Psammoperca waigiensis (Cuvier in Cuvier & Valenciennes 1828 )

Iwatsuki, Yukio, Newman, Stephen J., Tanaka, Fumiya & Russell, Barry C., 2018, Validity of Psammoperca datnioides Richardson 1848 and redescriptions of P. waigiensis Cuvier in Cuvier & Valenciennes 1828 and Hypopterus macropterus (Günther 1859) in the family Latidae (Perciformes) from the Indo-West Pacific, Zootaxa 4402 (3), pp. 467-486 : 471-476

Psammoperca waigiensis (Cuvier in Cuvier & Valenciennes 1828)

English name: Waigieu sea perch

Figs. 1C , 2A‒B , Table 1

Labrax waigiensis Cuvier in Cuvier & Valenciennes 1828:83 (type locality: Waigeo , Indonesia) ; Bauchot & Desoutter 1987:72 (Waigeo, Indonesia).

Cnidon chinensis Müller & Troschel 1849:21 (Manila, Philippines).

Psammoperca waigiensis ; Günther 1859:69 (Victoria [= Port Essington], NT, Australia and China; but Australian records questionable, probably P. datnioides ); Günther 1872:426 (Victoria [= Port Essington], NT, Australia, Torres Strait, NSW, Australia, questionable as noted above, Manila, and China); Bleeker 1871 –1876:108, pl. (28) 306, fig. 2 ( Singapore, Bintang, Banka, Java, Madura, Borneo Celebes Waigiu, Manila): Fowler & Bean 1930:181 ( Philippines); Greenwood 1976:77 ( Ceylon [ Sri Lanka], Madras [Chennai], Singapore, Hong Kong, Borneo, Cebu, Culion, Philippines); Fisher & Bianchi 1984:CENTRP Psamm 1 (Arabian Sea?, noted as perhaps extending to Arabian Sea, but questionable); Katayama in Masuda et al. 1984:123 (Ryukyu Islands, Japan); Mohsin & Ambak 1996:219 ( Malaysia and neighboring countries); Rainboth 1996:183 (Cambodian Mekong); Allen 1997:98 (tropical Australia [?] and south-east Asia); Chen et al. 1997:39 (Nansha Islands to South China Coastal Waters); Larson 1999:2430 (as vaigiensis ; tropical east Indo-West Pacific, from Bay of Bengal, Indo-Australian Archipelago and northern Australia [?], Philippines, Japan, and the China Sea. western Pacific); Nakabo 2000:679 (Ryukyu Islands, Japan); Lim in Randall & Lim 2000:608 (South China Sea); Sadovy & Cornish 2000:72 ( Hong Kong); Allen 2000:97 (Calamianes Islands, Philippines); Iwatsuki et al. 2000:98 (Makassar [= Ujung Pandang], South Sulawesi, Indonesia); Hutchins 2001:29 (Western Australia); Nakabo 2002:679 (Ryukyu Islands, Japan), Allen & Adrim 2003:30 ( Indonesia); Kimura et al. in Kimura & Matsuura 2003:43 (Bitung, northern tip of Sulawesi, Indonesia); Otero 2004:85 (coastal Indo-Pacific marine waters); Allen & Erdmann 2012:259 (the East Indies); Kottelat 2013:323 (Inland waters of southeast Asia); Nakae in Kimura et al. 2015:36 (northwestern Johor Strait, Peninsular Malaysia).

Psammoperca vaigiensis ; Boulenger 1895:365 (unjustified emendation of Cuvier’s Labrax waigiensis ).

Holotype. MNHN 0000-0564, 195 mm SL, Waigeo , Indonesia.

Non-type specimens (n = 14). MUFS 9882 , 179 mm SL, Singapore ; MUFS 36075 , 250 mm SL, Okinawa, Japan ; MUFS 43427 , 241 mm SL, Okinawa Island, Japan ; URM P. 979 at OCN, 299 mm SL, Okinawa Island, Japan ; URM P. 13418, Chantaburi fish Market , Thailand ; URM P. 26804, 247 mm SL, Naha fish market (Nahachiku-gyoren), Okinawa Island, Japan ; URM P. 31787 at OCF, 250 mm SL, Chinen , Okinawa Island, Japan ; URM P. 36597 at OCF, 197 mm SL, Itoman , Okinawa, Japan ; URM P. 41585, 208 mm SL, Naha , Okinawa, Japan ; URM P. 43585 at OCF, 272 mm SL, Okinawa Island, Japan ; URM P. 44173, Bintan Island , Indonesia ; ZMB 35 (holotype of Müller & Troschel’s Cnidon chinensis ), 375 mm TL, Manila, Philippines .

Diagnosis. Distinguished from Psammoperca datnioides by the following combination of characters: brownish dark head and body, often golden in colour in live specimens, pored lateral-line scales yellow-edged, Fig. 1C ) and similar yellowish brown colour in preserved specimens; lower counts of pored lateral line scales 46‒48; hind margin of maxilla reaching slightly short of hind margin of eye when mouth closed; tiny scales (not seen with naked eye] but visible through binocular scope) on posterior uppermost part of maxilla ( Fig. 3C‒D ); pelvic-fin spine subequal to 4th dorsal spine; 21 or 22 circumpeduncular scales; moderately lesser body depth (32‒36% [mean 36%] of SL); lower counts, 5½ / 7½‒9½ scale rows above and below lateral line; gill rakers (7 [including 6 rudiments] + 1 + 11 [including 3 rudiments] = 19 [9 rudiments]); last dorsal-fin spine / penultimate dorsal-fin spine 1.5‒1.7.

Description. Counts and proportional measurements as percent of SL of the type and other specimens of Psammoperca waigiensis (Cuvier in Cuvier & Valenciennes 1828) are shown in Table 1.

Body compressed, its depth 2.8‒3.1 times in SL, deepest at dorsal-fin origin; dorsal profile concave in interorbital region, rising steeply (convex) thereafter to dorsal-fin origin; head moderately acute, its length 2.6‒3.0 times in SL; eye oval, height less than width, orbit diameter 3.8‒4.1 times in head length; snout 3.3‒3.9 times in head length; interorbital space 54‒63% of eye diameter; mouth oblique, lower jaw projecting beyond upper one when mouth closed; maxilla progressively deeper posteriorly, extending to a vertical beyond posterior of black iris; villiform teeth present on jaws, palatines, pterygoids and vomer; tongue smooth anteriorly and present posteriorly, respectively; three sharp, strong spines on inferior margin of preopercle, the first antrorse; a retrorse spine at angle of preopercle, the posterior margin of which bears 26‒31 denticulations; a sharp spine at angle of operculum; nares level with middle of eye, separated from eye by distance subequal to horizontal orbit diameter of black iris; posteriormost part of cleithrum with one or two dull spines-like as often on exposed part without skin, just above base uppermost pectoral-fin ray ( Figs. 2A‒C ).

First dorsal fin commencing slightly behind pelvic-fin, third spine longest (III>IV>II>V>VI>VII>I); base of first dorsal fin less than that of second dorsal fin; third anal-fin spine longest (III>II>I) in specimens of 20 cm SL but subequal to third in specimens over 20 cm SL (III = II>I); pectoral fin 8 9‒100% of length of pelvic fin, which has one spine and five rays; distal profiles of pectoral, pelvic, anal and second-dorsal fins rounded; caudal fin rounded, with 9+ 8 in principal rays of upper lobe + lower lobe, respectively; dorsal and ventral procurrent rays (8‒10) + (6‒8), respectively; caudal peduncle depth 54‒58% of its length; scales ctenoid; body and head scaled, except for snout, throat, preorbital, and interorbital regions; dorsal and anal fins with a scaly sheath at base; seconddorsal, caudal, anal and lateral area of pelvic fin densely covered with minute scales; one row of pored lateral line scales on the caudal fin but one or two rows of a few pored scales on caudal fin sometimes present one above and one below median pored lateral line; vertebrae 11 + 14 = 25.

Colouration. In fresh specimens ( Fig. 1B , MUFS 43427, 241 mm SL and MUFS 9882, 179 mm SL), head and body yellowish brown, darker above lateral line and on dorsal region of head, lighter below, ventrally creamy white; fins yellowish brown, inter-radial membrane of dorsal fin yellowish hyaline, spinous portion dark brown; second dorsal fin, caudal and anal fins yellowish brown or hyaline, pectoral and pelvic fins somewhat paler yellowish brown, pelvic-fin spine whitish; vivid yellow longitudinal stripe present from upper part of maxilla to posteroventral part, with one spine as well as yellow posterior hind edge and hind margin of upper opercular yellow; pored lateral line outstandingly yellow. In preserved (70% ethanol) specimens, head and body yellowish tan; yellow longitudinal stripe and yellow on head, body and all fins absent.

Remarks. Psammoperca waigiensis (Cuvier in Cuvier & Valenciennes 1828) , well known as the Waigiou sea perch, is a widely distributed coastal species in the tropical eastern Indian Ocean and western Pacific Ocean (Katayama & Taki 1984; Larson 1999). Fisher & Bianchi (1984) listed P. waigiensis from the Western Indian Ocean, but Manilo & Bogorodsky (2003) considered this to be a questionable record.

Richardson (1848) established the genus Psammoperca (Type species Psammoperca datnioides Richardson 1848 ; type locality: ‘Australia’) in a publication on the fishes collected by the Erebus and Terror Expedition under Sir James Ross. This expedition, however, visited only Van Diemen’s Land (Tasmania) in Australia and, as P. datnioides is a tropical species, it is unlikely that it was collected during the expedition itself. This is confirmed in a footnote by Richardson (1848:1) who remarks: ‘To make the list of Australian species as complete as possible, a few undescribed fish from the western coasts of that country, discovered by the officers of the Beagle surveying ship, have been added to Sir James Ross’s collections’. The Beagle surveyed the coast of Northwestern Australia between the Gulf of Carpentaria and the Swan River (Perth, Western Australia) between 1838‒1841 (Stokes 1846), and it is likely that P. datnioides was collected during these voyages. Unfortunately the holotype of the species in the BMNH is lost (J. Maclaine and C. Fisher pers. comm.) but the original description includes a fine figure (pl. 57, fig. 1 for a whole specimen and fig. 2 for scale; Fig. 2B ). The figure clearly is Psammoperca datnioides , not P. waigiensis , because two clear diagnostic characters are shown: 12 soft dorsal-fin rays and the lower value (ca. 0.9) in last dorsal-fin spine / penultimate dorsal-fin spine (vs. 13 soft dorsal-fin rays and higher value [1.5‒1.7] of last dorsal-fin spine / penultimate dorsal-fin spine in P. waigiensis ), as redescribed above.

Paxton et al. (1989) and Allen et al. (2006) had synonymized P. datnioides under P. waigiensis , but we conclude that Psammoperca datnioides is a valid species of Psammoperca . Boulenger (1895) use of the name Psammoperca vaigiensis is an erroneous emendation of Cuvier’s Labrus waigiensis , and as such is invalid (Article 58A., ICZN 1999).

Müller & Troschel (1849) described Cnidon chinensis from Manila, Philippines, based on a specimen ZMB 35 ( Fig. 2C ; holotype of Cnidon chinensis Müller & Troschel 1849 ) collected around 1831 by Franz Julius Ferdinand Meyen (1804‒1840), Professor of Botany at the University of Berlin, who accompanied a circumnavigation of the world between 1830 and 1832 (P. Bartsch pers. comm.). Although Eschmeyer et al. (2016) regarded ZMB 35 as a non-typical specimen of 375 mm TL (J. Kapp & P. Bartsch [ZMB] pers. comm.), it fits the original description. The described holotype of Müller & Troschel (1849) is ‘14½ Zoll’ TL. Because, in Prussia at that time (after 1816) a Prussian Zoll was a 12th part of a Prussian foot (313.85 mm) ‒ and accordingly was 26.15 mm, and the TL of the specimen should be approximately 379 mm. Thus, allowing for some shrinkage in alcohol, the specimen (ZMB 35, 375 mm TL) is consistent with Müller & Troschel’s (1849) holotype (J. Kapp & P. Bartsch pers. comm.). This value was also reconfirmed from another of Müller & Troschel’s (1849) length measurements. For example, Pimelodus lateristrigus Müller & Troschel 1849 (= synonym of Pimelodella lateristriga [Lichtenstein 1823] ZMB 3038) is 4½ Zoll, meaning ca. 118 mm TL, and actual size is equal to 115 mm TL.

The descriptions of Cuvier and of Müller & Troschel conform rather well. The holotype of Cnidon chinensis has the following diagnostic characters of Psammoperca waigiensis (see Diagnosis below): 47 pored lateral-line scales (46‒48 in P. waigiensis vs. 49‒54 in Psammoperca datnioides ), 13 soft dorsal-fin rays (vs. 12 in P. datnioides ) and hind margin of maxilla slightly less than hind margin of eye when mouth closed (vs. hind margin of maxilla vertically behind center of eye when mouth closed in P. datnioides ). These three characters are not found in P. datnioides . Accordingly, we conclude that Cnidon chinensis Müller & Troschel 1849 is a junior synonym of Psammoperca waigiensis ( Cuvier 1828) .

As explained in the Introduction, the poorly known species Hypopterus macropterus ( Günther 1859) was placed in synonymy with Psammoperca waigiensis Cuvier in Cuvier & Valenciennes 1828 by Greenwood (1976, p. 77) who briefly mentioned this species as a member of the family Latidae . However, it was not discussed in subsequent work by Otero (2004) and its taxonomic identity has remained obscure. This species is redescribed below.

Biology. Shimose & Tachihara (2006) studied age, growth and reproductive biology of 291 specimens of Psammoperca waigiensis from around Okinawa Island, Japan. Opaque otolith zones formed every year (annual rings), correlated with spawning activity. Growth of this species was rapid during the first 2 years, reaching 186.2‒ 270.3 mm in SL. Females (196.6‒334.0 mm SL) were larger than males (186.2‒288.6 mm SL), a result of differential growth between sexes, which started before 2 years of age. Such biological information suggests that P. waigiensis does not grow to more than 50 cm TL.

Distribution. Currently known from Sri Lanka, Bay of Bengal, coasts of the Malay Peninsula, Indonesia, Malaysia, Vietnam, Philippines, China, Taiwan, and the Ryukyu Islands of Japan, but not Australia ( Fowler & Bean 1930; Larson et al. 2013). Günther (1859, 1872) reported this species from Victoria (= Port Essington), NT, New South Wales and Torres Strait, but the species is probably P. datnioides (See Distribution of P. datnioides above). Our study indicates P. waigiensis is unlikely to be distributed in Australia, but further work is needed to confirm its distribution in the region adjacent to northern Australia.

Figures

FIGURE 1. Psammoperca datnioides Richardson 1848 (A–B) and Psammoperca waigiensis (C). A, NTM S.16708-077, Neotype, 138 mm SL, Bullocky Point Reef, Darwin, Northern Territory; B, WAM P. 344852-001, 204 mm SL, Carnarvon Harbour, Western Australia; C, MUFS 43427, 241 mm SL, OkinawaIsland, Ryukyu Is., Japan.

FIGURE2. Holotype (A) of Labrax waigiensisCuvier in Cuvier& Valenciennes 1828, a plate 57(B) of Richardson(1848) for Psammoperca datnioides, and a holotype (C) of Cnidon chinensis Müller & Troschel 1849.

FIGURE3. Snout regionof four latidspecies: A, Lates japonicus, MUFS 11877, 253 mm SL, Miyazaki,Japan; B, Hypopterus macropterus, WAM P.30681-004, 138 mm SL, Shark Bay, Western Australia; C, Psammoperca datnioides, WAMP.34451-001 (024,X), 204 mmSL, Corronban Coastnear Shark Bay, Western Australia; D, Psammoperca waigiensis, Chantaburi,Thailand.