Parascombrops philippinensis ( Günther, 1880 ), Gunther, 1880

Schwarzhans, Werner W. & Prokofiev, Artem M., 2017, Reappraisal of Synagrops Günther, 1887 with rehabilitation and revision of Parascombrops Alcock, 1889 including description of seven new species and two new genera (Perciformes: Acropomatidae), Zootaxa 4260 (1), pp. 1-74 : 48-51

Parascombrops philippinensis ( Günther, 1880)

Figs. 2F , 7J , 8A , 10B , 11E , 13J , 15 G – L, 26, 27, 36, Tables 2–7

Acropoma philippinense Günther 1880: 51 (lectotype designated here: BMNH 1879.5.14.167; Philippines, Challenger station 201, 150– 186 m).

Synagrops malayanus Weber, 1913: 196 , fig. 52 (lectotype designated here: ZMA 112.453 , Timor Sea , Indonesia, Siboga: stn 302, 216 m).

Note: Synagrops philippinensis is one of the most commonly cited species of the group, often in check-lists without description or figure. If the published accounts did not contain information of most of characters used here for species diagnosis, we were unable to verify the record, so was not included.

Material examined (240 specimens): BMNH 1879.5 .14.167, lectotype, 70 mm SL, Philippines, Challenger station 201, 07°03’N, 121°48’E, 150–186 m ; BMNH 2016.9.14.1-4, 4 specimens, paralectotypes, 44, 63, 64, 74 mm SL, same data as lectotype ; BMNH 1986.10 .9.76, 64 mm SL, Sumbawa , Saleh Bay ; BMNH 1986.10 .9.88, 71 mm SL, off Bali; AMS I.21835-016, 68 mm SL, 09°21’S, 133°12’E, 156–164 m ; AMS I.21844-004, 58 mm SL, 10°01’S, 133°54’E, 80–84 m; BSKU 29045, 37603-04, 39445-49, 39477, 39507-10, 39513, 68618, 72773, 74367, 74985, 77449, 77814, 78371, 87910, 88320, 91384, 92536-42, 108395-98, 108634, 112682, 112684, 112688 -89, 112859, 112862, 116496, totally 47 specimens, 40–73 mm SL, all from Kochi ( Mimase and Saga fish markets) , Japan; CAS 238010, 67.5 mm SL, 13°08’N, 124°04’E, 363–385 m (otolith only) ; CAS 34418, 3 specimens, 59–65 mm SL, Philippines, off Batanga Islands ; NMMB-P3545, 2 specimens, 53–60 mm SL, Kaohsiung , Taiwan, 100 m; IOM, uncatalogued, 126 specimens, 20.5–75.0 mm SL, Nha Trang Bay , Vietnam; NMMB-P7474, 4 specimens, 60.5– 66 mm SL, Da-si fishmarket, Ilan , Taiwan; NMMB-P11346, 4 specimens, 43–63.5 mm SL, Dong-gang, Pingtung, southwestern Taiwan; NMMB-P22436, 54.5 mm SL, Da-si fishmarket, Iland , Taiwan; NMMB-P 23103, 54 mm SL, Dong-gang, Pingtung, southwestern Taiwan ; NSMT-P 30481, 59.5 mm SL, Tosa Bay; NSMT-P 53294, 3 specimens, 52–66.5 mm SL, 32°35’N, 129°52’E, 80 m ; NSMT-P 53618, 4 specimens, 54–74.5 mm SL, 32°33’N, 129°52’E, 80 m; NSMT-P 57379, 2 specimens, 33.5– 37 mm SL, 33°18’N, 133°33’E, 99 m; NSMT-P 59313, 3 specimens, 50.5– 57 mm SL, Tosa Bay, 150 m ; NSMT-P 65240, 76 mm SL, Kyushu, Kagoshima, Iwamoto fish market; NSMT-P 69717, 43.5 mm SL, 31°29’N, 127°10’E, 105 m ; NSMT-P 78079, 64 mm SL, Tosa Bay, 130 m ; NSMT-P 78351-52, 2 specimens, 56.5– 62 mm SL, Tosa Bay, 150 m ; NSMT-P 92053, 2 specimens, 34.5– 48 mm SL, Sagami Bay ; NSMT-P 96964, 3 specimens, 40.5– 45 mm SL, 07°50’N, 118°00’E; NSMT-P 101146, 56 mm SL, 21°50’N, 117°00’E; USNM 99325, 4 specimens, 72.5–80.5 mm SL, 08°48’N, 123°31’E, 366 m; ZMA 112.453 , lectotype of Synagrops malayanus , 52 mm SL, Timor Sea, Indonesia, Siboga : stn 302, 10°27’S, 123°28’E, 216 m ; ZMA 112.454 paralectotype of Synagrops malayanus , 54 mm SL, Savu Sea, Lobetobi Straits, Indonesia, Siboga : stn 303, 08°27’S, 122°54’E, 247 m ; ZMUC P44326-37 coll. Prokofiev, 12 specimens, 40–50 mm SL, Nha Trang, Vietnam .

Diagnosis. Small, moderately deep-bodied species with slightly convex dorsal head profile. Anal fin II + 7. Pectoral fin with 14–16 rays, pectoral-fin length 21.5–27.2% SL. Gill rakers 15–18. Pseudobranchial filaments 15–24. First anal-fin pterygiophore long, straight, with rather broad hollow tip. Palatines and ectopterygoids narrow, usually with 1, rarely 2, rows of denticles. Orbital diameter 9.9–12.9% SL. Snout about 65–90% of orbital diameter. Posterior edge of maxillary plate straight. No longitudinal ridges on preopercular lobe. Otolith moderately elongate (OL:OH = 1.6–1.85).

Description. Counts and measurements are summarized in Tables 2–7. Snout bluntly pointed; interorbital space slightly convex. Posterior edge of maxilla almost straight to very weakly concave, with postero-dorsal and postero-ventral angles rounded to sharpened, and postero-ventral angle not extended; posterior rim of the maxilla vertical. Preopercular lobe without longitudinal ridges; denticles of hind margin serration variably expressed, not extending into crests on preopercular lobe; inner edge of preopercle with 2–5 small denticles not extending along ventral branch. Two anal-fin spines. First anal-fin pterygiophore broad, long, straight, with broad hollow tip, reaching last pair of pleural ribs. First haemal spine with a variably developed but mostly weak posterior expansion. Pelvic fin spine serrated along its outer edge; all other fin spines smooth.

Dentition. Premaxilla with a pair of strong canines near symphysis, followed posteriorly by a wide band of minute teeth. Dentary with a pair of strong canines and a band of small conical teeth near symphysis, followed posteriorly by a row of small conical teeth and 2–4 enlarged canines. Vomer V-shaped with 1–2 rows of granular teeth anteriorly and 1–3 long teeth posteriorly on each side. Palatines narrow, with 1 row of granular long teeth anteriorly and 1–2 rows granular teeth posteriorly. Ectopterygoid narrow, with 1, rarely 2, rows of denticles. Tongue toothless.

Otolith morphology (n = 31). Otolith moderately slender, thin, up to about 6 mm length. OL:OH = 1.6–1.85; OL:OT about 7–8. Dorsal rim shallow, with feeble mid-dorsal angle and distinct postdorsal angle very close to posterior rim. Ventral rim deepest slightly in front of its middle, below collum of sulcus. Rostrum well developed, long, sharply pointed; weak excisura and antirostrum. Posterior rim blunt to nearly vertically cut. All rims smooth or slightly undulating. Inner face slightly convex with slightly supramedian-positioned sulcus. Sulcus shallow, with narrow, long, slightly deepened cauda and wide, shallow ostium. Ostium about twice as wide as cauda or less, with distinct colliculum. Cauda slightly widened posteriorly, almost straight and with rounded tip, terminating close to posterior tip of otolith. CaL:OsL = 1.05–1.25. Dorsal depression long, narrow. Ventral furrow distinct, close to ventral rim of otolith anteriorly, running parallel and at some distance to ventral otolith rim at mid section and posteriorly and approaching caudal tip. Outer face slightly concave, rather smooth.

Size. Small fishes, to about 80 mm SL.

Discussion. Parascombrops philippinensis is similar in size to P. pellucidus and P. parvidens (for distinction see respective discussions to the two species). It is tempting to interpret P. philippinensis as small, juvenile or subadult specimens of larger species such as P. argyreus or P. nakayamai , of which few similarly small specimens have been found. However, there are several significant differences that can not all be explained by ontogenetic changes. In the case of P. argyreus one of the distinctive characters, the number of pseudobranchial filaments, varies to some extent with ontogenetic growth and hence is difficult to use for diagnosis, requiring comparison of specimens of similar size ( Fig. 27 ). The most reliable characters for distinction are the absence of longitudinal ridges on the preopercle in P. philippinensis (vs present in P. argyreus ), the shape of maxilla (see Figs. 8A, E ; 11E, D) and the shape of the first anal-fin pterygiophore. Parascombrops philippinensis differs from P. nakayamai in a shorter snout (snout 65–90% of orbital diameter vs 87–113%), in the dentition of the vomer, palatines and ectopterygoids ( Figs. 9F , 10B ), notably in the narrower ectopterygoid tooth patch with 1–2 rows of granular teeth (vs 3–4 rows) and in the higher number of gill rakers (15–18 vs 11–15). The dentition of the vomer and the gillraker count can reflect ontogenetic changes and may not always be relaible characters for distinction ( Fig. 27 ). The snout / orbit diameter relation shows a rather sharp boundary ( Fig. 27 ) and therefore, together with the development of the dentition on the ectopterygoid, represent the best characters for distinction between the two species.

A re-examination of the type material of Synagrops malayanus ( Fig. 26C – D ) revealed no morphological differences to P. philippinensis and can be attributed to the range of variation seen in that species. We therefore follow Schultz (1940), Katayama (1960) and Mochizuki (1989) in synonymizing Synagrops malayanus with Parascombrops philippinensis .

Geographic and bathymetric distribution. Parascombrops philippinensis is a widely distributed and common species in the low latitude tropics of the Western Pacific, between 80 and 385 m depth, although probably less widely distributed than may be assumed from earlier published check-lists. The latter is a result of many small specimens of Synagrops having been recorded as this species, but have not been subject to this review. Parascombrops philippinensis is particularly common in the seas around the Philippines, Vietnam and eastern Indonesia and off the southern coast of Japan and Taiwan. We have no record of geographical overlap with P. argyreus , but there is geographical overlap with P. nakayamai in Taiwan and southern Japan. We also have no record of geographical overlap with P. pellucidus , but it does overlap with P. glossodon and P. parvidens across the entire area of distribution of the two latter (see Figs. 35–37 ).

Figures

FIGURE 2 A – F. Examples of cranial crests: A, Kaperangus microlepis n. gen., CAS 222834, SL 78.5 mm; B, Caraibops trispinosus n. gen., CAS 61040, 97.5 mm SL; C, Parascombrops ohei n. sp., BSKU 72614, holotype, 75 mm SL; D, Parascombrops argyreus, BPBM 23839, 118 mm SL; E, Parascombrops spinosus, USNM 395230, 95.5 mm SL; F, Parascombrops philippinensis, BMNH 1879.5. 14.167, lectotype, 70 mm SL. Lower images with crests highlighted.

FIGURE 7 A – Q. Head profiles of Parascombrops, Caraibops, Kaperangus, Synagrops and Ƒerilus pseudomicrolepis. A, Parascombrops analis, MNHN 1998 - 1056, 82 mm SL; B, Parascombrops argyreus, ZMUC P 44272, 122 mm SL; C, Parascombrops glossodon n. sp., BMNH 1986.10. 9.79. 85, paratype, 4 mm SL; D, Parascombrops madagascariensis n. sp., MNHN 2008 - 1769, holotype, 68.5 mm SL; E, Parascombrops mochizukii n. sp., NMMBA-P 782, holotype, 166 mm SL; F, Parascombrops nakayamai n. sp., BSKU 102255, holotype, SL 97 mm; G, Parascombrops ohei n. sp., BSKU 72614, holotype, 75 mm SL; H, Parascombrops parvidens n. sp., BSKU 98886, holotype, 57.5 mm SL; I, Parascombrops pellucidus, USNM 44428, 64.5 mm SL; J, Parascombrops philippinensis, ZMA 112.453 (paralectotype of P. malyanus), 52 mm SL; K, Parascombrops serratospinosus, CAS 238016, 67.5 mm SL; L, Parascombrops spinosus, USNM 395230, 95.5 mm SL; M, Parascombrops yamanouei n. sp., NMMB-P 23097, paratype, 70 mm SL; N, Caraibops trispinosus, CAS 61040, 97.5 mm SL; O, Kaperangus microlepis, CAS 222834, 78.5 mm SL; P, Synagrops japonicus, ZMUC P 44324, 159 mm SL; 7 Q, Ƒerilus pseudomicrolpis, CAS 61164, 91 mm SL.

FIGURE 8 A – F. Dentition and shape of maxillary: A, long, fang-like teeth on symphyses and laterally on dentary, maxillary with straight posterior edge (Parascombrops philippinensis, BMNH 1879.5. 14.167, lectotype, 70 mm SL); B, no long teeth on premaxillary or dentary, maxillary with straight posterior edge (Parascombrops parvidens n. sp., BSKU 98888, paratype, 62 mm SL); C, D, mouth cavity with teeth patches on vomer, palatine and ectopterygoid (8 c = Parascombrops nakayamai n. sp., CAS 34870, 81.5 mm SL; 8 D = Parascombrops serratospinosus, AMS I. 22822 - 017, 72 mm SL); E, maxillary with ventroposterior extension and concave posterior edge (Parascombrops argyreus, BPBM 23839, 118 mm SL); F, teeth patch on tongue (Parascombrops glossodon n. sp., ZMMGU 23812, paratype, 80 mm SL).

FIGURE 9 A – H. Dentition on vomer, palatines and ectopterygoids: A, Parascombrops analis, AMS I. 20919 - 038, 107.5 mm SL; B, Parascombrops argyreus (B 1 = view of mouth roof, BPBM 23829, 119 mm SL; B 2 = lateral close-up view of left palatine and ectopterygoid, MNHN 1998 - 1041, 92 mm SL); C, Parascombrops glossodon n. sp., AMS I. 21835 - 026, paratype, 72 mm SL; D, Parascombrops madagascariensis n. sp., MNHN 1988 - 1443, paratype, 62 mm SL; E, Parascombrops mochizukii n. sp., NMMB-P 3887, paratype, 98 mm SL; F, Parascombrops nakayamai n. sp., AMS I. 2449 - 008, paratype, 93.5 mm SL; G, Parascombrops parvidens n. sp., BSKU 99887, paratype, 58 mm SL; H, Parascombrops ohei n. sp., BSKU 0 63370, paratype, 87 mm SL.

FIGURE 10 A – H. Dentition of vomer, palatines and ectopterygoids: A, Parascombrops pellucidus, USNM 392727, 84 mm SL; B, Parascombrops philippinensis, BMNH 2016.9. 14.1, paralectotype, 74 mm SL; C, Parascombrops serratospinosus, BSKU 16827, 67 mm SL; 10 D, Parascombrops spinosus, MNHN 1989 - 0411, 82 mm SL; E, Kaperangus microlepis, CAS 222834, 96.5 mm SL; F, Parascombrops yamanouei n. sp., NMMB-P 11343, paratype, 82 mm SL; G, Caraibops trispinosus, CAS 61040, 97.5 mm SL; H, Ƒerilus pseudomicrolepis, CAS 61164, 91 mm SL.

FIGURE 11 A – E. Preopercular serrations: A, inner and outer edges serrated, serrations of outer edge extending onto preopercle as radial crests (Kaperangus microlepis, CAS 222834, 78.5 mm SL); B, inner and outer edges serrated, serration of outer edge extending onto preopercle as radial crests, and small spike present at upper central portion (Parascombrops analis, MNHN 1998 - 1056, 82 mm SL); C, inner and outer edges serrated, no serrations on preopercle, two ridges on central portion of preopercle (Parascombrops argyreus, BPBM 23839, 118 mm SL); D, inner and outer edges serrated, serrations of outer edge extending onto preopercle as radial crests, and one ridge on central portion of preopercle (Parascombrops parvidens n. sp., BSKU 15221, paratype, 74 mm SL); E, inner and outer edges serrated, no serrations or ridges on preopercle (Parascombrops philippinensis, BMNH 1879.5. 14.167, lectotype, 70 mm SL). Right images show characters enhanced by drawing and annotation.

FIGURE 13 A – Q. Anal-fin pterygiophores and shape of large first anal-fin pterygiophore of genera Parascombrops, Caraibops, Kaperangus, Synagrops and of Ƒerilus pseudomicrolepis. A, Parascombrops analis, BSKU 94343, 74.5 mm SL; B, Parascombrops argyreus, AMS I. 42758 - 006, 120 mm SL; C, Parascombrops glossodon n. sp., USNM 436684, paratype, 73 mm SL; D, Parascombrops madagascariensis n. sp., MNHN 2008 - 1769, holotype, 68.5 mm SL; E, Parascombrops mochizukii n. sp., NMMBA-P 3887, paratype, 98 mm SL; F, Parascombrops nakayamai n. sp., BSKU 102255, holotype, 97 mm SL; G, Parascombrops ohei n. sp., BSKU 72614, holotype, 75 mm SL; H, Parascombrops parvidens n. sp., BSKU 15219, paratype, 70 mm SL; I, Parascombrops pellucidus, USNM 44428, 65.5 mm SL; J, Parascombrops philippinensis, BMNH 2016.9. 14.1, paralectotype, 64 mm SL; K, Parascombrops serratospinosus, CAS 238016, 67.5 mm SL; L, Parascombrops spinosus, MNHN 1989 - 0411, 82 mm SL; M, Parascombrops yamanouei n. sp., NMMB-P 15571, paratype, 102 mm SL; N, Caraibops trispinosus, CAS 61040, 97.5 mm SL; O, Kaperangus microlepis, CAS 222834, 78.5 mm SL; P, Synagrops japonicus, MNHN 1998 - 0993, 84 mm SL; Q, Ƒerilus pseudomicrolpis, CAS 61164, 91 mm SL.

FIGURE 15 A – AC. Otoliths (inner faces and representative ventral views): A – F, Parascombrops pellucidus (A – C = BMNH 1939.5.24.892 - 894, D – F = BMNH 1939.5.24.880 - 891); G – L, Parascombrops philippinensis (G – H = NSMT-P 59313, I = BSKU 116496, J = NSMT-P 57379, K – L = ZMA 112. 454, Synagrops malayanus paralectotype); M – O, Parascombrops spinosus (M – N = MCZ 169467, O = USNM 395230); P – R, Parascombrops serratospinosus (USNM 99324); S – U, Parascombrops yamanouei n. sp. (NMMB-P 23097); V – X, Caraibops trispinosus (15 V – W = USNM 407771, 15 X = USNM 407745); Y – AA, Kaperangus microlepis (CAS 222834); AB – AC, Ƒerilus pseudomicrolepis (USNM 407758).

FIGURE 26 A – D. Parascombrops philippinensis (Günther, 1880); A – B, BMNH 1879.5. 14.167, lectotype, 70 mm SL; C – D, Synagrops malayanus Weber, 1913, lectotype, ZMS 112.453, 52 mm SL.

FIGURE 27 A – D. Number of pseudobranchial filaments (A) and gill rakers (B) against specimen size in Parascombrops argyreus, P. nakayamai and P. philippinensis; and snout length versus orbital diameter (C) and snout length as % of orbital diameter as a function of SL (D) in P. nakayamai and P. philippinensis.

FIGURE 35. Geographic distribution of Parascombrops analis, P. glossodon, P. madagascariensis and P. yamanouei. Large symbols indicate location of holotypes. One symbol may represent several samples.

FIGURE 36. Geographic distribution of Parascombrops argyreus, P. pellucidus and P. philippinensis. Large symbols indicate location of holotypes. One symbol may represent several samples.

FIGURE 37. Geographic distribution of Parascombrops mochizukii, P. nakayamai, P. ohei, P. parvidens, and P. serratospinosus. Large symbols indicate location of holotypes. One symbol may represent several samples.