Ch. spilopterus

Full morphological study. IORAS 03430 (1, 109 mm SL), 6°12’N 123°08’E, 5.04.1961 . IORAS 03431 (1, 132.5 mm SL), 11°29’S 122°30’E, 21.10.1959 . CAS 81227 (1, 205 mm SL), 9°44’-45’N, 118°47’-48’E, 9.05.1961 . CAS 82037 (1, 160 mm SL), 10°25’N 117°46’E, 2.02.1949 . CAS 82111 (2, 132– 169.5 mm SL), 4°45’N 119°53’E, 30.07.1948 . FRSKU 100393 (1, 215 mm SL), S285, Tong King Bay . FRSKU 100394 (1, 221 mm SL), S753, Tong King Bay . DASF ( Konedobu, PNG). No . NFO1963 * (1, 225 mm SL), Kitava I., Trobriands, 9.11.1969 .

Partial morphological study. USNM 308500 * (2, 217– 220 mm SL), Funafuti, Tuvalu, August 1989 . USNM Prereg No. 5411* (1, 150 mm SL), ~ Indonesia - Oceania . SIO 61-60 * (1, 163 mm SL), 7°10’S 127°22’E . SIO 61- 720 * (1, 207 mm SL), 11°11’S 122°29’E, 29.03.1961 . SIO 61-721 * (1, 151 mm SL), 10°33’S 124°17’E, 12.04.1961 . SIO 61-722 * (1, 216 mm SL), 9°19’S 126°29’E, 16.04.1961 . CAS 81958* (2, 209– 220 mm SL), 4°38’N 119°30’E, 11.12.1947 . CAS 81959* (1, 214 mm SL), 8°32’N 118°20’E, 6.12.1947 . CAS 81962* (1, 183 mm SL), 6°57’N 121°02’E, 4- 5.12.1947 . CAS 82021* (1, 213 mm SL), 2°00’N 120°00’E, 2.08.1948 . CAS 82133* (1, 227 mm SL), 13°22’N 124°36’E, 2.07.1948 .

Holotype (Fig. 4). ZMMU P-24078, R / V "Vityaz", Cruise 50, Sta. 6480, Sample 500, 7°32’N, 116°37’E, captured near the surface by a dip net on 10.06.1971 at 20.30-22.50. Juvenile, 108 mm SL. D 13, A 9, P I 12, Spred 30, Str 7½, Sp.br 23 (7 + 16), Vert 41 (26? + 15?). Measurements (in % SL): aA 77.0, aD 70.5, aV 57.5, cV 34.3, pV 39.0, c 24.1, po 10.2, o 8.3, ao 4.7, io 8.3, Hc 16.7, H 17.5, h 7.3, Dc 28.0, lP 68.5, lP 1 41.4, lV 32.3, lD 19.4, lA 11.3, HD -, HA 7.2, p 14.7. Body without bands, lower surface of head pale (with only few dots near lower jaw symphysis); dark specks below eye and on gill cover. Pectoral fins pale brown with pale “mirror” to the 4th ray and with 3–4 rows of dark spots. Tip of pectoral fin almost reaches middle of caudal peduncle. Pelvic fins pale, with brown pigmentation distally between 3rd – 4th rays (against which there is a darker spot) and sparse gray pigmentation proximally between 2nd – 3rd rays. Tip of pelvic fin protrudes slightly beyond end of anal-fin base. Dorsal fin gray, darker near ends of rays (except posterior rays); tip of last dorsal-fin ray reaches middle of caudal peduncle; penultimate ray shorter than ultimate. Anal fin transparent, first anal-fin ray below 6th dorsal-fin ray. Caudal fin pale brown, tip of upper lobe slightly darker. Lower jaw longer than upper, teeth large, conical and recurved, and arranged in two rows near the symphysis. Anterior of palatines with several teeth.

Paratypes. IORAS 03407, 6 specimens 36.5, 48, 55.5, 60, 62 and 67.5 mm SL, 1°58’N 105°53’E, 13.06.1971 . SIO 65-42, 163 mm SL, 6°51’N 103°22’E, 9.11.1964 . CAS 81226, 157 mm SL, 7°05’N 102°57’E, 11.08.1960, CAS 81188, 2 specimens 122 and 132 mm SL, 7°27’N 107°23’E, 2.10.1960 .

Description. Meristic and morphometric characters are given in Tables 1–7. D 11–14, A 9–11, P I 11–14 (usually I 12), Spred 25–31 (usually 27–30), Str 7–8½ (usually 7–7½), Sp.br 20–25 (4–7 + 15–18), Vert 41–42 (26– 27 + 14–16). Palatine teeth, if present (absent in approximately 50% of specimens), few in number: usually 1–2 teeth on anterior of palatines, and only a few specimens with up to 7 teeth; palatine teeth absent in all juveniles under 65 mm SL. Snout oblong, upper jaw slightly pointed at tip (Fig. 5g). Lower jaw longer than upper; teeth usually unicuspid. Only occasional specimens with several teeth with additional cusps (one specimen with single tricuspid tooth). Jaw teeth in adults numerous, large (usually easily visible to the naked eye), conical, often recurved. Teeth arranged in 2 or 3 rows. In juveniles, jaw teeth are not as numerous, in 1 or 2 rows, relatively small, conical or needle-shaped. Starting at about 60–70 mm SL teeth begin to recurve.

Body elongate. Greatest body depth does not change with growth, 5.1–6.2 in SL. Body width 1.07–1.34 and caudal peduncle depth 2.07–2.76 in greatest body depth. Greatest head depth and head length almost do not change with growth, 5.6–6.2 and 3.7–4.2 in SL, respectively. Head length 0.92–1.22 in dorso-caudal distance. Eyes are large, eye diameter of juveniles 35–100 mm SL decreases strongly as the fish grows, in fish about 100–130 mm SL, diameter almost does not change with growth, and in largest fish (135–165 mm SL) even increases again. In juveniles 35–100 mm SL eye diameter 8.8–12.3 in SL, 2.2–3.1 in head length and 1.1–1.3 in interorbital space, in fish 125–165 mm SL, eye diameter 10.8–13.3 in SL, 2.8–3.3 in c, and 1.0–1.2 times in io.

Pectoral fins relatively long, their length decreasing with growth: in juveniles (35–100 mm SL) 1.4–1.6 times, in large fish (125–165 mm SL) 1.5–1.7 in SL. Tip of pectoral fin reaches from posterior of dorsal-fin base to middle of caudal peduncle. First pectoral-fin ray unbranched, its length virtually does not change with growth: 2.2– 2.8 in SL and 1.5–1.85 in lP. Pelvic fin base slightly closer to posterior edge of head than to beginning of lower caudal-fin lobe (cV / pV = 0.74–0.96); pelvic fins shift slightly posteriorly as fish grows (Fig. 2). Pelvic fin length decreasing quite strongly from juveniles to adults: in juveniles (35–100 mm SL) 2.4–3.1 times in SL and 1.6–2.0 times in lP; in large fish (125–165 mm SL) 3.35–3.8 and 2.2–2.55 times, respectively. Tip of pelvic fin of small juveniles 35–65 (occasionally up to 73) mm SL reaches, or almost reaches, beginning of lower caudal-fin lobe (in few fishes it protrudes slightly beyond); in juveniles 65–115 mm SL it reaches from end of anal-fin base to middle of caudal peduncle or a little further; in fish 120 mm SL or more it reaches from middle to end of anal-fin base (and only in one fish it slightly protruded beyond end of anal-fin base).

Anal-fin origin far behind dorsal-fin origin (1st anal-fin ray under 4 th –7 th dorsal-fin ray, usually under 5th or 6th). Dorsal fin with 1–4 rays more than anal fin. Height of dorsal and anal fins decreases markedly as fish grow. In juveniles (35–100 mm SL) HD 5.3–8.8, HA 8.7–14.3 in SL; in large fish (125–165 mm SL) HD 8.5–11.6, HA 13.1–16.3 in SL. Longest ray of dorsal and anal fins is 2nd or, occasionally, 3rd. Tip of last dorsal-fin ray reaches middle of caudal peduncle - the beginning of lower caudal fin lobe. Middle and posterior rays of dorsal fin not elongate (in juveniles 25–65 mm SL penultimate rays are slightly elongated and extend slightly beyond tip of last ray).

Pigmentation. Five to six bands on the lower part of body in juveniles 24–55 mm SL (Figs. 5 a–d): near pectoral-fin base (often barely noticeable), between pectoral and pelvic fins, near pelvic-fin base, above anus, above middle of anal fin and near caudal-fin base. In addition, some fish with aggregation of melanophores on gill cover. In fish 55–75 mm SL the bands begin to disappear and, usually, there are no more than 3–4 poorly discernible bands on the underside of body. Fish larger than 80 mm SL have no body bands (Fig. 5e).

The head of juveniles 35–90 mm SL dark, its lower surface is densely covered with dots, only chin is paler, with only a few melanophores. Later the pigment on the lower surface of head begins to disappear, and in fish 110 mm SL lower surface of head pale with only few dots near the lower jaw symphysis. Adults usually with a few small specks on gill cover and under eye.

Pectoral fins (Fig. 6) of juveniles 25–55 mm SL pale with dark oval area on distal part of fin near lower edge and with slightly pigmented area on base near upper rays. Some fish also with small dark spot near tip of first ray. In fish 55–90 mm SL, these pigmented areas expand and merge near the upper edge of fin (Fig. 6b); fin becomes dark brown (proximally may be paler than distally) with pale transverse band (“mirror”) reaching up to 1st–3rd ray. A few darker spots usually visible on fin. In fish 115 mm SL and more, fin brown or dark brown with narrow posterior pale edge, a “mirror”, reaching to 3rd–5th (8th) ray (sometimes the “mirror” is feint or entirely inconspicuous), and dark spots, varying greatly in size and number. Spots are usually numerous, but some fish do not have more than 3–5 spots; only in one of 43 fish studied spots completely absent (Fig. 6f).

Pelvic fins of juveniles 25–70 mm SL (Fig. 7a) pale with large dark spot distally and few dots near bases of outer rays. In larger fish, 65–115 mm SL (Fig. 7b, c), dots on fin base disappear, and only the spot distally between 2nd–5th rays remains (sometimes with 1–2 darker, oval spots superimposed on this spot). In fish 115–135 mm SL (Fig. 7d), the distal spot disappears, although a vestige remains as dots between rays in some fish. In larger fish, pelvic fins transparent.

The dorsal fin of juveniles 25–85 mm SL pale with dark spot on upper (or posterior-upper) part or with dark upper margin. A few dots scattered near bases of anterior rays and on central part of fin in some fishes. In fish 85– 120 mm SL, the spot begins to disappear, and only its remains near the tips of rays persist. In fish over 125 mm SL, spot disappears completely and fin becomes uniformly gray (in one specimen 145 mm SL ends of dorsal-fin rays were blackish).

The anal fin of juveniles 25–80 mm SL transparent with small spot (or aggregation of dots) posteriorly near ends of rays. In fish 80–115 mm SL the spot disappears, remaining in some fish as a few dots. In fish over 115 mm SL anal fin transparent or, more rarely, grayish.

Caudal fin of fish 25–55 mm SL (Figs. 5 a–d) pale, with pigmentation along lower lobe rays (except uppermost rays), sometimes forming 1–2 aggregations (bands). In fish 55–80 mm SL, pigmentation appears on base of upper lobe and, in some fish, also on its end. In fish 80–115 mm SL pigment extends onto fin and fin becomes pale brown with paler fork and, sometimes, with darker tip of the upper lobe. In fish over 115 mm SL fin becomes more or less uniformly pale brown.

Coloration in life. No data.

Maximum size. The maximum length of Ch. f. pseudospilopterus in our material was 163 mm SL (SIO 65-42, 6°51’N 103°22’E, 9.11.1964). Females and males do not differ in size.

Etymology. The name " pseudospilopterus " reflects the extreme similarity of adults of this subspecies with Cheilopogon spilopterus .

Common names. The name “spot-winged toothy flying fish” (Russian: “пятнoкpылый зyбacтый длиннoкpыл”) is proposed here.

TABLE 1. Dorsal, anal and pectoral fin ray counts for flying fishes Cheilopogon formosus, Ch. marisrubri, Ch. spilopterus, Ch. katherinae, Cheilopogon sp.1, Cheilopogon sp.2, Cypselurus bosha and C. olpar .

TABLE 2. Transverse scales counts and dorsal minus anal rays difference for flying fishes Cheilopogon formosus, Ch. marisrubri, Ch. spilopterus, Ch. katherinae, Cheilopogon sp.1, Cheilopogon sp.2, Cypselurus bosha and C. olpar .

SpredSpecies (subspecies), locality232425262728293031323334 Ch. formosus formosus--1--1------ Ch. formosus pseudospilopterus--11131813154--- Ch. formosus andamanicus--1-9131021--- Ch. formosus (species total) --31223223175--- Ch. marisrubri--38915102---- Ch. spilopterus----174342-- Ch. katherinae-------44112 Cheilopogon (Ptenichthys) sp.1 ----122511-- Cheilopogon (Ptenichthys) sp.2 ----1--1---- C. bosha (northern Vietnam and China) ---91213102---- C. bosha (Indonesia, Philippines, Thailand, Malaysia) -1110943----- C. bosha (Kenya and Madagascar) ---3212---- C. bosha (Red Sea) ---31221---- C. bosha (species total) -11252420173---- C. olpar (western and northern Australia) 113981111---- C. olpar (eastern Australia) 14431111---- C. olpar (species total) 2571291222----

TABLE 4. Gill raker counts for flying fishes Cheilopogon formosus, Ch. marisrubri, Ch. spilopterus, Ch. katherinae, Cheilopogon sp.1, Cheilopogon sp.2, Cypselurus bosha and C. olpar .

Species (subspecies) S L, mm n aA aD aV cV cV 1 Measurements, % S L pV c po o ao ao 1 io Ch. formosus 68.8 36.0-99.0 29 77.0 72.7-78.8 69.5 66.3-72.3 57.7 55.3-59.8 32.8 30.5-35.3 - 40.2 39.0-42.5 25.0 23.9-26.1 10.6 9.1-11.3 9.5 8.1-11.3 4.9 3.6-5.5 - 10.1 9.1-11.4 pseudospilopterusCh. formosusandamanicus Ch. formosus(both subspecies) Ch. marisrubri142.3 101.0-163.040 78.5 (n=28) 77.0-80.2 71.2 (n=28) 69.1-74.4 59.2 (n=28) 56.4-61.6 33.9 (n=24) 31.4-35.7 34.4 (n=16) 32.7-36.838.7 36.6-41.325.6 24.1-26.710.5 (n=29) 9.8-11.28.5 (n=32) 7.5-9.25.7(n=22) 4.7-6.27.1 (n=4) 6.6-7.89.3 (n=23) 8.3-10.057.7 41.0-76.012 77.0 75.2-78.8 70.3 67.6-71.7 58.4 56.3-60.4 33.2 31.5-34.7 -39.3 37.9-41.625.4 (n=9) 24.3-26.010.7 9.7-12.39.6 8.9-10.64.4(n=9) 4.0-4.7-9.9 9.3-10.5147.7 101.0-179.022 79.0 (n=9) 77.7-80.0 71.6 (n=9) 69.6-73.7 59.9 (n=9) 58.0-61.3 34.6 (n=9) 32.2-35.8 36.2 (n=13) 34.6-38.137.8 35.7-41.525.1 24.2-26.310.7 (n=9) 10.2-11.28.3 (n=16) 7.1-9.15.6 (n=9) 5.0-6.1-9.4 (n=9) 8.8-10.165.2 36.0-99.041 77.0 72.7-78.8 69.8 66.3-72.3 57.9 55.3-60.4 32.9 30.5-35.3 -40.0 37.9-42.525.1 23.9-26.110.7 9.1-12.39.5 8.1-11.34.8 3.6-5.5-10.0 9.1-11.4 144.4 101.0-179.0 62 78.6 (n=37) 71.3 (n=37) 59.4 (n=37) 34.1 (n=33) 35.2(n=29) 32.7-38.1 38.4 35.7-41.5 25.4 24.1-26.7 10.5 (n=38) 9.8-11.2 8.4 (n=48) 7.1-9.2 5.6 (n=31) 4.7-6.2 7.1 (n=4) 6.6-7.8 9.3 (n=32) 8.3-10.166.733.0-99.029-42.1 (n=20)40.4-44.124.8 (n=19)23.7-26.010.2 (n=19)9.0-11.39.7 (n=20)8.4-11.84.6 (n=12)4.1-5.4-10.6 (n=20)9.1-11.8140.1102.5-163.0 30 33.4 (n=2)32.9-33.940.4 (n=29)38.7-42.125.4 (n=29)24.0-27.210.9 (n=28)10.2-11.78.07.6-8.75.7 (n=26)5.1-6.3 - 9.2 (n=28)8.3-10.0150.0 109.0-183.0932.1 (n=4) 31.1-32.842.0 40.2-43.224.8 22.0-26.911.1 (n=5) 10.4-11.87.9 (n=7) 7.2-8.55.3 (n=5) 3.8-6.2-10.0 (n=5) 9.2-11.1 Ch. spilopterus 216.1 13 205.0-227.0 31.6 (n=10) 41.2 25.8 10.8 (n=4) 7.9 (n=8) 5.9 (n=3) 7.5 9.3 (n=7) 30.3-33.7 39.3-42.3 24.1-27.2 10.6-11.0 7.2-8.9 5.4-6.3 n=1 8.3-9.7 Ch. katherinae173.7 139.0-212.01235.4 (n=6) 33.2-36.741.5 (n=11) 40.3-43.024.3 23.5-25.110.6 (n=8) 10.3-11.27.5 (n=11) 7.0-8.25.0 (n=5) 4.3-5.87.2 (n=5) 6.6-7.68.2 (n=5) 8.0-8.5 Cheilopogon sp.1 58.6 33.0-103.014 72.1-76.6 66.0-71.0 49.3-54.6 26.6-33.3 32.2 (n=2) 31.5-33.042.3 40.9-44.823.5 (n=13) 21.8-24.79.5 (n=12) 8.4-10.59.4 8.0-10.64.4 (n=11) 4.0-4.8-9.3 (n=10) 8.1-11.2 Cheilopogon sp.2 51.4 31.0-72.55 75.9 75.5-76.7 67.8 64.8-70.3 53.4 51.9-54.5 29.4 28.1-30.3 -43.2 (n=4) 43.0-43.523.9 23.0-25.29.9 9.3-10.69.8 8.8-10.44.3 3.5-4.7-9.8 9.1-10.6

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Species (subspecies) S L, mm io 1 Hc H h Dc lP Measurements, % S L lP 1 lV lD lA HD HA p lcir Ch. formosus 68.8 36.0-99.0 - 17.1 16.1-17.9 17.3 16.1-18.0 7.3 6.4-8.1 27.4 25.6-29.3 67.7 64.2-72.2 40.5 37.6-44.6 37.3 32.8-41.9 19.6 17.9-21.2 12.5 10.8-14.4 14.3 (n=17) 11.3-18.8 9.1 (n=19) 7.0-11.5 14.4 13.3-15.0 - pseudospilopterusCh. formosusandamanicus Ch. formosus(both subspecies)

Ch. marisrubri Comparative remarks. Adults of Ch. f. pseudospilopterus are similar to representatives of Ch. spilopterus of the same size, and these two taxa are very difficult to distinguish. In most of the collections studied, the adults of Ch. f. pseudospilopterus were identified as Ch. spilopterus (including those determined by the second author of this work, see Parin, 1996:305). These species differ somewhat in pigmentation of the pectoral fins ( Ch. spilopterus usually has no “mirror”, and frequently has a large darker spot or stripe near the posterior end of the fin, compare Figs. 6 and 8), in total number of rays in dorsal, anal and pectoral fins (in Ch. spilopterus D + A + P = 37–40, in Ch. f. pseudospilopterus, 33–38, usually 36 or less), and in a number of morphometric characters (Tables 6, 7): most strongly in cV / pV, aA, lA; there are also differences in aD, h, aV, cV, pV, H, Dc, Dc / c, lD, HD (Figs. 2 a–e). These species also differ in the number of vertebrae: according to Parin (1961a, 1961b), Ch. spilopterus has 42–44 (26–28 precaudal + 13–16 caudal) vertebrae. Juveniles of these two species are easily distinguished because Ch. spilopterus has paired chin barbels, to about 100 mm SL (juveniles of Ch. formosus pseudospilopterus have no barbels), a greatly enlarged dorsal fin, and different pigmentation on body and fins (see Parin 1961a, fig. 10).

Ch. spilopterus 216.1 17.3 (n=3) 19.5 (n=7) 7.6 (n=6) 27.5 67.3 (n=6) 39.3 (n=5) 25.2 (n=6) 19.0 (n=4) 12.7 (n=5) 11.2 (n=5) 6.6 (n=3) 14.6 (n=6)

- - 205.0-227.0 16.6-18.1 18.6-20.3 7.2-8.2 26.5-28.5 63.5-69.5 36.6-44.7 23.1-27.4 18.2-19.6 12.1-13.5 9.5-12.7 5.4-8.5 14.0-15.2

Note: Here and elsewhere the mean is given above the line, the range of variation – under the line.

TABLE 7. Some indexes of measurements for flying fishes Cheilopogon formosus, Ch. marisrubri, Ch. spilopterus, Ch. katherinae, Cheilopogon sp.1, Cheilopogon sp.2, Cypselurus

bosha and C. olpar .

Biology. The only available mature female of this species, 162 mm SL, was captured off southern Vietnam (5°59’N 105°23’E) in June 1960. The only mature male 158 mm SL was caught in Gulf of Thailand (7°37’N 103°38’E) in January 1961. One female (142 mm SL, captured 11.08.1960, 7°15’E 103°06’E) was close to maturity. Early juveniles, 23.5–28.5 mm SL, were captured in the South China Sea in March and June. Based on the above, the spawning season of Ch. f. pseudospilopterus is probably greatly extended and appears to last at least from January to February and from June to August.

Distribution. This subspecies is distributed (Fig. 3) in the southern shallow-waters of the South China Sea, from the Gulf of Thailand to northern Borneo and eastwards to the southern Philippines (ZMUC, uncat., 38 mm SL, 9°36’N 125°46’E, 30- 31.07.1951). Several fish were also caught south of Java (extreme south point of capture at 10°18’S 110°23’E - IORAS, uncat., 136 mm SL, 31.10– 2.11.1959). The distribution of this subspecies (as well as the other two subspecies of Ch. formosus) is neritic.