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Perciformes: Trachinoidei: Pinguipedidae 3501

PINGUIPEDIDAE (= Parapercidae, Mugiloididae) by J.E. Randall iagnostic characters (based on ): Body moderately elongate and little compressed, Dexcept posteriorly (size to about 30 cm); head pointed; eyes often jutting slightly above dorsal profile of head, oriented as much dorsally as laterally. Mouth moderately large, protractile, and terminal or with lower jaw slightly projecting; recurved canine teeth in an outer row at front of jaws, those in lower jaw with 3 to 5 on each side (see Fig. 1 in key), the most lateral largest; a band of villiform teeth behind anterior canines, soon narrowing on side of jaws; an outer row of incurved conical teeth on side of jaws, a few as large recurved canines; small teeth on vomer in a chevron-shaped patch of 1 to 3 rows; teeth on palatines present or absent. A strong sharp spine posteriorly on opercle, and a lesser spine at upper end of margin of subopercle. Gill membranes united, free from isthmus except anteriorly; branchiostegal rays 6. Along dorsal fin with IV or V spines and 20 to 24 soft rays, the soft portion clearly higher than the spinous; anal fin with I spine and 16 to 20 soft rays; caudal fin variable in shape, from slightly rounded to lunate, usually with 15 branched rays; pectoral fins with 14 to 21 rays; pelvic fins with I spine and 5 soft rays, the fourth ray longest, the fins inserted below or anterior to base of pectoral fins. Lateral line complete, slightly arched above pectoral fins, the pored scales 38 to 84; scales ctenoid, often becoming cycloid ventrally on abdomen and chest; small scales on opercle and cheeks, none on occiput, interorbital, or snout; no scales on dorsal or anal fins. Colour: most species distinctively though rarely brightly coloured, frequently with dark bars and/or small spots; most, if not all, are sexually dichromatic, though the differences may be slight (males of some species with oblique lines on cheek and operculum, in contrast to spots or no markings on females).

Habitat, biology, and fisheries: The species of Parapercis, the only in the area, live mainly on open sedimentary or rubble bottoms, some in the vicinity of reefs, others on mud substrata; they rest upon the bottom, propping themselves on their pelvic fins. All are carnivorous, feeding principally on benthic crustaceans, especially crabs and shrimps, occasionally on small fishes. They are easily approached, in general, underwater. Several have been shown to be protogynous hermaphrodites, and it is expected that this will apply to all members of the genus. Males are territorial and maintain harems. Few species are large enough to be of commercial importance as food (and then only of limited value).

Similar families occurring in the area Percophidae (especially Bembrops and Chrionema): 2 separate dorsal fins, the first with VI spines (IV or V in Pinguipedidae).

Percophidae Key to the species of Pinguipedidae occurring in the area 1a. Lateral-line scales 68 to 76; caudal fin emarginate; 2 prominent dark brown to black spots, one above other, at base of caudal fin; maximum standard length at least 25 cm .....® 2 1b. Lateral-line scales 38 to 65; caudal fin slightly rounded, truncate, slightly double emargi- nate, or with a slightly prolonged upper lobe (except emarginate to lunate in P. schauin- slandii); 2 black spots at base of caudal fin present or absent; maximum standard length 23 cm, rarely more than 20 cm ...... ® 3 3502 Bony Fishes

2a. Body very elongate, its depth 7.3 in standard length; dorsal part of body yellow-olive without dark spots; head violet with some orange colour below eye and 3 yellow lines extending anteriorly from eye; no black on spinous portion of dorsal fin . . . . . Parapercis elongata (Viet Nam) 2b. Body not very elongate, its depth 4.7 to 6.1 in standard length; whitish with 6 large quadrangular brown spots on back (may contain small pale blotches and may be red in fish from deep water) and a second series of 6 similar but more round dark spots on lower side of body; head with blue lines extending anteriorly from eye and across interorbital; spinous portion of dorsal fin black between first and fifth spines ...... Parapercis nebulosa (Western Australia, Queensland, and New South Wales)

3a. Palatine teeth present ...... ® 4 3b. Palatine teeth absent ...... ® 7

4a. Lateral-line scales 38 to 44; pectoral-fin rays 14 or 15 (usually 14); 3 U-shaped dark bars dorsally on body followed by 2 more posteriorly with the bottom of U missing (hence as double bars); a series of 9 dusky bars on side of body, every other one extending from the bottom of the dorsal markings ...... (western Pacific from southern Japan to Queensland) 4b. Lateral-line scales 48 to 64; pectoral-fin rays 15 to 18; colour not as above ...... ® 5

5a. Third dorsal-fin spine longest; dorsal-fin rays 21; anal-fin rays 17; pectoral-fin rays 15; lateral-line scales 48 to 52; canine teeth at front of lower jaw 10; a series of 9 vertically elongate dark brown bars on ventral half of body with a pair of dark blotches (or groups of dark brown spots) in upper part of each pale interspace; a narrow dusky Y-shaped bar linking each ventral dark bar to a series of dark spots on back ...... Parapercis cylindrica (western Pacific east to Fiji and the Marshall Islands) 5b. Fifth dorsal-fin spine longest; dorsal-fin rays 23; anal-fin rays 18 to 21; pectoral-fin rays 17 to 20; lateral-line scales 53 to 64; canine teeth at front of lower jaw 6 to 8; colour not as above ...... ® 6

6a. Lateral-line scales 53 to 58; canine teeth at front of lower jaw 8; red with 5 narrow slightly oblique blackish bars on upper half of body; no large black spot in spinous part of dorsal fin ...... Parapercis muronis (southern Japan to Philippines) 6b. Lateral-line scales 64; canine teeth at front of upper jaw 6; no dark bars on body; scales of dorsal part of body and postorbital head with a brown spot, thus forming longitudinal rows; a large black spot basally in spinous part of dorsal fin ...... Parapercis striolata (a single specimen, 15.5 cm total length, taken at 310 m off Kai Islands, Indonesia)

7a. Anterior rays of soft portion of dorsal fin prolonged and filamentous (all but the last 5 or 6 rays filamentous in large adults), the second to fourth longest, 1.7 to 3.8 in standard length (over the range of 6.7 to 11.4 cm standard length); 7 dusky bars on upper half of body containing 3 or 4 dark brown dots in their lower part, the last 6 as 3 pairs; below each pair, on lower side, a large dusky blotch; 2 small dark brown spots, one above the other, at base of caudal fin ...... Parapercis filamentosa (Hainan to Thailand and Indonesia) 7b. No elongate filamentous rays in soft portion of dorsal fin; colour not as above ...... ® 8

8a. Caudal fin emarginate to lunate; 8 or 9 subquadrangular red to dark brown spots along back alternating with similar but more vertically elongate red spots on lower side; 2 narrow bright red bars at pectoral-fin base; spinous portion of dorsal fin black at base, deep red distally; a longitudinal row of black spots along middle of soft portion of dorsal fin ...... Parapercis schauinslandii (Indo-Pacific) 8b. Caudal fin rounded, truncate, slightly double emarginate, or with a short prolonged upper lobe; colour not as above ...... ® 9 Perciformes: Trachinoidei: Pinguipedidae 3503

9a. Dorsal-fin spines IV; vertical dark bars or elongate spots on ventral half of body ...... ® 10 9b. Dorsal-fin spines V; vertical dark bars on ventral half of body present or absent ...... ® 12

10a. Dorsal-fin rays 22 or 23; anal-fin rays 19 or 20; caudal fin, including corners, rounded; a narrow brown stripe on upper side of body, bordered above by a narrower, nearly white stripe, ending in a white-edged black spot about size of pupil on upper base of caudal fin, the spot followed by 2 yellow bars; 12 narrow yellow bars below brown stripe on ventral half of body ...... Parapercis fuscolineata (depths of 170 to 187 m, Philippines) 10b. Dorsal-fin rays 20 or 21; anal-fin rays 17; ventral half of body white, with a series of 10 brown to brownish red bars or vertically elongate spots, each containing a black spot; posterior half of caudal fin usually with a large white to yellow spot, sometimes containing small dark brown to black spots ...... ® 11

11a. Total gill rakers on first gill arch 17 to 19; a row of small blackish spots in outer third of anal fin; males without an ocellus above opercle ...... Parapercis clathrata (western Pacific east to Marshall Islands and Samoa) 11b. Total gill rakers on first gill arch 14 to 17 (rarely 17); no row of small blackish spots in outer third of anal fin; males with an ocellated black spot above opercle next to upper end of gill opening (females with a smaller dark brown spot at this location) ...... Parapercis millepunctata (Pitcairn Group and Marshall Islands to Mauritius and Maldives)

12a. Canine teeth in outer row at front of lower jaw 6 (3 on each side; Fig. 1)...... ® 13 12b. Canine teeth in outer row at front of lower jaw 8 (4 on each side) ...... ® 19

13a. Anal-fin rays 19; pectoral-fin rays 18 or 19 ...... ® 14 upper jaw 13b. Anal-fin rays 16 or 17; pectoral-fin rays 17 or 18 ...... ® 15

14a. Eye not very large, its diameter more than 4 times in lower head length of adults; lateral-line scales 57 to 60; jaw third or fourth dorsal-fin spine longest; last interspi- nous membrane of dorsal fin connected to base of first soft ray; light red dorsally, shading to white ven- trally, the scale edges brown except ventrally; 2 lon- gitudinal rows of indistinct light red blotches on body, ending in 2 dark red spots on base of caudal fin; snout light blue with oblique yellow lines . . . . . (Philippines to Indonesia and northwestern Australia, west on continental 3 canine teeth on each side shelf to Persian Gulf) 14b. Eye very large, about 3 times in head length; lateral- Fig. 1 dentition of Parapercis line scales 64; fifth dorsal-fin spine longest; last inter- schauinslandii spinous membrane of dorsal fin connected horizon- tally to first ray; body with yellow bars; lateral line grey; snout and eye yellow-olive; dorsal and anal fins golden yellow ...... Parapercis flavescens (2 specimens, 7.4 and 8.9 cm, taken at 360 m off Isle of Pines, New Caledonia)

15a. Lateral-line scales 49 to 55; no dark brown bars on lower part of body; a single row of small dark spots in soft portion of dorsal fin ...... ® 16 15b. Lateral-line scales 56 to 65; a series of 9 or 10 dark brown bars on lower part of body; 3 rows of small dark spots in soft portion of dorsal fin ...... ® 18 3504 Bony Fishes

16a. Eye very large, its diameter longer than snout, 2.7 to 2.95 times in head length; caudal fin truncate with rounded corners; total gill rakers on first gill arch 17; 2 rows of dark brown spots about pupil size along side of body, each row continuing as 2 more spots in caudal fin; a series of small dusky spots near base of dorsal fin ...... Parapercis biordinis (Western Australia at 23ºS, and Philippines) 16b. Eye not very large, usually shorter than snout, 3.2 to 3.8 times in head length; caudal fin truncate, the upper 3 branched rays prolonged as a short lobe; total gill rakers on first gill arch 14 to 16; no rows of small dark brown spots on side of body; a series of 6 to 8 subquadrangular dusky bars along back, the scales within bars with dark brown flecks on edges ...... ® 17

17a. Body elongate, its depth about 6.2 times in standard length; eye moderately large, about 3.2 times in head length; no dark bar under eye (only a dusky spot on preorbital below front of eye), and no large dusky area on side of snout; no broad zone of dark brown spots from nape and occiput to operculum; a row of small dark brown spots, one per membrane, in outer third of soft portion of dorsal fin ...... Parapercis rufa (depths of 85 to 90 m, Philippines) 17b. Body not elongate, its depth 4.85 to 5.1 times in standard length; eye not large, 3.5 to 3.8 times in head length; a triangular brown bar below eye, and side of snout broadly dusky; a broad zone of small dark brown spots from nape and occiput to posterior part of operculum; a dark brown spot at base of each soft ray of dorsal fin (Somalia, Indonesia, and southern Japan) ...... Parapercis somaliensis

18a. Lateral-line scales 59 to 65; 3 longitudinal series of large whitish spots on body, the dark grey-brown interspace a coarse reticulum; a dark grey-brown band from nape to interop- ercle containing a black ring just above gill opening; a broad dark grey-brown bar below eye; 3 black spots on each side of abdomen ...... Parapercis tetracantha (Japan to Indonesia and Bay of Bengal) 18b. Lateral-line scales 56 to 60; a midlateral white stripe on body (sometimes interrupted by narrow vertical dusky extensions of dark brown bars of lower half of body), extending nearly to end of caudal fin where bordered above and below by black spots (also a few small black spots within white on fin); scales on back above white stripe with dark edges, many with outer half dark brown; an orange-yellow bar at base of pectoral fins ...... Parapercis xanthozona (Japan to New South Wales, east to New Caledonia, Solomons, and Palau; East Africa)

19a. Posterior margin of preopercle and opercle strongly serrate; cheek scales ctenoid; body elongate, its depth 5.7 to 6.9 in standard length; edges of scales of upper two-fifths of body dark brown, the centres pale; 8 V-shaped dark bars faintly visible on upper part of body, becoming darker at the lower part of each V; a row of 8 large dark brown spots on lower side of body; 2 prominent oval black spots, one above the other, basally in caudal fin ...... Parapercis diplospilus (a single specimen, 7.5 cm total length, taken at 38 m, Philippines) 19b. Margin of preopercle and opercle not serrate; scales on cheek cycloid; body not elongate, its depth 4.8 to 5.8 times in standard length; colour not as above ...... ® 20

20a. Pectoral-fin rays 14 to 16 (usually 15); anal-fin rays 16 or 17; scales on opercle cycloid; lower two-thirds of body whitish with 8 narrow red bars, each containing 2 dark spots (black or darker red), one at top of bar and one at level of lower third of body; a small dark red to black spot on abdomen above base of pelvic fins; spinous portion of dorsal fin cream with a broad oblique black bar (except posteriorly) ...... Parapercis multiplicata (southern Japan to Great Barrier Reef, east to Pitcairn, west to Western Australia) 20b. Pectoral-fin rays 16 to 19; anal-fin rays 17 to 19; scales on opercle ctenoid; colour not as above ...... ® 21 Perciformes: Trachinoidei: Pinguipedidae 3505

21a. Pectoral-fin rays 17 to 19; anal-fin rays usually 17; last membrane of spinous portion of dorsal fin connected to first dorsal-fin ray nearly at height of fifth dorsal-fin spine; upper third of body greenish to brownish grey, flecked with dark brown, the lower two-thirds white with 3 or 4 longitudinal rows of small dark brown spots, sometimes linked by narrow dark stripes except lower row as discrete yellow-edged black spots; caudal fin with a very large black spot centrobasally in fin ...... (East Africa to Fiji) 21b. Pectoral-fin rays 16 or 17; anal-fin rays usually 18; last membrane of spinous portion of dorsal fin connected to first dorsal-fin ray nearly at its base; light brownish red to grey-brown dorsally, shading to whitish ventrally, with 6 large subquadrangular dark reddish brown spots on ventral half of body, narrowly linked to less distinct, saddle-like, dark spots dorsally on body; lower third to fourth of caudal fin dark brownish red ...... Parapercis maculata (southern Japan to Persian Gulf)

List of species occurring in the area The symbol 0 is given when species accounts are included. 0 Parapercis alboguttata (Günther, 1872) Parapercis biordinis Allen, 1976 0 Parapercis clathrata Ogilby, 1910 0 Parapercis cylindrica (Bloch, 1797) Parapercis diplospilus Gomon, 1981 Parapercis elongata Fourmanoir, 1967 0 Parapercis filamentosa (Steindachner, 1879) Parapercis flavescens Fourmanoir and Rivaton, 1979 Parapercis fuscolineata Fourmanoir, 1985 0 Parapercis hexophtalma (Cuvier, 1829) Parapercis maculata (Bloch and Schneider, 1801) 0 Parapercis millepunctata (Günther, 1860) 0 Parapercis multiplicata Randall, 1984 Parapercis muronis (Tanaka, 1918) 0 Parapercis nebulosa (Quoy and Gaimard, 1824) Parapercis rufa,newname1/ 0 Parapercis schauinslandii (Steindachner, 1900) 0 Parapercis snyderi Jordan and Starks, 1905 0 Parapercis somaliensis Schultz, 1968 Parapercis striolata (Weber, 1913) 0 Parapercis tetracantha (Lacepède, 1802) 0 Parapercis xanthozona (Bleeker, 1849)

References Cantwell, G.E. 1964. A revision of the genus Parapercis, family Mugiloididae. Pac. Sci., 18(3):239-280. Randall, J.E. 1984. Two new mugiloidid fishes of the genus Parapercis. Fresh. Mar. Aquar., 7(12):41-49.

1/ Parapercis rosea Fourmanoir, 1985 is a junior homonym of Parapercis rosea (Liénard, 1829) (Percis rosé Liénard corrected to P. rosea Liénard by Sauvage, 1891:317). Fourmanoir described the colour of Parapercis rosea as rose-brown. The new name rufa is proposed, taken from the Latin for reddish. 3506 Bony Fishes

Parapercis alboguttata (Günther, 1860) En - Bluenose . Maximum total length about 27 cm. Usually taken by trawling in depth range of 50 to 120 m. A coastal species of the continental shelf of Asia, northwestern Australia, and the large islands of Indonesia. Parapercis smithii (Regan), P. tesselata (Herre), and P. cephalus Kotthaus are syno- nyms.

Parapercis clathrata Obilgy, 1910 En - Latticed sandperch. Largest specimen, 17.5 cm. A shallow-water species generally found on sand or rubble near reefs. Occurs from the Marshall Islands and Samoa to the western Pacific where it ranges from the Ryukyu Islands to the Great Barrier Reef.

Parapercis cylindrica (Bloch, 1797) En - Sharpnose sandperch. Maximum total length about 23 cm. Occurs in calm areas on sand, rubble, seagrass, or weedy bottoms, generally in the vicinity of shallow reefs. Marshall Islands and Fiji to the western Pacific from southern Japan to central New South Wales. Perciformes: Trachinoidei: Pinguipedidae 3507

Parapercis filamentosa (Steindachner, 1879 En - Threadfin sandperch. Maximum total length about 18 cm. Taken by trawling; the probable habitat mud. Known from Singapore, Thailand, Hainan, and Java. Parapercis hainanensis Lin and P. longifilis Herre are synonyms.

Parapercis hexophtalma (Cuvier, 1829) En - Speckled sandperch. Maximum total length about 28 cm. An inshore species generally found on sand or rubble substrata near coral reefs. Fished throughout its range, but no special fishery exists. Occurs from Samoa and the Marshall Islands to the Red Sea and coast of East Africa south to Natal; in the western Pacific from the Ryukyu Islands to the southern Great Barrier Reef. Parapercis polyophtalma (Cuvier) is a synonym.

female

male

Parapercis millepunctata (Günther, 1860) En - Spotted sandperch. Maximum total length at least 18 cm total length (largest specimen). A common shallow-water species usually found in clear water on sand and rubble around coral reefs. Known throughout the islands of Oceania except Hawaii and Easter Island; in the western Pacific from the Ryukyu Islands to the Great Barrier Reef; ranges west in the Indian Ocean to the Maldives and Mauritius. Parapercis cephalopunctata (Seale) is a synonym. 3508 Bony Fishes

Parapercis multiplicata Randall, 1984 En - Redbarred sandperch. Maximum total length about 12 cm. Usually found on rubble or sand near coral reefs in more than 30 m. The only known localities to date: Ryukyu Islands, Indonesia, Western Australia, Great Barrier Reef, New Caledonia, and Pitcairn.

Parapercis nebulosa (Quoy and Gaimard, 1824) En - Barred sandperch. Maximum total length about 25 cm. Usually found on silty sand or weedy bottoms in shallow bays, but has been taken by trawling in 30 m. Caught throughout its range, but no special fishery exists. Occurs from Western Australia to Queensland and New South Wales. Records from the western Indian Ocean are misidentifications of Parapercis robinsoni Fowler. P. emeryana (Richardson) is a synonym.

Parapercis schauinslandii (Steindachner, 1900) En - Redspotted sandperch. Maximum total length about 13 cm. A species of clear water on sand or rubble bottoms, often near coral reefs, generally at depths greater than 20 m. Occurs throughout the islands of Oceania, including Hawaii and Pitcairn Group (but not Easter Island); in the western Pacific from Honshu to the Great Barrier Reef; west in the Indian Ocean to the coast of East Africa south to Natal. Not recorded from the seas of the Arabian Peninsula or the southern coast of Asia. Perciformes: Trachinoidei: Pinguipedidae 3509

Parapercis snyderi Jordan and Starks, 1905 En - U-mark sandperch. Maximum total length about 11 cm. Generally found on silty sand or rubble areas in the vicinity of coral reefs. Western Pacific from southern Japan and Korea to the southern Great Barrier Reef.

Parapercis somaliensis Schultz, 1968 En - Somali sandperch. Maximum total length 15 cm. Taken by trawling in 50 to 72 m. Known only from 3 localities: Somalia, Red Sea, and south of Sumbawa, Indonesia.

(from Schultz, 1968)

Parapercis tetracantha (Lacepède, 1802) En - Reticulated sandperch. Attains a total length of about 26 cm. A clear-water species found in and around shallow coral reefs. Occurs from southern Japan to Indonesia and the Bay of Bengal. Parapercis cancellata (Cuvier, 1829) is an invalid replacement name. 3510 Bony Fishes

Parapercis xanthozona (Bleeker, 1849) En - Yellowbar sandperch. Maximum total length about 23 cm. Generally found in the protected waters of bays or lagoons. Western Pacific from Honshu to central New South Wales, east to Fiji. Also 2 records from East Africa (Zanzibar and Natal). The East African form is a little different in colour and warrants further investigation. The eastern Australian form may represent a distinct species, Parapercis stricticeps (De Vis).

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