Cypselurus poecilopterus crassus, Shakhovskoy & Parin, 2022

Shakhovskoy, Ilia B. & Parin, Nikolay V., 2022, A review of the flying fish genus Cypselurus (Beloniformes: Exocoetidae). Part 2. Revision of the subgenus Poecilocypselurus Bruun, 1935 with descriptions of three new species and five new subspecies and reinstatement of Exocoetus apus Valenciennes and E. neglectus Bleeker, Zootaxa 5117 (1), pp. 1-109 : 8-9

Cypselurus poecilopterus crassus subsp. nov.

Synonymy and bibliography.

Exocoetus poecilopterus . Günther 1866: 291–292 (description; Taiwan).

Cypsilurus poecilopterus . Jordan & Starks 1903: 542 (short description; Hakodate, Japan).

Cypselurus poecilopterus . Lindberg 1947: 144 (listed; Japan, Korea). Woods & Schultz 1953: 184–185, 187 (morphometry; in part: Japan). Abe 1954: 984–990, Pl. 196 (description, distribution; West Pacific; in part). Tomiyama et al. 1958: 243 (short description, figure; Japan). Imai 1958b: 42, Pl. 40 figs. 1–4 (early life history stages; Japan). Imai 1959: 81–85, Pls. 39–41 (description, distribution, early life history stages; Japan, north to Hokkaido). Tsukahara 1959: 183–184, figs. 42–43 (juveniles; Amakusa Is., Japan). Abe 1960: 149 (listed; Japan, Kuroshio Current). Parin 1960a: 228–231, 278–279, 281, fig. 9 (description; North-West Pacific; in part). Parin 1962: 225–229 (distribution; Sea of Japan and adjacent waters; in part). Lindberg & Legeza 1965: 221, 228, 230–231, fig. 217 (short description (after Abe 1954), distribution; Japan). Nishimura 1965: 66 (listed; Sea of Japan). Ueno 1971: 77 (listed; Hokkaido and adjacent waters). Gushiken 1973: 60, fig. 318 (Okinawa). Masuda et al. 1975: 180 (short description; southern Japan). Yoshino et al. 1975: 71 (listed; Ryukyu Is.). Chen 1978: 291–294, 298–299 (osteology, distribution, identification key; Taiwan). Parin 1984: EXOC Cyp 12 (in part: figure (after Abe 1954)). Chen 1987: 13, 19, 23, 25, 52, 151, Figs. 3-12, 3-33G, 3-34E, 4-6, 6-9, 6-20 , Tabs. 2-1, 3-1, 4-6 (description, distribution, early life history stages; North-West Pacific; in part). Chen 1988: 282–283 (early life history stages, figures, distribution; Japan). Nagasawa & Torisawa 1991: 365 (listed; Hokkaido). Shao et al. 1992: 34, figs. E–F ( Taiwan (Ken-Ting National Park)). Chen 1993: 191–192, Pl. 43 fig. 2 (short description; Taiwan). Senou et al. 2006: 431 (listed; Sagami Sea). Ichimaru 2007: 39 (listed; Goto Is.). Mok 2008: 24 (listed; Eastern Taiwan). Nishida et al. 2008: 287, 290 (listed as associated with drifting seaweeds; Northern Kyushu, Japan). Nagase et al. 2009: 812 (listed; Tanegashima, Japan). Motomura et al. 2010: 82, fig. 55 (listed; Yaku-shima I.). Shakhovskoy & Parin 2010: 581–602 [559–579], figs. 1–2 (description, morphometry, distribution; in part: waters of Japan and Taiwan). Chang et al. 2012: 542–545, 547 (distribution, fishery; Taiwan). Huang & Ou, 2012: 513, fig. 1 (fishing grounds; Taiwan). Iwatsuki et al. 2017: 34 (listed; Hyuga Nada, Southwestern Japan). Motomura & Harazaki 2017: 15 (listed; Yaku-shima I., Japan). Ueda et al. 2017: 2 (karyotype; Iejima, Japan). Koeda & Akita 2018: 2, 4, fig.1D (listed, photo; northeast of Kubura, Japan). Nakae et al. 2018: 229 (listed; Amami-oshima I., Ryukyu Is., Japan). Shakhovskoy 2018: 56–59, 64–67, 78, figs. 14–15, 18, S4 (distribution according to oceanographic factors; in part: north-western Pacific). Chang 2020: 8 (listed; Orchid I., Taiwan). Hata 2020: 159 (short description, photos, fishery; Osumi, Japan).

? Cypselurus oligolepis (non Bleeker). Chen 1993: Pl. 43 fig. 1 ( Taiwan; in part).

Holotype ( Fig. 1c ). ZMMU P-24450 , R/V Vityaz, Cruise 46 , Sta. 6165 , sample 19, 37°28’N 146°36’E, 20 August 1969, captured with dip-net. Length 80 mm SL (juvenile). D 12, A 9, P I 14, Spred 26, Str 10, Sp.br 25 (7 + 18), Vert 42 (26 + 16). Measurements (in % SL): aA 77.5, aD 68.7, aV 55.2, cV 33.8, pV 41.5, c 22.8, po 11.3, o 8.7, ao 2.1, io 12.8, Hc 22.3, H 23.7, h 9.3, Dc 30.0, lP 69.8, lP 1 36.3, lV 39.7, lD 21.8, lA 11.1, HD 13.7 ?, HA 9.5, p 20.3. Body robust, snout very short, jaws of equal size. Jaw teeth small, unicuspid. Palatine teeth present. Body pale brown to brown. Pectoral fins with large elongated dark brown spots arranged in oblique rows upon paler background. Tip of pectoral fin protruding slightly beyond end of dorsal-fin base. Pelvic fins pigmented similar to pectoral fins, their tips almost reaching origin of caudal-fin lower lobe. Dorsal fin brown with a row of paler spots near base. Tip of last dorsal-fin ray protruding slightly beyond origin of caudal-fin upper lobe, tip of penultimate ray still a little further back. Anal fin pale brown except for pale area between first three rays. First anal-fin ray beneath 6 th dorsal-fin ray. Caudal fin pale with brown base and two faint dark bands (aggregations of melanophores) on lower lobe.

Paratypes. IORAS 04001 , 3 specimens 53, 71 and 119 mm SL, 30°05’N 142°43’E, 6–7 October 1955 .

Differential diagnosis. Cypselurus poecilopterus crassus differs from other subspecies of C. poecilopterus in wider body and head in juveniles: in fish 25–80 mm SL p usually> 20 vs. usually <20% SL and io usually> 14.5 vs. usually <14.5% SL; in juveniles 80–120 mm SL p> 18 vs. usually <18% SL and io ≥ 12 vs. usually <12% SL. In addition, juveniles of this subspecies have on average a shorter snout and pelvic fins (see Shakhovskoy & Parin 2010).

Cypselurus p. crassus also differs from other subspecies ecologically, occurring in colder waters of North-West Pacific (see Shakhovskoy 2018: fig. S4).

Coloration in life. According to Abe (1954), the body of adults prior to preservation is dark blue dorsally and whitish ventrally. The inner side of the pectoral fin is bluish and has (to the 8th ray) several irregular rows of rounded, brownish-black spots and several irregular rows of rounded, yellow spots between the former. The lower part of the pectoral fin (below the 8 th ray) is pale bluish. The upper side of the first rays of pelvic fins is blue and the remainder is hyaline. In some smaller specimens the membranes of pelvic fins have a few rows of rounded, brownish-black spots. Chen (1993) described the coloration of the paired fins in fish from waters of Taiwan as: pectoral fins yellowish-green (except in the lower part) with brown, slightly reddish, spots; pelvic fins with yellowish-brown spots. According to Imai (1959), the body of some specimens are violet in color while others have coloration usual for flying fishes. In juveniles about 35 mm SL, the dorsal, pelvic and pectoral fins are greenish-yellow, and the black spots on the pectoral fins are fringed with red. Such pectoral fin coloration is retained to a certain extent in adults. Tsukahara (1959) reported that juveniles 60–70 mm FL are brownish-black in life, their pectoral and pelvic fins dark brownish with brownish-black spots, the dorsal and anal fins dark brownish.

Biology. According to Imai (1958b, 1959), the spawning season in Kagoshima Prefecture is July–August. Small juveniles (6.0– 20.5 mm SL) are captured in July–August near Yakushima and Tanegashima Is. Near Kyushu, juveniles are found in July–September. Tsukahara (1959) suggested that spawning takes place considerably south of Amakusa Is. According to Chen (1987) spawning occurs from late May to July in coastal areas between Taiwan and Kagoshima I. Larvae appeared in the waters off Taiwan in June and in Japanese waters in June–September at temperatures 24.0–28.8°C (greatest number occurred in 25–27°C).

Etymology. The subspecies name reflects extremely wide and deep body in juveniles (from Latin crassus — stout, nourished).

Common names. The name“Japanese spotwing flying fish”(Russian:“ЯПонский ПЯтнокрылый стрижехвост”) is proposed here for C. p. crassus.

Figures

FIGURE 1. Holotypes of Cypselurus poecilopterus subspecies: (A)—holotype of C. p. arabicus subsp. nov. (ZMMU P-24449, 191 mm SL, 16°10’N 60°45’E); (B)—the same fish, pelvic fin; (С)—holotype of C. p. crassus subsp. nov. (ZMMU P-24450, 80 mm SL, 37°28’N 146°36’E).

FIGURE 3. Distribution of Cypselurus starksi (□), C. oligolepis oligolepis (○), C. o. persicus (◊), C. o. georgii ( ) and C . izumii (Δ). Filled symbols indicate capture location for fishes ≤ 100 mm SL, open symbols indicate capture location for fishes> 100 mm SL. Positions of holotypes marked with asterisk.

FIGURE 4. Flying fish drawings from Bleeker 1866-1872: (A)—Exocoetus oligolepis; (B)—E. brachysoma; (C)—E. neglectus.

FIGURE 5. Cypselurus oligolepis oligolepis: (A)—37 mm SL, IORAS 03985, 15°47’N 123°28’E; (B)—43 mm SL, ZMUC uncat., 4°30’N 103°28’E; (C)—76 mm SL, IORAS 03975, 12°28’S 128°05’E; (D)—94 mm SL, IORAS 03978, 5°37’N 94°46’E; (E)—124 mm SL, IORAS 03972, Singapore; (F)—166 mm SL, IORAS 03967, Nha Trang.

FIGURE 6. Size variation of morphometric characters (in % SL) in Cypselurus oligolepis (□— C. o. oligolepis, ○— C. o. apus, ∆— C. o. persicus, +—C. o. georgii), C. clariangulatus (◊) and C. izumii (x): (A)—diameter of eye (o); (B)—width of interorbital space (io); (C)—length of pectoral fin (lP); (D)—length of pelvic fin (lV); (E)—postorbital distance divided by diameter of eye (index po/o); (F)—pectoral fin length divided by pelvic fin length (index lP/lV).

FIGURE 7. Pectoral fins in Cypselurus oligolepis oligolepis: (A)—31.5 mm SL, IORAS 03985, 15°47’N 123°28’E; (B)—76 mm SL, IORAS 03975, 12°28’S 128°05’E; (C)—94 mm SL, IORAS 03978, 5°37’N 94°46’E; (D)—140 mm SL, IORAS 03972, Singapore.

FIGURE 8. Pelvic fins in Cypselurus oligolepis oligolepis: (A)—31.5 mm SL, IORAS 03985, 15°47’N 123°28’E; (B)—70.5 mm SL, CAS 81864, Linapacan I.; (C)—99 mm SL, CAS 81829, 6°56’N 121°45’E; (D)—124 mm SL, IORAS 03972, Singapore.

FIGURE 9. Cypselurus oligolepis persicus: (A)—181 mm SL, USNM 147840 (holotype), Ras Tanura, West Pier; (B)—same specimen, pectoral fin; (C)—182 mm SL, USNM FIN 28135, Saudi Arabia, Jubail (photo by J.E. Randall, from the USNM website); (D)—171 mm SL, MTUF uncat, Kuwait fishmarket (drawing by T.N. Steyker).

FIGURE 10. Holotype of Cypselurus oligolepis georgii (145 mm SL, ZMMU P-24451, road of Suez): (A)—side view; (B)— close up of head; (C)—pectoral fin.

FIGURE 11. Cypselurus oligolepis georgii: (A)—39 mm SL, IORAS 03990, 14°05’N 42°20’E; (B)—56 mm SL, IORAS 03990, 14°05’N 42°20’E; (C)—71.5 mm SL, IORAS 03991, Gulf of Aden; (D)—76 mm SL, IORAS 03990, 14°05’N 42°20’E; (E)—113 mm SL, IORAS 03989, 14°05’N 42°20’E.

FIGURE 12. Pectoral fins in Cypselurus oligolepis georgii: (A)—39 mm SL, IORAS 03990, 14°05’N 42°20’E; (B)—56 mm SL, IORAS 03990, same place; (C)—71.5 mm SL, IORAS 03991, Gulf of Aden; (D)—76.5 mm SL, IORAS 03989, 14°05’N 42°20’E; (E)—111 mm SL, IORAS 03988, 13°14’N 42°59’E; (F)—123.5 mm SL, IORAS 03988, same place.

FIGURE 13. Pelvic fins in Cypselurus oligolepis georgii: (A)—71.5 mm SL, IORAS 03991, Gulf of Aden; (B)—76 mm SL, IORAS 03990, 14°05’N 42°20’E; (C)—103.5 mm SL, IORAS 03989, 14°05’N 42°20’E; (D)—117 mm SL, IORAS 03986, 13°03’N 42°58’E.

FIGURE 14. Cypselurus oligolepis subsp. (131 mm SL, IORAS 03993, Red Sea?).

FIGURE 15. Type of Exocoetus apus (122 mm SL, MNHN B834, Macao): (A)—side view; (B)—view from above. Photos provided by Jonathan Pfliger and Zouhaira Gabsi (MNHN).

FIGURE 16. Cypselurus oligolepis apus: (A)—32 mm SL, AMS uncat., Sek Harbour; (B)—41 mm SL, SOSC uncat., 00°51’S 135°35’E; (C)—94.5 mm SL, SOSC uncat., 01°53’N 127°39’E; (D)—116.5 mm SL, SOSC uncat., 01°20’N 128°01’E; (E)— 134 mm SL, IORAS 04006, Rabaul.

FIGURE 17. Pectoral (A–C) and pelvic (D–F) fins in Cypselurus oligolepis apus: (A)—41 mm SL, SOSC uncat., 00°51’S 135°35’E; (B)—92.5 mm SL, SOSC uncat., 1°00’N 127°56’E; (C)—144 mm SL, IORAS 04006, Rabaul; (D)—32 mm SL, AMS uncat., Sek Harbour; (E)—43 mm SL, CAS 128237 (SU 28237), Madang; (F)—110 mm SL, IORAS 04006, Rabaul.

FIGURE 18. Distribution of Cypselurus oligolepis apus (○) and C. clariangulatus (◊). Filled symbols indicate capture loca- tion for fish ≤ 100 mm SL, open symbols indicate capture location for fish> 100 mm SL. Positions of holotypes marked with asterisk.

FIGURE 19. Cypselurus izumii: (A)—187 mm SL (holotype), IORAS 03995, 20°01’S 115°00’E; (B)—pectoral fin of specimen 174 mm SL (paratype), IORAS 03997, 26°27’S 112°44’E.

FIGURE 20. Holotype of Cypselurus clariangulatus (50 mm SL, ZMMU P-24452, 8°18’N 120°25’E): (A)—side view; (B)— pectoral fin; (C)—pelvic fin.

FIGURE 21. Cypselurus clariangulatus: (A)—32 mm SL, CAS 81823, 8°43’N 122°40’E; (B)—61 mm SL, ZMUC uncat., 2°54’N 120°04’E; (C)—92 mm SL, CAS 81829, 6°56’N 121°45’E; (D)—128.5 mm SL, IORAS 03999, Philippines (Camotes Sea?); (E)—147 mm SL, CAS 81829, 6°56’N 121°45’E.

FIGURE 22. Pectoral (A–B) and pelvic (C–E) fins in Cypselurus clariangulatus: (A)—55.5 mm SL, IORAS 04000, 7°45’N 120°25’E; (B)—147 mm SL, CAS 81829, 6°56’N 121°45’E; (C)—44 mm SL, IORAS 04003, ~ 7°N 135°31’E; (D)—89 mm SL, CAS 81829, 6°56’N 121°45’E; (E)—128 mm SL, CAS 81904, Minis I.

FIGURE 23. Cypselurus neglectus neglectus: (A)—160.5 mm SL, IORAS 04007, Tonkin Bay; (B)—83 mm SL, IORAS 04013, Tonkin Bay. After Parin and Besednov 1965.

FIGURE 24. Size variation of morphometric characters (in % SL) in Cypselurus oligolepis (□), C. clariangulatus (♦), C. izumii (x), C. neglectus (○), C. starksi (◊) and C. nossibe (▲): (A)—cephalo-ventral (head to pelvic-fin origin) distance (cV); (B)—postventral distance (pV), (C)—cephalo-ventral distance divided by postventral distance (index cV/pV); (D)—head length (c); (E)—greatest body depth (H); (F)—greatest head depth (Hc).

FIGURE 25. Cypselurus neglectus neglectus: (A)—33.5 mm SL, IORAS 04024, 3°04’N 109°28’E; (B)—47.5 mm SL, IO- RAS 04029, Surabaya; (C)—65 mm SL, SIO 61-634, 12°04’N 101°59’E; (D)—83 mm SL, CAS 81190, 10°02’N 109°48’E; (E)—128 mm SL, IORAS 04012, Tonkin Bay; (F)—149 mm SL, SOSC Ref.No.190, 6°30’N 118°45’E.

FIGURE 26. Size variation of morphometric characters (in % SL) in Cypselurus neglectus neglectus (+—Tonkin Bay, x—other areas) and C. n. shcherbachevi (□—Indonesia, ○—Andaman Sea, ∆—Bay of Bengal and Arabian Sea): (A)—eye diameter (o); (B)—snout length (ao), (C)—pelvic fin length (lV); (D)—dorsal fin height (HD); (E)—cephalo-ventral distance divided by postventral distance (index cV/pV); (F)—pelvic fin length divided by snout length (index lV/ao).

FIGURE 27. Pectoral (A–C) and pelvic (D–F) fins in Cypselurus neglectus neglectus: (A)—37 mm SL, IORAS 04028, Kieta Harbour, Bougainville; (B)—65 mm SL, SIO 61-634, 12°04’N 101°59’E; (C)—133 mm SL, IORAS 04025, 1°58’N 105°53’E; (D)—37 mm SL, IORAS 04028, Kieta Harbour, Bougainville; (E)—83 mm SL, CAS 81190, 10°02’N 109°48’E; (F)—126 mm SL, SOSC Ref.No.190, 6°30’N 118°45’E.

FIGURE 28. Distribution of Cypselurus neglectus neglectus (○), C. n. shcherbachevi (◊) and C. nossibe (∆). Filled symbols indicate capture location for fishes ≤ 100 mm SL, open symbols indicate capture location for fishes> 100 mm SL. Positions of holotypes marked with asterisk.

FIGURE 29. Holotype of Cypselurus neglectus shcherbachevi (123 mm SL, ZMMU 24453, 14°11’N 82°53’E): (A)—side view; (B)—pectoral fin; (C)—pelvic fin.

FIGURE 30. Cypselurus neglectus shcherbachevi: (A)—33 mm SL, ZMUC uncat., 6°06’N 96°00’E; (B)—48 mm SL, IORAS 04032, 5°00’N 90°52’E; (C)—78 mm SL, same place; (D)—105 mm SL, IORAS 04033, 9°36’N 90°53’E; (E)—122.5 mm SL, IORAS 04034, 13°37’N 86°53’E; (F)—138 mm SL, IORAS 04031 (paratype), 6°33’N 82°18’E.

FIGURE 31. Pectoral fins of Cypselurus neglectus shcherbachevi: (A)—47 mm SL, IORAS 04046, ~2°25’S 107°36’E; (B)— 80 mm SL, IORAS 04034, 13°37’N 86°53’E; (C)—105 mm SL, IORAS 04033, 9°36’N 90°53’E; (D)—122.5 mm SL, IORAS 04034, 13°37’N 86°53’E.

FIGURE 32. Pelvic fins of Cypselurus neglectus shcherbachevi: (A)—81 mm SL (fixed with Bouin solution), IORAS 04041, 6°04’N 98°29’E; (B)—105 mm SL, IORAS 04033, 9°36’N 90°53’E; (C)—138 mm SL, IORAS 04031 (paratype), 6°33’N 82°18’E.

FIGURE 33. Cypselurus nossibe: (A)—47 mm SL, IORAS 04052, 6°58’S 39°40’E; (B)—same specimen, close-up of chin barbel; (C) –63 mm SL, IORAS 04050, 5°37’S 39°54’E; (D)—115 mm SL, IORAS 04049, 8°51’S 41°52’E; (E)—124.5 mm SL, IORAS 04049, same place; (F)—172 mm SL, ZMMU 24454 (holotype), Nossi-Be I., Madagascar; (G)—same specimen, close-up of head.

FIGURE 34. Pectoral fins of Cypselurus nossibe: (A)—47 mm SL, IORAS 04052, 6°58’S 39°40’E; (B)—63 mm SL, IO- RAS 04050, 5°37’S 39°54’E; (C)—115 mm SL, IORAS 04049, 8°51’S 41°52’E; (D)—117,5 mm SL, IORAS 04048, 5°11’S 52°09’E; (E)—172 mm SL, ZMMU 24454 (holotype), Nossi-Be I., Madagascar; (F)—183 mm SL, USNM 294610 (paratype), Shimoni, Kenya.