Cypselurus oligolepis oligolepis ( Bleeker, 1865 )

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 : 16-29

Cypselurus oligolepis oligolepis ( Bleeker, 1865)

Synonymy and bibliography.

Exocoetus unicolor (non Valenciennes). Bleeker 1852: 11, 21 (description; Batavia (Java )).

Exocoetus oligolepis Bleeker, 1865: 107 , 109–111 (original description; Indonesia, Singapore; in part?). Günther 1866: 296–297 (description of type; East-Indian Archipelago). Bleeker 1866 –1872: 67, 69–70, Tab. CCXLVIII Pl.Scombres II fig. 3 (description; Indonesia, Singapore; in part?). Günther 1910: 370 (short description; West Pacific; in part).

Cypsilurus oligolepis . Weber & de Beaufort 1922: 180, 189–190 (description; Indo-Australian Archipelago; in part).

Cypselurus oligolepis . Woods & Schultz 1953: 184–185, 187 (morphometry; Siam; in part: USNM 103337).? Chandy 1954: 177, 179–180, fig. 1c (short description; India ( Bombay, Ratnagiri)). Parin 1960a: 222, 254, 255, 279 (description (after Weber & de Beaufort 1922 and Woods & Schultz 1953); western Pacific; in part). Parin & Gorbunova 1964: 232–233, fig. 4a (eggs; Singapore). Jones & Kumaran 1965: 111–112, fig. 5 (short description; Laccadive Archipelago: Androth, Chetlat, Kadamat). Parin & Besednov 1965: 107–108 (description of a juvenile; Tonkin Bay; in part: only specimen 107 mm SL). Munro 1967: 117, 121 (New Guinea; short description; in part). Jinadasa 1971: 15 (short description; Sri Lanka; in part). Hyodo & Tsukahara 1977: 9, 13 (description; north Arabian Sea).? Mai & Bui 1978: 251, 261 (listed; Vietnam; in part?). Jones & Kumaran 1980: 146–147, fig. 124 (short description; Laccadive Archipelago: Androth, Chetlat, Kadamat). Parin 1984: EXOC Cyp 11 (description, figures, distribution; Western Indian Ocean; in part). Talwar & Kacker 1984: 298, 300–301, text-fig. 122 (short description, fishery; India). Chen 1987: 15, Fig. 3-20 , Tab. 3-1, (short description; North-West Pacific; in part). Chen 1993: 191 (short description, figure; Taiwan; in part). Lakshminaraina 1993: 26–27, Fig. 4a (short description, distribution; “Indian seas”; in part). Parin & Lakshminaraina 1993: 55 (description, distribution; South-East India). Rao et al. 2000: 86–87 (short description; Andaman Is., Mayabunder). Barman & Mishra 2006: 3, 11, 13–14 (description; India; in part). Matsunuma 2018: 67 (short description, photo; Vietnam, Ha Long Bay; in part).

Cypselurus starksi (non Abe).? Imai 1958a: 41, Pl. 39 fig. 3 (juvenile; Makurazaki).? Imai 1959: 75–81, Pl. 38b (juveniles; Japan; in part: only juveniles with bands on body and stripes on pectoral and pelvic fins). Kyushin et al. 1977: 40–41 (description, photo; Andaman Sea).

Cypselurus bruuni Kotthaus, 1969: 15–17 , figs. 167, 174 (original description; off Goa, India). Shakhovskoy & Parin 2019: 67–68 (description of the holotype and paratype).

Cypselurus naresii (non Günther).? Yoshida et al. 2013: 60 (photos; Thailand (Chantha Buri); in part: KAUM-I.33236).

Material examined. Two hundred specimens 11–175 mm SL.

South China Sea. Full morphological study . IORAS 03967 (1, 166 mm SL), Nha Trang Bay , 20.06.2003 . IORAS 03968 (1, 44 mm), 3°11’N 109°06’E, 19.03.1977 . IORAS 03969 (1, 139 mm SL), Tonkin Bay , 30.07.1960 . IORAS 03970 (1, 106 mm SL), 20°20’N 106°47’E . CAS 81098* (4, 135– 144 mm SL), 9°31’N 99°55’E, 6- 8.11.1957 . CAS 81124* (3, 141– 150.5 mm SL), 12°33’N 101°16’E, 27.10.1957 . CAS 81179 (1, 50 mm SL), 11°55’N 102°13’E, 20.11.1960 . CAS 81204* (1, 148.3), Gulf of Thailand, Market at Chol Buri , 1.12.1957 . CAS 81943 (1, 64 mm SL), 10°13’N 103°55’E, 10.02.1961 . ZMUC uncatalogued (8, 43– 111 mm SL), 4°30’N 103°28’E, 6- 7.06.1951 .

Partial morphological study. IORAS uncat* (1, 111 mm SL), 18°35’N 106°20’E, 14.07.1961 . CAS 81121* (3, 136– 160 mm SL), 7°22’N 100°43’E, 2.04.1968 . CAS 81170* (1, 154 mm SL), 10°27’N 99°15’E, 27.05.1960 . Institute of Marine Research ( Vietnam) No. 51789* (6, 108– 164 mm SL), Can Bong, 1.08.1976 . ISH 4-1993 * (1, 130 mm SL), off Chumpon, Gulf of Thailand, 6.12.1993 . ISH 5-1993 * (1, 115 mm SL), off Chumpon, Gulf of Thailand, 6.12.1993 . SMF 28116* (1, 109 mm SL), Gulf of Thailand, December 1974 . ZMUC uncat. (1, 63 mm SL), 4°30’N 103°28’E, 6- 7.06.1951 . Unknown collection. No. 1673* (1, 167 mm SL), Thailand, Songhla Lake , 1982 .

Singapore. Full morphological study. IORAS 03971 (1, 46.5 mm SL), 1°58’N 105°53’E, 13.07.1971 . IORAS 03972 (11, 124– 142.5 mm SL), Singapore, 3- 4.11.1962 . IORAS 03973 (11, 118.5– 146 mm SL), Singapore, 8- 9.03.1975 . IORAS 03974 (1, 128 mm SL), Singapore, 23– 26.02.1973 . CAS 130720 ( SU 30722 )* (1, 143 mm SL), Singapore, 18.03.1934 . ZMUC uncat. (5, 43– 143 mm SL), 1°35’N 103°01’E, 11.05.1951 .

Partial morphological study. T. Abe collection No. 10660* (1,128 mm SL), Singapore . ZMUC uncat. (2, 117– 130 mm SL), 1°46’N 104°25’E, 21.06.1951 .

Indonesia and Australia. Full morphological study. IORAS 03975 (1, 76 mm SL), 12°28’S 128°05’E, 3- 4.04.1973 . AMS I.16937-002 (1, 50.5 mm SL), Australia, NT, Darwin , 1970 . AMS I.22137-015 (1, 61 mm SL), 18°46’S 146°26’E, 6.12.1980 . FRSKU 112656 (1, 141 mm SL), 20°30’S 114°51’E, 11.01.1988 . FRSKU 112657 (1, 147.5 mm SL), 20°30’S 114°51’E, 11.01.1988 . FRSKU 112658 (1, 166 mm SL), 20°30’S 114°51’E, 11.01.1988 . FRSKU 112835 (1, 143 mm SL), 15°11’S 122°47’E, 12.01.1989 . FRSKU 112836 (1, 136 mm SL), 15°11’S 122°47’E, 12.01.1989 . FRSKU 112837 (1, 148 mm SL), 15°11’S 122°47’E . FRSKU 112838 (1, 150 mm SL), no data . NTM S.11951-003 (1, 51 mm SL), N of Table Head , Port Essington, N. T., 17.09.1985 .

Partial morphological study. IORAS 03976 (1, 32 mm SL), Surabaya, 24.12.1959 . IORAS 03977 (1, 78 mm SL), 6°59’S 104°49’E, 24.03.1961 . AMS I.33759-008* (1, 31.5 mm SL), 10°11’S 142°56’E, 30.01.1993 . AMS I.23423-004* (2, 126– 127 mm SL), 18°01’S 118°23’E, 1.08.1982 . CSIRO C.726* (1, 170 mm SL), Maret Island , W.A., 17.09.1949 . CSIRO CA.1805* (1, 144 mm SL), Passage Islands , 9.12.1979 . FRSKU 112891 * (1, 146 mm SL), 18°29’S 118°59’E . FRSKU 113560 * (1, 147.5 mm SL), 20°00’S 116°59’E, 9.01.1989 . FRSKU 113896 * (1, 142 mm SL), 20°00’S 117°04’E, 27.11.1989 . NTM S.10593.002* (2, 137– 147 mm SL), West side of Mc Cluer, N . T., 17.10.1982 . NTM S.11661-001* (1, 120 mm SL), Jensen Is., Wessel Is. Group, 4.02.1985 . QM I.6728–6729* (2, 130 mm SL), Thursday Id., N.Q., 26.05.1939 . QM I.16068* (1, 141 mm SL), Haggerstone Id., 19.02.1979 . QM I.17171* (1, 140 mm SL), Torres Strait, 8.11.1974 . SIO 61-56 * (4, 133– 138 mm SL), 10°11’S 139°31’E, 12.10.1960 . USNM 072565 * (1, 147 mm SL), Java, Batavia, 2.04.1909 . WAM 29667.005 * (6, 157– 170 mm SL), 20°29’S 114°54’E, 11.01.1988 .

Andaman Sea and Padang Harbour. Full morphological study. IORAS 03978 (4, 71.5–94 mm SL), 5°37’N 94°46’E, 9.03.1961 . IORAS 03979 (1, 99 mm SL), Andaman Sea, 17.03.1964 . SOSC Ref. No. 360 (1, 55.5 mm SL), 7°11’N 99°22’E, 9.02.1966 . SOSC uncat. (1, 145 mm SL), 7°11’N 99°22’E, 9.02.1966 . ZMUC P34931 (1, 134 mm SL), Padang Harbour, 26.09– 5.10.1929 .

Partial morphological study. ISH 10-1993 * (10, 130– 148 mm SL), Ranong, Thailand, 10.12.1993 .

Bay of Bengal and Arabian Sea. Full morphological study. IORAS 03980 (1, 156 mm SL), Yendi, Karwar , 5.10.1979 . IORAS 03981 (1, 149 mm SL), Yendi, Karwar , 5.10.1979 . IORAS 03982 (1, 170 mm SL), Vizinjam , December 1979 . IORAS 03983 (1, 144 mm SL), 5°20’N 80°01’E, 10- 11.11.1964 . IORAS 03984 (1, 157 mm SL), Thirumullaivasal , July 1991 . CAS 141378 (SU 41378), (1, 158.5 mm SL) , Gulf of Mannar, Crusadai I., 22.01.1941 .

Partial morphological study. CMFRI uncat** (4, 150– 175 mm SL), Elathur Estuary , Calicut 2.11.1979 . CMFRI uncat** (2, 153– 154 mm SL), Karwar , 3.12.1979 . CMFRI uncat** (1, 160 mm SL), Veraval , Chamad, 11.10.1979 . ZMH 4983 * (1, 80 mm SL), off Goa , India, 15.02.1965 . ZMH 4984 * (1, 78.5 mm SL), off Goa , India, 15.02.1965 .

Philippines. Full morphological study . IORAS 03985 (2, 31.5–37 mm SL), 15°47’N 123°28’E, 10.09.1957 . CAS 81829 (11, 36– 99 mm SL), 6°56’N 121°45’E, 6.05.1948 . CAS 81864 (1, 70.5 mm SL), Palawan Prov., Linapacan I., San Miguel, 25.09.1949 . CAS 81904 (1, 69 mm SL), Zamboanga Del Sur, Pilas Group, Minis I., 7.05.1948 . SOSC Ref.No. 190 (3, 107– 136 mm SL), 6°30’N 118°45’E, 27.01.1965 . ZMUC uncat (1, 49.5 mm SL), 9°36’N 125°46’E, 30- 31.07.1951 .

Partial morphological study. IORAS 03985 (8, 10.9–17.2 mm SL), 15°47’N 123°28’E, 10.09.1957 . BSKU 17726 * (1, 34.5 mm SL), 8°19’N 118°09’E, 27.05.1972 . USNM 135803 * # (2, 125 mm SL), Catbalogan, Samar I., 15.04.1908 . USNM 135804 * # (3, 122–134 mm SL), Subig Bay, Luzon I., 6- 8.01.1908 . USNM 135805 * # (2, 121–123.5 mm SL), Catanduanes I., Agoho Pt., 10.06.1909 . USNM 135806 * # (5, 124–132.5 mm SL), San Miguel Bay, Luzon, 5.07.1909 . USNM 294551 * (1, 81.5 mm SL), Luzon I., Albay Bay, 3.05.1976 . USNM 435573 # (1, 123 mm SL), near Dauis, Panglao, 31.07.2016 . ZMUC uncat (3, 26.5–31 mm SL), 9°36’N 125°46’E, 30- 31.07.1951 .

Other areas. Full morphological study. ZIN 4827* (1, 122 mm SL), Pacific Ocean, Dr. Mertens. ZIN 5509* (1, 121 mm SL), Oc. Pacificus—Dr. Schrenk, 1854.

Type material. Full morphological study. BMNH 1866.5.2.31* (1, 147 mm SL), E. Ind. Arch.

Partial morphological study. AMS I.57* (1, 137 mm SL), Malay Arch. RMNH 12054* (8, 126– 147 mm SL), no data.

Types. Ten syntypes of Exocoetus oligolepis Bleeker were studied by the second author (nine have questionable type status). In this work we designate the syntype BMNH 1866.5.2.31 as the lectotype of Cypselurus oligolepis (Bleeker) to clarify the application of the name to the taxon.

The lectotype (ex syntype) of Exocoetus oligolepis Bleeker ( BMNH 1866.5.2.31, E. Ind. Arch.). Length 147 mm SL. D 12, A 8, P I 16, Spred 23, Str 8?, Sp.br 21 (4 + 17), Vert 40 (26 + 14). Measurements (in % SL): aA 81.2, aD 71.4, aV 58.5, cV 1 34.3, pV 40.1, c 25.3, o 9.0, ao 1 7.2, io 1 9.0, H 19.8, Dc 26.8, lP 64.7, lV 32.4, lD 18.7, lA -, HD 12.3, HA -. Pectoral fins gray to 9 th ray, with tip pale. The first anal-fin ray beneath 6 th dorsal-fin ray. Second dorsal-fin ray the longest. Lower jaw shorter than upper jaw. Jaw teeth tricuspid. Palatine teeth present. This specimen was also described by Günther (1866).

Paralectotype (ex syntype (?)) of Exocoetus oligolepis Bleeker ( AMS I.57, Malay Arch., Purch. Mr. Fransis Day 1886). Length 137 mm SL. D 12, A 10, P I 15, Spred 27?, Sp.br 22 (5 + 17), Vert 41 (27 + 14). Measurements (in % SL): cV 1 33.5, pV 43.1, c 25.2, o 9.3, io 1 10.1, Dc 28.5, lP 62.0, lP 1 44.9, lV 29.2. The fish is in bad condition (shrunken). First anal-fin ray beneath 5 th dorsal-fin ray. Jaw teeth in many rows, some teeth tricuspid. Palatine teeth present. This specimen is probably C. oligolepis oligolepis , although we are not certain.

Paralectotypes (ex syntypes (?)) of Exocoetus oligolepis Bleeker ( RMNH 12054, Coll. Dr. Bleeker). Eight specimens 126–147 mm SL have the following counts: D 12, 12, 12, 12, 12, 12, 12, 11; A 8, 8, 8, 8, 8, 8, 8, 8; Spred 27, 26, 25, 24, 26, 25, 23, 26. Judging by the characters provided the vast majority of fish (if not all of them) belong to C. oligolepis oligolepis , although it is possible that some are other subspecies (or even C. neglectus ).

We regard Cypselurus bruuni Kotthaus, 1969 as a junior synonym of C. oligolepis oligolepis . The holotype and the only paratype of this species were studied by the second author, and their characters were given in the first part of this review ( Shakhovskoy & Parin 2019: 67–68).

Description. Meristic and morphometric characters are given in Tables 1–5 , 7 , 10. D 10 –13 (usually 12–13), A 8–9 (rarely 7), P I 12–16 (usually I 13–15), Spred 22–28 (usually 24–26), Str 8–9½ (very rarely 7 or 7½), Sp.br 19–26 (4–8 + 14–18), usually 21–24 (5–6 + 16–17), Vert 39–43 (25–27 + 14–17), usually 40–42 (25–26 + 15–16). Snout short ( Fig. 5 ), lower jaw shorter than upper or of equal length (rarely lower jaw may be slightly longer, especially in juveniles). Jaw teeth small to medium-sized (not visible or barely visible with naked eyes), mainly tricuspid (some specimens also have conical teeth or teeth with additional cusps). Juveniles <90 mm have mainly conical teeth. Teeth arranged in 2–5 rows, in juveniles in 1–3 rows. Palatine teeth always present, numerous (sparse in juveniles <40 mm SL); one juvenile 68.5 mm SL with two teeth on vomer.

Body rather deep. Greatest body depth changes slightly with growth: in juveniles 30–100 mm SL body depth 4.6–6.2 in SL; in fish 120–170 mm SL, 4.0– 5.5 in SL. Body width 0.96–1.52 and caudal peduncle depth 2.01–3.00 in greatest body depth. Greatest head depth and head length do not change with growth, 4.7–5.9 and 3.7–4.4 in SL, respectively. Head length 3.7–4.4 in SL and 0.97–1.27 in dorso-caudal distance. Eyes large, eye diameter decreases strongly with growth ( Fig. 6a ): in juveniles (30–100 mm SL) 7.7–12.6 in SL, 2.0– 3.1 in head length, 1.0– 1.5 in interorbital width and 0.85–1.4 in postorbital distance ( Fig. 6e ); in fish 120–170 mm SL, eye diameter 10.5–13.9 in SL, 2.7–3.4 in с, 0.95–1.45 times in io and 1.1–1.45 in po.

Pectoral fins relatively long, their length increasing with growth to about 100 mm SL, and then nearly constant ( Fig. 6c ): in juveniles 30–100 mm SL pectoral fin 1.5–1.85 in SL and in fish 120–170 mm SL, 1.45–1.65 in SL. Tip of pectoral fin reaching posterior half of dorsal-fin base or slightly beyond, not exceeding middle of caudal peduncle; in juveniles <50 mm SL to about middle of dorsal-fin base. First pectoral-fin ray unbranched, its length increasing with growth: in juveniles 30–100 mm SL it fits 2.55–3.35 in SL and 1.57–1.93 in lP; in fish 120–170 mm SL, 2.3–3.1 in SL and 1.55–1.90 in lP. Pelvic-fin base much closer to posterior edge of head than to origin of caudal-fin lower lobe (cV / pV = 0.63–0.80); pelvic fins not changing position as fish grows. Pelvic-fin length decreasing strongly from juveniles to adults ( Fig. 6d ): in juveniles 30–100 mm SL pelvic fin 2.15–2.8 in SL and 1.22–1.77 in lP ( Fig. 6f ); in fish 120–170 mm SL, 2.8–3.4 in SL and 1.82–2.22 in lP. Tip of pelvic fin of juveniles 30–70 mm SL reaching (nearly reaching) origin of caudal-fin lower lobe or protruding beyond; in juveniles 70–100 mm SL reaching from middle of caudal peduncle to origin of caudal-fin lower lobe; in fish 100–130 mm SL reaching from end of anal-fin base to middle of caudal peduncle and in fish> 130 mm SL reaching from middle to end of anal-fin base (very rarely further).

Anal-fin origin far posterior to dorsal-fin origin (1st anal-fin ray under 4 th –8 th dorsal-fin ray, usually under 5th–7th). Dorsal fin with 2–6 rays more than anal fin. Height of dorsal and, particularly, anal fins decreasing markedly as fish grow. In juveniles 30–100 mm SL, HD 6.3–9.1 and HA 8.3–13.9 in SL; in fish 120–170 mm SL, HD 7.25–10.85 and HA 10.75–14.7 in SL. Longest ray of dorsal and anal fins—2 nd or 3 rd (in a juvenile 46 mm SL 4 th dorsal ray the longest). Tip of last dorsal-fin ray reaching from middle of caudal peduncle to origin of caudal-fin upper lobe (rarely slightly further). Middle and posterior rays of dorsal fin not elongate, penultimate rays usually not extending beyond tip of last ray.

Pigmentation. Body of juveniles ( Fig. 5a–d ) brown or pale brown, ventral side usually paler than dorsal one. In juveniles 25–80 mm SL dark body bands absent or one to four bands on the lower part of body present: near pectoral-fin base, between pectoral and pelvic fins, near pelvic-fin base; in some fish also under eye or above anus. Fish> 80 mm SL without body bands.

The lower surface of the head of juveniles 25–90 mm SL brown with paler chin and distal parts of branchiostegal rays, or in some fish pale with pigmentation mainly under the eye. With growth pigment on lower surface of head begins to disappear (persisting as dots under the eye), and in fish> 105 mm SL, lower surface of head pale with only few dots near lower jaw symphysis. Adults with a few small specks on gill cover and (or) under eye or without specks.

Pectoral fins ( Fig. 7 ) of juveniles 10–80 mm SL brown or dark brown with pale tip and usually with pale transverse band or a row of pale spots (sometimes with only 1–2 spots) in central part ( Fig. 7a–b ). Juveniles from Philippine Sea (IORAS 03985, 15°47’N 123°28’E) also have large pale spots distally ( Fig. 7a ). Occasional specimens with entirely dark fins. In fish (75) 80–110 mm SL, the fins become brown or dark brown to 10 th –11 th ray with pale tip and small pale band (“mirror”) reaching up to 6 th –10 th ray ( Fig. 7c ). In fish ≥ 110 mm SL, pectoral fins brown or dark brown (rarely dark gray) to 8 th –9 th (rarely to 7 th or 10 th –11 th) ray, with narrow pale posterior edging and broad pale tip ( Fig. 7d ). Pigmentation usually extending distally one ray lower than proximally ( Fig. 7d ) or a small “mirror” present.

Pelvic fins of juveniles 10–50 mm SL ( Fig. 8a ) entirely brown or dark brown. Fish from Philippine Sea sometimes with a paler area near tip of 1 st –2 nd ray. Fish (45) 55–90 mm SL usually with a row of 1–5 pale spots ( Fig. 8b ). In fish 95–115 mm SL pigmentation begins to disappear starting with proximal part of the fin and between 5 th –6 th rays ( Fig. 8c ). In fish ≥ 115 mm SL pelvic fins pale ( Fig. 8d ), although few dots or a line between rays may persist occasionally.

Dorsal fin of juveniles 10–45 mm SL pale with melanophores near bases of anterior rays and along fin’s upper margin; posterior margin devoid of pigmentation. In fish 45–95 mm SL, dorsal fin is covered with melanophores (darker near fin base anteriorly and along upper margin of the fin, paler or entirely pale near fin base posteriorly and along last ray). In fish 95–110 mm SL, pigmentation starts to disappear and dorsal fin becomes gray with dark upper margin. In fish> 110 mm SL, dorsal fin gray or translucent, usually with yellowish or brownish tinge near its base.

Anal fin of juveniles 10–17 mm SL transparent, sometimes with few black dots between 2–3 posterior rays distally or near fin base anteriorly. In fish 30–90 mm SL, anal fin pale with a large dark spot posteriorly (occasionally absent) and, sometimes, with melanophores near the fin base anteriorly. In fish 90–100 mm SL, pigmentation starts to disappear, and in fish> 100 mm SL anal fin translucent or, rarely, grayish (in fish 100–110 mm SL a few dots sometimes persist distally).

Caudal fin of fish 10–35 mm SL pale, with dots along rays of lower lobe (except uppermost rays), and sparse dots between upper lobe rays and on fin base. In fish 35–90 mm SL, caudal fin pale with pale brown or brown base and dots between rays of lower lobe and, usually, on upper lobe distally. The dots usually form 1–3 aggregations (bands) on lower lobe and, in fish> 55 mm SL, also 1 band on upper lobe. In fish ≥ 90 mm SL caudal fin pale brown, brown or gray, usually with the distal part of upper lobe darker and paler near fork ( Fig. 5d–f ).

Coloration in life. According to Bleeker (1852), pectoral fins in adults are a beautiful dark blue, and caudal fin also blue.

Maximum size. The maximum length of C. o. oligolepis in the material examined was 175 mm SL (CMFRI uncat, Elathur Estuary, Calicut). The largest female and male were 170 and 157 mm SL, respectively.

Common names. The name “largescale flying fish” is most frequently used for C. oligolepis . The name “common largescale flying fish” (Russian: обыкновенный малочеШуйный стрижехвост) is proposed here for the subspecies C. o. oligolepis .

Intrasubspecies variation. Specimens from different regions slightly differ in frequencies of some counts ( Tables 1–5 ). For example, C. o. oligolepis from Australia and Indonesia have a slightly higher number of anal-fin rays (usually 9 vs. usually 8). Fishes from Philippines more frequently have 11 dorsal-fin rays, and fishes from Indian waters have fewer vertebrae. There are also subtle differences in several measurements— H, h, io, p, HA and lA ( Table 7 ). These minor differences suggest some degree of isolation between populations. Juveniles from the Philippine Sea ( IORAS 03985 , 15°47’N 123°28’E) differ from other juveniles in pigmentation of pectoral (with large pale area near tip, Fig. 7a ) and pelvic (with pale area near 1 st ray) fins, and in higher values of HD and HA. Fishes from the Philippine Sea are worth study based on more representative material.

Biology. Males are mature by 124 mm SL, females by 126 mm SL. Close to mature or mature fish were captured off Nha Trang , Vietnam in June ( IORAS 03967 ) and off Can Bong in August ( Institute of Marine Research No. 51789), in the Gulf of Thailand in October ( CAS 81124), near Singapore in March ( IORAS 03973 ) and November ( IORAS 03972 ), in Padang Harbour in September–October ( ZMUC P34931), near Vizinjam , India in December ( IORAS 03982 ), in the Gulf of Mannar in January ( CAS 141378 ). Small juveniles 17–31 mm SL were captured on 11 May 1951 at 1°35’N 103°01’E, on 6–7 June 1951 at 4°30’N 103°28’E and on 30–31 July 1951 at 9°36’N 125°46’E (all ZMUC uncat.). Juveniles 11–37 mm SL were captured in the Philippine Sea in September ( IORAS 03985 ). These data suggest year-round spawning of C. o. oligolepis , probably with some local seasonality.

Distribution. According to our data ( Fig. 3 ), C. o. oligolepis occurs in the neritic waters of India, Sri Lanka, Indonesia, Thailand, Malaysia, Cambodia, Australia, New Guinea, Philippines and Vietnam, from Veraval, India (CMFRI uncat.) to eastern Philippines (CAS 138417, Mindanao I., Talissayan) and eastern Australia (AMS I.22137- 015, 18°46’S 146°26’E), and from Tonkin Bay (Institute of Marine Research ( Vietnam) No. 51789, Can Bong) to NW Australia (CSIRO CA.1805, Passage Islands).

Cypselurus o. oligolepis has also been reported by Hyodo & Tsukahara (1977) from the northern Arabian Sea at about 20– 22°N 60– 70°E, by Jones & Kumaran (1965) from the Laccadive Archipelago, by Rao et al. (2000) from Mayabunder (Andaman Islands) and by Chen (1993) from Taiwan. Imai (1958a, 1959) described (as C. starksi ) juveniles presumably of C. o. oligolepis from the waters of southern Japan (Makurazaki). Additional occurrences are listed in synonymy and bibliography.

Günther (1910) reported C. oligolepis from Tahiti. However, it is probably a misidentification because there is no evidence of any subspecies of C. oligolepis from Tahiti.

Figures

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.