... ® 2

rostral hook 2-3 lateral lines no lateral line Fig. 1 Symphurus Fig. 2 3892 Bony Fishes

2a. Lips on eyed side of head not fringed with labial papillae ...... (Cynoglossus) ® 3 2b. Lips on eyed side of head distinctly fringed with labial papillae (Fig. 3) ...... (Paraplagusia) ® 23

3a. Caudal fin with 8 or 9 rays (caudal-fin rays best fringed lips observed from blind side using transmitted light) .....® 4 3b. Caudal fin with more than 9 rays ...... ® 9

4a. Two lateral lines on eyed side of body ...... ® 5 4b. Three lateral lines on eyed side of body ...... ® 6 Fig. 3 Paraplagusia

5a. Scales ctenoid on both sides of body; scales between lateral lines more numerous (10 to 13); 2 rounded, conspicuously pigmented spots on posterior one-third of eyed side of body ...... Cynoglossus maculipinnis 5b. Scales ctenoid on eyed side of body, cycloid on blind side; scales between lateral lines fewer (8); no obviously pigmented spots on eyed side of body ...... Cynoglossus ogilbyi (not yet reported from the area) upper eye advanced upper eye not advanced 6a. Upper eye (migratory eye) much in advance of fixed eye (Fig. 4a) ...... Cynoglossus microlepis 6b. Upper eye (migratory eye) not in advance of fixed eye (Fig. 4b) ...... ® 7

7a. Posterior region of peritoneum darkly pigmented (usually show- ing through abdomen on both a) b) sides of body) . . . . Cynoglossus mccullochi Fig. 4 7b. Peritoneum unpigmented ...... ® 8

8a. Body comparatively slender (19 to 23% standard length); maxilla extending posteriorly beyond fixed eye; corner of mouth nearer to branchial opening than to tip of snout (Fig. 5) ...... Cynoglossus gracilis (not yet reported from the area) 8b. Body comparatively deep (23 to 28% standard length); maxilla extending only to point equal with verticals through middle and posterior margin of fixed eye; corner of mouth nearer to tip of snout than to branchial opening (Fig. 6) ...... (not yet reported from the area)

Fig. 5 Cynoglossus gracilis Fig. 6 Cynoglossus abbreviatus

9a. Caudal fin usually with 10 rays ...... ® 10 9b. Caudal fin usually with 12 rays ...... ® 22

10a. Only 2 lateral lines (dorsal and medial) on eyed side of body ...... ® 11 10b. Three lateral lines (dorsal, medial, and ventral) on eyed side of body ...... ® 17 Pleuronectiformes: Cynoglossidae 3893

11a. Eyes small, pedunculate (Fig. 7) ...... Cynoglossus monopus 11b. Eyes not pedunculate ...... ® 12

12a. Eyes contiguous, or nearly so (Fig. 8a) ...... Cynoglossus kopsi 12b. Eyes distinctly separate (Fig. 8b) ...... ® 13 eyes pedunculate separate eyes eyes contiguous

a) b) Fig. 7 Cynoglossus monopus Fig. 8

13a. Scales ctenoid on both sides of body ...... ® 14 13b. Scales ctenoid on eyed side of body, and cycloid on blind side ...... ® 16

14a. Rostral hook comparatively short, extending only to point between verticals through anterior nostril and just anterior to anterior margin of fixed eye ...... ® 15 14b. Rostral hook comparatively long, extending at least to vertical through middle of fixed eye ...... Cynoglossus lida

15a. Dorsolateral line usually undulating; no dark blotches or irregular cross bands on body; eyed-side lower jaw with crenulate fleshy ridge comprised of several folds, or jaw with a single broadly triangular-shaped fleshy ridge (Fig. 9) ...... 15b. Dorsolateral line slightly undulating; eyed side with dark blotches forming irregular cross bands; eyed-side lower jaw with low, broadly rounded fleshy ridge (Fig. 10) . dorsolateral line undulating irregular cross bands

Fig. 9 Cynoglossus cynoglossus Fig. 10 Cynoglossus puncticeps

16a. Midlateral-line scales numerous (90 to 101); 11 or 12 scales between lateral lines ...... 16b. Midlateral-line scales fewer (56 to 70); 7 to 9 scales between lateral lines . . . . .

17a. Midlateral-line scales less than 100 ...... ® 18 17b. Midlateral-line scales 100 or greater ...... ® 19

18a. Blind side with ctenoid scales, except head region with cycloid or weakly ctenoid scales; only 17 or 18 scales between dorsolateral and midlateral lines ...... 18b. Blind side of body and head region with ctenoid scales throughout; about 24 scales between dorsolateral and midlateral lines ...... Cynoglossus waandersi

19a. One nostril on eyed side of head ...... Cynoglossus macrophthalmus 19b. Two nostrils on eyed side of head ...... ® 20 3894 Bony Fishes

20a. Dorsal-fin rays fewer than 110 ...... Cynoglossus kapuasensis (reported only from Kapuas River) 20b. Dorsal-fin rays 110 or greater ...... ® 21

21a. Anal-fin rays 86 to 89; only 15 or 16 scales between dorsolateral and midlateral lines; total vertebrae 51 or 52 ...... Cynoglossus heterolepis 21b. Anal-fin rays 92 to 105; 19 to 22 scales between dorsolateral and midlateral lines; total vertebrae 57 to 61 ...... Cynoglossus suyeni

22a. Blind side with 2 lateral lines (dorsal and medial) ...... Cynoglossus bilineatus 22b. Blind side with only a medial lateral line ...... Cynoglossus borneensis

23a. Labial papillae unbranched (Fig. 11a); caudal vertebrae 44 or more ...... ® 24 23b. Labial papillae branched (Fig. 11b); caudal vertebrae usually 44 or less ...... ® 25 labial papillae 24a. Two lateral lines on eyed side unbranched labial papillae of body; dorsal-fin rays 133 to a) b) branched 143; anal-fin rays 102 to 112 ...... Paraplagusia longirostris Fig. 11 mouth 24b. Three lateral lines on eyed side of body; dorsal-fin rays 133 to 143; anal-fin rays 87 to 91...... Paraplagusia sinerama

25a. Eyed side of body pale uniformly brown without ocelli; anal-fin rays 66 to 80; dorsal-fin rays 105 or fewer ...... Paraplagusia blochii 25b. Eyed side of body brown with pale ocelli of various sizes; anal-fin rays 81 to 88; dorsal-fin rays more than 105 ...... Paraplagusia bilineata

List of species occurring in the area The symbol 0 is given when species accounts are included. A question mark indicates that occurrence in the area is uncertain. ? Cynoglossus abbreviatus (Gray, 1834) Cynoglossus arel (Schneider, 1801) 0 Cynoglossus bilineatus (Lacepède, 1802) Cynoglossus borneensis (Bleeker, 1858) 0 Cynoglossus cynoglossus (Hamilton-Buchanan, 1822) Cynoglossus feldmanni (Bleeker, 1853) ? Cynoglossus gracilis Günther, 1873 Cynoglossus heterolepis Weber, 1910 ? Cynoglossus interruptus Günther, 1880 ? Cynoglossus itinus (Snyder, 1909) Cynoglossus kapuasensis Fowler, 1905 Cynoglossus kopsi (Bleeker, 1851) Cynoglossus lida (Bleeker, 1851) 0 Cynoglossus lingua Hamilton-Buchanan, 1822 Cynoglossus macrophthalmus Norman, 1926 Cynoglossus maculipinnis Rendahl, 1921 Cynoglossus mccullochi Norman, 1926 Pleuronectiformes: Cynoglossidae 3895

Cynoglossus microlepis (Bleeker, 1851) Cynoglossus monopus (Bleeker, 1849) ? Cynoglossus ogilbyi Norman, 1926 0 Cynoglossus puncticeps (Richardson, 1846) ? Cynoglossus semilaevis Günther, 1873 Cynoglossus suyeni Fowler, 1934 Cynoglossus waandersi (Bleeker, 1854) 0 Paraplagusia bilineata (Bloch, 1787) Paraplagusia blochii (Bleeker, 1851) Paraplagusia longirostris Chapleau, Renaud, and Kailola, 1991 Paraplagusia sinerama Chapleau and Renaud, 1993 Symphurus strictus Gilbert, 1905 Symphurus gilesii (Alcock, 1889) Symphurus luzonensis Chabanaud, 1955 Symphurus marmoratus Fowler, 1934 Symphurus microrhynchus (Weber, 1913) Symphurus orientalis (Bleeker, 1879) Symphurus regani Weber and De Beaufort, 1929 Symphurus schultzi Chabanaud, 1955 Symphurus septemstriatus (Alcock, 1891) Symphurus strictus Gilbert, 1905 Symphurus undatus Gilbert, 1905 Symphurus woodmasoni (Alcock, 1889)

References Chapleau, F. and C.B. Renaud. 1993. Paraplagusia sinerama (Pleuronectiformes, Cynoglossidae), a new Indo-Pacific tongue with a revised key to species of the . Copeia, 1993(3):798-807. Chapleau, F., C.B. Renaud, and P.J. Kailola. 1991. Paraplagusia longirostris, a new (Cynoglossidae) from Australia and Papua New Guinea. Japan. J. Ichthyol., 38(3):239-244. Menon, A.G.K. 1977. A systematic monograph of the tongue soles of the genus Cynoglossus Hamilton-Buchanan (Pisces: Cynoglossidae). Smithson. Contrib. Zool., (238):129 p. Munroe, T.A. 1992. Interdigitation pattern of dorsal-fin pterygiophores and neural spines, an important diagnostic character for symphurine (Symphurus: Cynoglossidae: Pleuronectiformes). Bull. Mar. Sci., 50(3):357-403. 3896 Bony Fishes

Cynoglossus arel (Schneider, 1801) Frequent synonyms / misidentifications: Cynoglossus macrolepidotus (Bleeker, 1851) / None. FAO names: En - Largescale tonguesole; Fr - Langue à grandes écailles; Sp - Lengua escamuda.

Diagnostic characters: Body elongate, its depth 20 to 26% standard length. Eyes with a small scaly interorbital space. Snout obtusely pointed; rostral hook short; corner of mouth reaching posteriorly to or beyond lower eye, about midway between gill opening and tip of snout. Dorsal-fin rays 116 to 130. Anal-fin rays 85 to 98. Caudal-fin rays usually 10. Two lateral lines on eyed side of body; none on blind side. Midlateral-line scales 56 to 70. Scales large, ctenoid on eyed side of body; cycloid (smooth) on blind side; 7 to 9 scale rows between lateral lines on eyed side of body. Total vertebrae 50 to 57. Colour: eyed side uniform brown, with a dark patch on gill cover, blind side white. Size: Maximum total length about 38 cm, commonly 20 to 30 cm. Habitat, biology, and fisheries: Inhabits muddy and sandy bottoms on the continental shelf to depths of about 125 m. Feeds predominantly on bottom-living invertebrates. Caught mainly with bottom trawls. Marketed mostly fresh or frozen; also dried-salted. Distribution: Indo-West Pacific from Red Sea and western coast of India to Indonesia, China (including Taiwan Province), southern Japan, and the Philippines; not to New Guinea or Australia. Pleuronectiformes: Cynoglossidae 3897

Cynoglossus bilineatus (Lacepède, 1802) Frequent synonyms / misidentifications: Cynoglossus quadrilineata (Bleeker, 1851) / None. FAO names: En - Fourlined tonguesole; Fr - Langue à quatre lignes; Sp - Lengua de cuatro líneas.

Diagnostic characters: Body elongate, its depth 22 to 29% standard length. Eyes with a small scaly interorbital space. Snout rounded; rostral hook short; corner of mouth reaching posteriorly beyond lower eye, nearer to gill opening than to tip of snout. Dorsal-fin rays 107 to 113. Anal-fin rays 80 to 88. Caudal-fin rays usually 12. Two lateral lines on both sides of body. Midlateral-line scales 88 to 96. Scales ctenoid on eyed side of body; cycloid on blind side; 13 to 16 scale rows between lateral lines on eyed side of body. Total vertebrae 51 to 53. Colour: eyed side brown with an irregular dark blotch on gill cover, blind side white. Size: Maximum total length about 40 cm, commonly 15 to 30 cm. Habitat, biology, and fisheries: Inhabits muddy and sandy bottoms on the inner continental shelf to about 80 m. Feeds predominantly on bottom-living invertebrates. Caught mainly with bottom trawls. Described as excellent table fish. Marketed mostly fresh and frozen; also dried-salted. Distribution: Widespread in Indo-West Pacific along entire northern part of the area to Taiwan Province of China, southward to Philippines and northern coast of Australia and New Guinea (but absent from waters to the east of New Guinea); also, westward to Pakistan and southern and west coasts of India. 3898 Bony Fishes

Cynoglossus cynoglossus (Hamilton-Buchanan, 1822) Frequent synonyms / misidentifications: Cynoglossus sumatranus (Bleeker, 1851); C. bengalensis (Bleeker, 1853) / None. FAO names: En - Bengal tonguesole.

Diagnostic characters: Body elongate, its depth 22 to 34% standard length. Eyes not contiguous. Snout rounded; rostral hook short; corner of mouth not reaching posteriorly beyond vertical through posterior margin of lower eye, nearer to tip of snout than to gill opening. Dorsal-fin rays 95 to 102. Anal-fin rays 72 to 78. Caudal-fin rays usually 10. Two lateral lines on eyed side of body, none on blind side. Midlateral-line scales 70 to 90. Scales on both sides of body ctenoid; 12 to 14 scale rows between lateral lines of eyed side. Total vertebrae 44 to 48. Colour: eyed side uniformly brown-grey, with vague dark marbling. Size: Maximum total length about 20 cm, commonly 10 to 15 cm. Habitat, biology, and fisheries: Inhabits muddy and sandy bottoms, often in shallow areas, including estuaries and brackish waters. Feeds mostly on a variety of bottom-living invertebrates. Caught mainly with bottom trawls, and also with beach seines in estuaries. Marketed mostly fresh or frozen; also dried-salted. Distribution: Indo-West Pacific region from Pakistan and northeastern and central coasts of India eastward to Gulf of Thailand, Indonesia, and the Philippines; apparently not found in New Guinea or northern Australia. Pleuronectiformes: Cynoglossidae 3899

Cynoglossus lingua (Hamilton-Buchanan, 1822) Frequent synonyms / misidentifications: None / None. FAO names: En - Long tonguesole.

Diagnostic characters: Body very elongate, its depth 17 to 22% standard length. Eyes with a small interorbital space. Snout obtusely pointed; rostral hook short; corner of mouth reaching well beyond vertical through posterior margin of lower eye, much nearer to gill opening than to tip of snout. Dorsal-fin rays 126 to 138. Anal-fin rays 97 to 114. Caudal-fin rays 10. Two lateral lines on eyed side of body, none on blind side. Midlateral-line scales 90 to 101. Scales comparatively large, ctenoid on eyed side of body, cycloid (smooth) on blind side; only 11 or 12 scale rows between lateral lines on eyed side of body. Total vertebrae 57 to 66. Colour: eyed side reddish brown, sometimes with irregular brown-black patches, with a large black blotch on gill cover. Size: Maximum total length around 40 cm, commonly 20 to 30 cm. Habitat, biology, and fisheries: Inhabits mainly shallow muddy and sandy bottoms on the inner continental shelf, often entering estuaries. Feeds predominantly on a variety of benthic invertebrates. Taken mainly with bottom trawls. Marketed mostly fresh and frozen, also dried-salted. Distribution: Widespread in Indo-West Pacific from Red Sea and western coast of India to Gulf of Thailand, to the Philippines and Indonesia; not to New Guinea or Australia. 3900 Bony Fishes

Cynoglossus puncticeps (Richardson, 1846) Frequent synonyms / misidentifications: None / None. FAO names: En - Speckled tonguesole; Fr - Langue tachetée; Sp - Lengua moteada.

Diagnostic characters: Body elongate. Eyes with a narrow interorbital space. Snout rounded; rostral hook short; angle of jaws not reaching posteriorly beyond vertical through posterior margin of lower eye, a little nearer to tip of snout than to gill opening. Dorsal-fin rays 90 to 100. Anal-fin rays 72 to 78. Caudal-fin rays usually 10. Two lateral lines on eyed side of body; none on blind side. Midlateral-line scales 78 to 99. Scales ctenoid on both sides of body; 15 to 19 scale rows between lateral lines on eyed side. Total vertebrae 44 to 49. Colour: eyed side yellow-brown, with very distinct irregular dark brown blotches, often forming irregular cross bands; some rays of dorsal and anal fins dashed with dark brown. Size: Maximum total lengths around 18 cm, commonly 8 to 10 cm. Habitat, biology, and fisheries: Inhabits muddy and sandy bottoms in estuaries and on the continental shelf to about 140 m. Feeds predominantly on bottom-living invertebrates. Caught mainly with bottom trawls and beach seines. Marketed mostly fresh and frozen; also dried-salted. Distribution: Widespread in Indo-West Pacific throughout most of northern part of area from Gulf of Thailand to Taiwan Province of China and southern China, southwards to northwestern and central coasts of northern Australia and New Guinea, and eastward to the Philippines. Pleuronectiformes: Cynoglossidae 3901

Paraplagusia bilineata (Bloch, 1787) Frequent synonyms / misidentifications: Paraplagusia marmorata (Bleeker, 1851) / None. FAO names: En - Doublelined tonguesole; Fr - Langue à deux lignes; Sp - Lengua de dos líneas.

Diagnostic characters: Body elongate. Eyes with a scaly interorbital space. Snout rounded; rostral hook long and reaching posteriorly well beyond vertical through posterior margin of lower eye; corner of mouth not reaching beyond vertical through posterior margin of lower eye, nearer to gill opening than to tip of snout. Mouth fringed with branched labial papillae. Dorsal-fin rays more than 105. Anal-fin rays 81 to 88. Two or 3 lateral lines on eyed side of body, none on blind side. Scales ctenoid on both sides of body. Colour: eyed side brown, sometimes spotted or marbled with darker patches; blind side tinged with yellow. Size: Maximum total length about 35 cm, commonly 15 to 25 cm. Habitat, biology, and fisheries: Inhabits muddy and sandy bottoms on the inner continental shelf, often found in shallow and even estuarine (brackish) waters. Feeds predominantly on bottom-living invertebrates. Caught mainly with bottom trawls, occasionally also with beach seines. Marketed mostly fresh or frozen; also dried-salted. Distribution: Widespread in Indo-West Pacific throughout most of northern part of area to Taiwan Province of China, southern China and southern Japan, eastward to the Philippines, and southward to New Guinea and northern and eastern Australia; westward to Red Sea.

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