Cypselurus comatus ( Mitchill, 1815 )

Shakhovskoy, Ilia B. & Parin, Nikolay V., 2024, A review of the flying fish genus Cypselurus (Beloniformes: Exocoetidae). Part 3. Revision of the subgenus Cypselurus sensu stricto with descriptions of one new species and four new subspecies and reinstatement of Cypselurus crockeri Seale and Exocoetus socotranus Steindachner, Zootaxa 5473 (1), pp. 1-125 : 97-115

Cypselurus comatus ( Mitchill, 1815)

Synonymy and bibliography.

Exocoetus comatus Mitchill, 1815: 448 , pl. 5 fig. 1 (original description; New York). Valenciennes in Cuvier & Valenciennes 1847: 133–135 (description; South Carolina). Günther 1866: 286 (short description; southern Atlantic). Lütken 1876: 393, 399, 400, Tab. 10 fig. 1 (short description; southern Atlantic, 11°S 36°W).

Exocetus appendiculatus Wood, 1825: 283 (original description; locality unknown).

Cypsilurus appendiculatus . Swainson 1838: 299, fig.63 (placed in a new genus Cypsilurus ).

Exocetus comatus . DeKay 1842: 231, pl. 36 fig. 115 (short description; New York). Storer 1846: 440 (short description; New York).

Cypselurus comatus . Weinland 1858: 385 (listed). Jordan & Gilbert 1883: 381 (short description; Atlantic Ocean). Breder & Nichols 1934: 40, 42 (discussion on validity). Breder 1938: 38, 51–55, figs. 25–28 (description, early life history stages, distribution; western Atlantic). Imai 1955: 99–101, 103–104, fig. 5 (comparison with C. naresii ).? Erdman 1956: 324 (listed; Puerto Rico). Duarte-Bello 1959: 45 (listed; Cuba). Parin 1961b: 118, 129, 137, 140, 165, 180 (morphology, systematics). Staiger 1965: 679–685, figs. 1–2 [description; western Atlantic; in part: specimen from Gulf of Mexico is Cheilopogon melanurus (Valenciennes) ]. Parin 1967: 55 (distribution). Kotthaus 1969: 11 (morphometry; in part: only Atlantic specimens). Gibbs & Staiger 1970: 461 (distribution; western Atlantic, in part). Rass 1972: 8 (eggs; Cuba). Fahay 1975: 19 (in key). Guitart 1975: 263–264, fig. 198 (short description; Cuba). Gibbs 1978: EXOC CYP 1 (short description, figure, distribution; West Central Atlantic; in part). Kovalevskaya 1980: 224 (listed). Kovalevskaya 1982: 108–110, figs. 1–3 (early life history stages; western Atlantic). Kovalevskaya 1983b: 1–2 (early life history stages). Grudtsev 1987: 38, 44, 58, 79, 140, 199, 201, 203, 205, 207 (distribution; western Atlantic). Bruce & Bowman 1989: 16 (listed as host of Cymothoidae parasites). Lao 1989: 105 (listed; Barbados). Oxenford 1993: 36 (listed; Barbados). Collette et al. 1997: 27 (South Carolina). Monteiro et al. 1998: 398–399 (short description; North-eastern Brazil). Carvalho-Filho 1999: 78 (short description; Brazil). Parin 2003: 1126 (description, distribution, figure; West Central Atlantic). Quattrini et al. 2004: 161– 162, pl. 1 fig. E (listed; North Carolina). Cotten & Comyns 2006: 957, 978–979 (early life history stages). Schwartz 2006: 65, 67 (listed; off Cape Lookout). Fahay 2007: 808–809 (early life history stages). Hunte et al. 2007: 97 (listed; juveniles off Barbados). Casazza & Ross 2008: 352, 355 (listed; off North Carolina). Rudershausen et al. 2010: 1350, 1351, 1354, 1355 (listed as prey of large fishes; North Carolina). Parin et al. 2019: 186–191, figs. 5.12–5.13 (description, distribution; western Atlantic).

Exocoetus appendiculatus . Jordan & Meek 1885: 60 (stated to be identical with E. comatus ).

Exocoetus heterurus (non Rafinesque). Jordan & Meek 1885: 59–60 (in part). Jordan & Evermann 1896: 735–736 (description; in part; western Atlantic). Bean 1903: 331–332 (description; New York; in part: only juveniles with a chin barbel).

Cypsilurus bahiensis (non Ranzani).? Evermann & Marsh 1900: 104 (short description; Puerto Rico).

Cypselurus vitropinna Breder, 1927: 20–21 , fig. 11 (original description; West Indies). Breder & Nichols 1930: 7–9, fig. 5 (jaw teeth, comparison with C. lutkeni (= Ch. melanurus ); West Indies). Nichols & Breder 1930: 5–7 (in key; comparison with other Atlantic species). Parr 1930: 21, 23–27 (description, morphometry; Bahamas and Turks Is.). Breder & Nichols 1934: 40, 42, 44, fig. 1g (vertebral counts; synonymy).

Cypselurus antarei Beebe & Hollister, 1933: 83–84 , fig.15 (original description; 21°50’N 63°32’W). Breder & Nichols 1934: 40 (discussion on validity). Collette et al. 1992: 6 (description of holotype, synonymy with C. comatus ).

Cypsilurus comatus . Bruun 1933: 382 (not synonymous with C. heterurus ). Bruun 1935: 52–56, 96, text-figs. 9, 10, 28, pl. 2 fig. 3 (description, distribution, western Atlantic). Fowler 1944: 440 (Bahama Is.).

Cypselurus alienus Herre, 1935: 392–393 (original description; Cuba near Santiago, Caicos Passage). Herre 1936: 11, fig. 1 (listed; Cuba and Caicos Passage).

Probable misidentifications. Fowler (1959) described C. comatus from waters of Fiji, but it is a misidentification and he dealt with Cheilopogon cyanopterus (Valenciennes) . Sauskan (1973) reported C. comatus as most plentiful species of flying fishes in neritic waters of the eastern tropical Atlantic, but this author had probably dealt with Ch. melanurus . Pinheiro et al. (2020: 5, fig. 4e) reported a juvenile of C. comatus from Saint Peter and Saint Paul’s Archipelago, but it is a misidentification of a species of Cheilopogon , subgenus Ptenichthys Müller (probably also Ch. melanurus ).

Material examined. Study material is similar to Parin et al. 2019. Eighty six specimens 27.5–210 mm SL (of which three specimens were studied by R.H. Gibbs and 21 specimens by J.C. Staiger [compare with list of material in Staiger 1965]).

North-western Atlantic Ocean. Full morphological study. IORAS 04356 (1, 173 mm SL), 19°12’N 68°04’W, 10.02.1973 . IORAS 04357 (1, 180 mm SL), 19°38’N 68°09’W, 31.01.- 1.02.1973 . IORAS 04358 (1, 173 mm SL), 19°50’N 68°14’W, 3- 4.03.1973 . IORAS 04144 (1, 183.5 mm SL), road of Plymouth , Montserrat I., 14.03.1969 . IORAS 04360 (1, 160 mm SL), 16°24’N 62°43’W . IORAS 04361 (1, 183 mm SL), ~ 20°25’N 60°36’W, 11- 13.03.1964 . IORAS 04366 (1, 80 mm SL), 19°40’N 76°42’W, 10- 11.06.1982 . USNM 170321 *g (3, 183– 195 mm SL), 17°27’N 63°13’W, 13.04.1956 . ZMUC P34718 (1, 167 mm SL), between St. Thomas and St. Croix, 23.03.1922 .

Partial morphological study. AMNH 27113 (B.O.C. 167)* (paratype of C. vitropinna ) (1, 169 mm SL), West Indies , February–April 1925 . ANSP 67723 * (1, 108 mm SL), 25°N 80°W, 13.07.1935 . ANSP 150179 * (1, 197 mm SL), Bahamas: Hog Island, 1.08.1967 . ANSP 150180 * (1, 172 mm SL), Bahamas: Crooked Island Passage, 26.05.1962 . CAS 124419 ( SU 24419 )* (paratype of C. alienus ) (1, 165 mm SL), Bahama Is., Caicos Passage, 27.11.1928 . FMNH 17224 * (holotype of C. alienus , 158.5 mm SL), Cuba, near Santiago, 28.11.1928 . MCZ 41190 * (2, 181– 186 mm SL), Bahamas, June 1945 . UMML 13816 x (3, 183– 205 mm SL), Bahamas . UMML 16364 x (5, ~ 169–198 mm SL), Bahamas . UMML G-50s (2, 31–46.5 mm SL), 25°38’N 79°25’W . UMML G-70s (2, 34.5–55.5 mm SL), 25°23’N 79°41’W . UMML uncats (1, 27.5 mm SL), 27°40’N 79°18’W, 11.09.1954 . UMML uncats (1, 29.5 mm SL), 26°25’N 76°40’W, 24.01.1954 . UMML uncats (1, 38 mm SL), 27°00’N 79°18’W, 22.04.1953 . UMML uncats (1, 40.5 mm SL), 26°25’N 76°40’W, 13- 14.06.1954 . UMML uncats (1, 44 mm SL), 28°19’N 79°26’W, 24.04.1953 . UMML uncats (1, 60 mm SL), 20°43’N 73°33’W, 22.12.1961 . UMML uncats (1, 60 mm SL), 30°21’N 79°26’W, 27.06.1954 . UMML uncats (1, 96 mm SL), 22°35’N 78°08’W, 29.11.1951 . UMML uncats (4, 169.5– 179.5 mm SL), off Bimini, 6- 7.07.1961 . UMML uncats (1, 181 mm SL), 27°40’N 79°18’W, 23.06.1954 . UMML uncats (3, 182– 204.5 mm SL), Cab Cay— West End , 21.05- 3.06.1962 . USNM 53085 * (1, 180 mm SL), near Cat I., Bahamas, 8.07.1903 . USNM 126426 x (1, ~ 164 mm SL), Puerto Rico, 22.02.1899 . USNM 170921 * (holotype of C. antarei , 67 mm SL), N of Sombrero I., 30.07.1932 . USNM 178024 x (1, ~ 162 mm SL), Bahamas: 5 miles west of Crooked I., 25- 26.01.1936 . USNM 197842 *s (1, 151 mm SL), between Gd. Cayman I. and Swan I., 15.01.1950 . USNM 198024 * (1, length not measured), no data . USNM 200802 x (1, ~ 54 mm SL), Florida Straits, 15.05.1966 . USNM 251937 -F53s (1, 71 mm SL), 25°20’N 80°00’W, 30.08.1962 . USNM 268483 *s (1, 81 mm SL), 20°10’N 60°05’W, 10.12.1956 . USNM 294681g (1, 172 mm SL), 13°35’N 59°54’W, 6.02.1966 . USNM 294693 * (7, 149– 181 mm SL), 20°40’N 69°56’W, 29- 30.05.1965 . YPM 167 * (paratypes of C. vitropinna ) (5, 176– 210 mm SL), either Florida Keys, Bahamas, or western Caribbean Sea , 5.02.– 20.04.1925 . YPM 459 * (holotype of C. vitropinna , 177 mm SL), either Florida Keys, Bahamas, or western Caribbean Sea , 5.02.– 20.04.1925 .

South-western Atlantic Ocean. Full morphological study. IORAS 04393 (1, 181.5 mm SL), 20°32’S 34°49’W, 10.12.1981 . IORAS 04359 (1, 134 mm SL), 19°14’S 34°02’W, 6.01.1978 . IORAS 04362 (1, 195 mm SL), 20°41’– 21°00’S 34°46’– 34°48’W, 5.11.2004. IORAS 04363 (1, 193 mm SL), same data . IORAS 04364 (1, 201.5 mm SL), same data . IORAS 04365 (1, 196 mm SL), same data . IORAS 04367 (1, 99 mm SL), 17°46’S 35°30’W, 22.01.1964 . IORAS 04368 (1, 66 mm SL), 13°32’S 32°57’W, 4.01.1978 . USNM TABL 103136g (2, 199 – 208 mm SL), 14°29’S 38°51’W, 16.09.1966 .

Partial morphological study. IORAS 04370 (1, 61 mm SL), 22°51’S 35°08’W, 7.01.1978 . USNM 294694 * (1, 184.5 mm SL), 18°05’S 35°57’W, 5.10.1966 . ZMH ( ISH 238 /51)* (1, 188 mm SL), 18°49’S 38°32’W, 17.12.1951 .

Atlantic Ocean (precise locality unknown). Full morphological study. IORAS 04394 (1, 174 mm SL) . IORAS 04369 (1, 144 mm SL), 31.05.1965 .

Partial morphological study. ISH 238-1951s (1, ~ 190 mm SL).

Notes for Material examined section:

x These specimens were studied by the first author based on X-Rays only.

s These specimens were studied by J.C. Staiger (data from scientific archive of N. V. Parin).

g These specimens were studied by R.H. Gibbs (data from scientific archive of N. V. Parin).

Types. The holotype of C. comatus is unknown ( Fricke et al. 2023). The holotypes of C. alienus , C. vitropinna and C. antarei were studied by the second author and their characters are given below.

Holotype of C. alienus, FMNH 17224, south coast of Cuba, near Santiago , 28 November 1928. Length 158.5 mm SL, D 12, A 9, P I 14, Spred 26, Str 7½, Sp.br 22 (5 + 17), Vert 42 (28 + 14). Measurements (in % SL): cV 1 35.8, pV 37.9, c 26.1, po 11.1, o 9.1, io 1 8.7, H 18.9, h 7.3, Dc 27.2, lP 64.9+, lV 32.4, HD 11.6 , p 12.8. First anal-fin ray beneath 6th dorsal-fin ray. Longest dorsal-fin ray—2nd. Jaw teeth unicuspid, palatine teeth present.

Holotype of C. vitropinna, YPM ICH 000459, West Indies. Length 177 mm SL, D 12; A 9, P I 12, Spred 26, Str 8½, Sp.br 22 (5 + 17), Vert 42 (27 + 15). Measurements (in % SL): cV 1 35.0, pV 38.1, c 25.6, o 8.3, io 1 8.1, H 18.9, h 7.2, Dc 27.0, lP 67.4, lP1 44.6, lV 30.0, HD 11.2, p 13.1. First anal-fin ray beneath 5th dorsal-fin ray. Pectoral fins lightly pigmented to 7th ray.

Holotype of C. antarei, USNM 170921, 21°50’N 63°32’W, 30 July 1932. Length 67 mm SL, D 13, A 9, P I 14, Spred 24, Sp.br 23 (7 + 16), Vert 43 (28 + 15). Measurements (in % SL): cV 1 35.8, pV 35.0, c 25.8, o 9.1, io 1 9.8, H 17.9, Dc 27.7, lP 64.9, lV 40.4, HD 13.5, lcir 101.4. Pelvic fins dark. Longest dorsal-fin ray—2nd. See also original description of the holotype of C. antarei in Beebe & Hollister (1933).

Diagnosis. A moderately large species of Cypselurus with few dorsal-fin rays, predorsal scales and vertebrae. Body rather deep, head and eyes large, pelvic fins long, pectoral and dorsal fins rather short. Pelvic-fin base about midway between posterior edge of head and origin of caudal-fin lower lobe. Jaw teeth mainly conical and with additional cusps, palatine teeth present. Juveniles with a long, ribbon-like chin barbel and low dorsal fin; ventral side of body paler than dorsal one. In adults, pelvic and anal fins pale. D 11–13, Spred 22–30, Str 6–8½, Vert 41–44 (26–29 + 13–16).

Description. Meristic and morphometric characters are given in Tables 1–7 and 13–14. D 11–13 (usually 12)16, A 7–9 (usually 8–9), P I 12–15 (usually I 13–14), Spred 22–30 (usually 26–28), Str 6–8½ (usually 7–7½), Sp.br 19–25 (4–7 + 15–18), usually 21–24 (5–7 + 16–17), Vert 41–44 (26–29 + 13–16), usually 41–43 (27–28 + 14–16). Snout moderate, lower jaw usually shorter than upper jaw or of equal size, sometimes slightly longer; upper jaw not pointed anteriorly ( Fig. 45 ). Juveniles ≤ 150–160 mm SL17 with a long ribbon-like chin barbel ( Figs. 45a–d , 46 ), usually protruding beyond pectoral-fin base and, in some fish 55–75 mm SL, as long as body length ( Figs. 4f , 46a ; see also Kovalevskaya 1982: fig. 2). Chin barbel is wide proximally and narrowed distally (like in C. naresii

16 One specimen 149 mm SL ( USNM 294693 ) has 9 dorsal-fin rays, but it is undoubtedly a mutation .

17 The largest barbelled juvenile of C. comatus in our material was 151 mm SL (USNM 197842), but Bruun (1935) reported a juvenile 161.3 mm SL with a chin barbel as long as 67.5% SL.

Measurements, % SL

Region SL, mm n

aA aD aV cV cV1 pV c po o ao ao1 io 61.8 35.7 26.2 8.9

55.6 79.6 (n=6) 72.5 (n=6) 32.3 36.5 (n=3) 12.7 10.1

17 (n=7) (n=2) (n=16) (n=15) 5.1 (n=1) -

27.5–96.0 79.0–80.8 70.6–73.9 (n=1) 35.0–37.9 (n=1) (n=1)

North–western 60.6–63.4 35.6–35.8 24.8–27.7 7.6–9.9

Atlantic 79.0 71.8 60.2 34.9 35.3 36.8 25.4 11.3 8.2 6.8 9.2

175.8 5.4 (n=7)

40 (n=15) (n=15) (n=16) (n=7) (n=24) (n=31) (n=39) (n=9) (n=31) (n=4) (n=7)

108.0–210.0 4.8–6.2

77.9–80.1 70.3–73.3 58.8–62.0 33.1–36.5 32.5–38.4 35.3–38.3 23.7–27.2 11.0–11.8 7.1–9.4 6.6–7.3 8.8–9.8 9.9

82.5 77.7 72.1 60.1 34.7 36.7 25.9 11.9 8.7 4.7

2 - - 9.8–

66.0–99.0 77.5–77.9 71.5–72.8 59.7–60.6 34.7–34.8 35.4–38.0 25.7–26.1 11.9–12.0 8.6–8.8 4.6–4.8

South–western 10.0

Atlantic 60.6 35.6 37.5 11.2 7.9 9.1

188.0 78.0 (n=8) 71.1 (n=8) 36.7 24.9 7.7 5.4 (n=6)

10 (n=8) (n=6) (n=4) (n=8) (n=2) (n=6)

134.0–208.0 76.8–79.4 70.3–72.9 35.3–37.7 24.0–25.6 7.3–8.2 5.1–5.7

59.6–61.5 34.8–36.8 36.7–38.8 10.5–11.8 7.8–8.1 8.8–9.7

Measurements, % SL

Region SL, mm

io1 Hc H h Dc lP lP1 lV lD lA HD HA p lcir 9.3 7.8 26.5 63.3 39.1 18.2 10.2 12.9 10.2 14.7 54.5 55.6 19.0 19.4 (n=8) 39.1 (n=2)

(n=7) (n=6) (n=3) (n=16) (n=15) (n=6) (n=6) (n=12) (n=10) (n=7) (n=16) 27.5–96.0 (n=1) 17.9–20.7 37.6–40.6

North–western 8.4–9.8 7.5–8.1 25.1–27.7 59.1–68.2 35.4–41.3 17.5–19.3 9.0–11.3 11.8–15.1 7.8–12.9 13.3–15.3 25.8–101.4 Atlantic 8.4 17.6 19.0 6.9 26.4 64.3 40.6 29.2 17.6 9.9 10.8 6.9 13.8

175.8 54.4 (n=2) (n=23) (n=6) (n=33) (n=30) (n=31) (n=29) (n=18) (n=32) (n=15) (n=15) (n=22) (n=12) (n=33)

108.0–210.0 39.4–69.4 7.0–9.4 16.9–18.0 16.6–21.1 6.1–7.6 25.3–28.2 60.4–67.5 35.2–44.6 25.6–34.5 17.0–18.6 8.6–11.4 9.6–11.9 6.0–8.2 12.8–14.9

82.5 18.2 18.3 7.5 27.0 65.1 41.1 39.3 17.3 10.0 11.7 8.7 14.1 75.5

-

66.0–99.0 18.1–18.3 18.3–18.3 7.4–7.7 27.0–27.1 65.0–65.3 39.5–42.8 38.0–40.7 16.9–17.7 9.5–10.6 11.3–12.1 8.2–9.3 13.9–14.3 53.3– 97.7 South –western

Atlantic 188.0 7.7 18.1 19.5 (n=9) 6.7 26.5 66.1 40.0 (n=9) 30.0 17.9 10.2 10.0 7.0 14.1

(n=4) (n=5) (n=9) (n=9) (n=8) (n=8) (n=7) (n=8) (n=9) 42.6 (n=1) 134.0–208.0 17.5–21.6 6.2–7.2 26.0–27.6 37.5–41.7

7.5–8.2 17.9–18.4 65.2–67.9 27.6–35.9 17.3–18.8 9.1–11.3 9.1–10.9 6.2–7.7 12.7–14.7

C. hiraii (Pacific Ocean)

C. hiraii (Yellow and East China seas) Cypselurus hiraii (all regions)

C. o. opisthopus (Indian Ocean)

C. o. opisthopus ( Philippines)

C. o. opisthopus (New Guinea)

164.9 (n=62) 1.21(n=54) 1.03 (n=58) 2.15 (n=36) 1.65 (n=35) 1.40 (n=57) 3.13 (n=58) 102.0–186.0 1.09–1.38 0.90–1.16 1.68–2.37 1.52–1.89 1.12–1.58 2.79–3.44

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C. o. crockeri

Cypselurus opisthopus (species total)

C. n. naresii

C. n. ordinarius (western Pacific Ocean)

C. n. ordinarius (eastern Indian Ocean)

C. n. ordinarius (Bay of Bengal and Andaman Sea) 182.9 (n=42) 1.05 (n=37) 0.98 (n=41) 2.33 (n=29) 1.60 (n=33) 1.47 (n=36) 3.29 (n=38) 101.0–215.0 0.88–1.19 0.91–1.11 1.84–2.52 1.45–1.75 1.30–1.63 3.02–3.54

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(all regions)

169.4 (n=99) 1.06 (n=75) 0.99 (n=93) 2.21 (n=64) 1.60 (n=61) 1.48 (n=76) 3.29 (n=81) 101.0–240.0 0.88–1.22 0.89–1.11 1.82–2.52 1.44–1.75 1.30–1.68 2.94–3.61 62.8 (n=9) 0.99 (n=8) 1.01 1.62 (n=8) 1.53 (n=8) 1.51 (n=8) 3.20 (n=8) 50.0–86.0 0.91–1.06 1.00–1.05 1.55–1.77 1.46–1.60 1.41–1.75 2.95–3.38 190.0 (n=43) 1.03 (n=32) 1.00 2.24 (n=26) 1.61 (n=24) 1.49 (n=30) 3.29 (n=30) 101.0–258.0 0.95–1.12 0.90–1.13 1.78–2.61 1.50–1.74 1.30–1.68 2.96–3.57 62.8 (n=9) 0.99 (n=8) 1.01 1.62 (n=8) 1.53 (n=8) 1.51 (n=8) 3.20 (n=8) 50.0–86.0 0.91–1.06 1.00–1.05 1.55–1.77 1.46–1.60 1.41–1.75 2.95–3.38 194.8 (n=58) 1.02 (n=42) 1.01 2.24 (n=39) 1.60 (n=32) 1.53 (n=41) 3.35 (n=41) 101.0–258.0 0.91–1.12 0.90–1.13 1.78–2.61 1.47–1.74 1.30–1.92 2.96–3.80 143.1 (n=15) 0.97 (n=9) 1.02 (n=14) 2.40 (n=13) 1.57 (n=9) 1.49 (n=9) 3.27 109.0–182.0 0.91–1.07 0.94–1.10 2.07–2.62 1.45–1.66 1.33–1.60 3.02–3.63 62.8 (n=64) 1.04 (n=59) 1.01 (n=55) 1.62 (n=55) 1.57 (n=50) 1.37 (n=44) 2.96 (n=44) 33.0–96.0 0.81–1.20 0.90–1.13 1.35–1.87 1.46–1.71 1.08–1.75 2.53–3.38 154.6 (n=119) 1.04 (n=89) 1.00 (n=114) 2.17 (n=79) 1.60 (n=71) 1.49 (n=89) 3.28 (n=98) 101.0–200.0 0.88–1.22 0.89–1.13 1.78–2.62 1.44–1.75 1.30–1.77 2.94–3.72

C. n. albitaenia

(Indian Ocean)

C. n. albitaenia

(all regions)

C. n. socotranus b

Cypselurus naresii (species total)

221.3 (n=54) 1.05 (n=38) 1.01 2.41 (n=38) 1.60 (n=33) 1.52 (n=38) 3.35 (n=41) 201.0–258.0 0.97–1.13 0.91–1.11 2.10–2.61 1.45–1.74 1.30–1.92 2.92–3.80

C. persimilis 156.3 (n=7) 1.03 (n=4) 0.97 2.29 (n=5) 1.63 (n=5) 1.32 (n=4) 3.03 (Indian Ocean) 129.0–176.0 0.97–1.09 0.92–1.02 2.01–2.41 1.53–1.70 1.29–1.37 2.93–3.24

C. persimilis 153.2 (n=24) 1.13 (n=17) 0.94 2.38 (n=19) 1.59 (n=19) 1.40 (n=16) 3.17 (n=19) ( Philippines) 140.0–166.0 1.01–1.22 0.87–1.02 2.27–2.53 1.49–1.72 1.30–1.55 2.95–3.44

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76.0 (n=1) 0.97 1.15 1.87 1.67 1.19 2.70 C. a. folletti (eastern Pacific Ocean) C. a. angusticeps

(Polynesia) C. a. angusticeps (Hawaii)

C. a. angusticeps (western Pacific Ocean)

C. a. angusticeps (eastern Indian Ocean)

C. a. angusticeps (all regions)

Cypselurus angusticeps (species total)

C. comatus (north–western Atlantic)

C. comatus (south–western Atlantic)

C. comatus naresii ); it has a median keel along the full length (or nearly so) and two large triangular lobes basally. Jaw teeth small to medium-sized (not visible or barely visible to naked eyes), mainly conical and with additional cusps, in some fish tricuspid teeth may also be present (occasionally tricuspid teeth prevail); juveniles ≤ 80 mm SL with conical teeth only. Teeth arranged in 2–3 rows, in juveniles ≤ 80 mm SL in 1–2 rows. Palatine teeth present, sparse and diminutive (in fish> 190 mm SL absent in 4 of 7 specimens studied).

Body rather deep both in juveniles and adults, body depth 4.6–6.0 in SL. Body width 1.28–1.40 and caudal peduncle depth 2.28–3.12 in greatest body depth. Greatest head depth barely changing with growth, 5.25–5.9 in SL. Head length hardly changing with growth ( Fig. 47b ), 3.6–4.2 in SL and 0.90–1.11 in dorso-caudal distance. Eyes large, eye diameter slightly decreasing with growth ( Fig. 47c ), 10.1–14.1 in SL, 2.6–3.4 in head length, 1.3–1.45 in interorbital width and 1.2–1.6 in postorbital distance. Snout length barely increasing with growth.

Pectoral fins rather long, their length barely increasing with growth ( Fig. 4c ), 1.45–1.7 in SL. Tip of pectoral fin reaching from end of dorsal-fin base (or nearly so) to middle of caudal peduncle. First pectoral-fin ray unbranched, its length not changing with growth ( Fig. 4d ), 2.25–2.85 in SL and 1.46–1.79 in lP. Pelvic-fin base about midway ( Fig. 4a ) between posterior edge of head and origin of caudal-fin lower lobe (cV / pV = 0.88–1.03). Pelvic-fin length decreasing strongly with growth ( Fig. 47d ): in juveniles 25–100 mm SL pelvic fin 2.4–2.85 in SL and 1.50–1.82 in lP; in fish 135–210 mm SL, 2.8–3.9 in SL and 1.83–2.43 in lP. Tip of pelvic fin in juveniles 65–100 mm SL protruding slightly beyond origin of caudal-fin lower lobe. In fish ≥ 135 mm SL pelvic-fin tip reaches from end of anal-fin base (or nearly so) to middle of caudal peduncle.

Anal-fin origin well behind dorsal-fin origin (1st anal-fin ray under 5th–7th dorsal-fin ray). Dorsal fin with 2–5 (usually 3–4) rays more than anal fin. Height of dorsal ( Figs. 4e , 47e ) and anal fins decreasing with growth: in juveniles 25–100 mm SL, HD 6.6–8.85 in SL and HA 7.75–12.8 in SL; in fish 135–210 mm SL, HD 8.4–11.0 in SL and HA 12.2–16.65 in SL. Longest dorsal- and anal-fin ray—2nd (rarely 3rd). In fish> 65 mm SL tip of last dorsal-fin ray reaching from middle of caudal peduncle to origin of caudal-fin upper lobe; middle and posterior rays of dorsal fin not elongated, penultimate rays not extending beyond tip of last ray ( Fig. 45 ).

Pigmentation. Body of fish> 65 mm SL with typical “pelagic” pigmentation, body bands always absent ( Fig. 45 ). Ventral surface of head in juveniles 65–100 mm SL pale with pigmentation only on chin, in fish> 130 mm SL entirely pale. Dark stripe through the eye in juveniles absent, but snout usually with darker pigmentation. A few small dark specks may be present on gill cover and/or under eye of adults.

Chin barbel ( Figs. 45a–d , 46 ) pale brown to brown with a pale median keel, in some fish barbel is paler distally. Opposite (ventral) side of barbel pigmented similarly, but usually more intensely, with at most a trace of pale line along the keel (if any).

Pectoral fins in juveniles and adults ( Fig. 48a–c ) brown to pale-grayish (often nearly transparent in adults) to 7th–9th ray, usually with a pale tip; pale posterior edging absent or very narrow. Pectoral-fin pigmentation usually darker distally and extending one-two rays lower than proximally, occasionally a small “mirror” present. Pelvic fins of juveniles 65–150 mm SL pale brown to brown, much paler proximally and between 5th–6th rays ( Fig. 48d–f ); in some fish a pale edging between tips of 2nd and 3rd rays present. In fish ≥ 160 mm SL pelvic fins pale ( Fig. 48g ), occasionally with a few dots distally.

Dorsal fin of juveniles 65–100 mm SL pale with two weak aggregations of melanophores anteriorly near fin base and along the upper margin. In fish ≥ 135 mm SL dorsal fin gray to pale-brownish. Anal fin of juveniles 65–145 mm SL pale with aggregation of brown dots in the posterior part distally. In fish ≥ 160 mm SL anal fin unpigmented.

Caudal fin of juveniles 65–100 mm SL ( Fig. 45a–c ) pale with pale brown base and weak pale brown pigmentation along lower-lobe rays (in a juvenile 80 mm SL a faint dark band present on lower lobe distally). In fish> 130 mm SL caudal fin ( Fig. 45d–e ) pale brownish to brown, sometimes with slightly darker fin base and tip of upper lobe.

Coloration in life. No data.

Maximum size. The maximum length of C. comatus in the material examined was 210 mm SL (YPM 167 paratype of C. vitropinna , not dissected for sex determination). The largest female was 208 mm SL (USNM TABL 103136), the largest male— 195 mm SL (IORAS 04362).

Intraspecific variation. Fish from north-western and south-western Atlantic slightly differ in number of predorsal scales and vertebrae (see Tables 3, 6), as well as in preanal distance, head length, eye diameter, pelvic-fin length and dorsal-fin height ( Fig. 47 ). These differences indicate that populations from these regions partially isolated (by waters of Amazon River outflow?), but they do not merit a subspecies status.

Comparative remarks. Cypselurus comatus differs from C. hiraii in fewer vertebrae, predorsal and transverse scales and gill rakers (see Tables 1–6), longer head (in juveniles <150 mm SL head length 24.8–27.7 vs. 21.7–24.7% SL; in fish> 150 mm SL, 23.7–27.2 vs. 21.2–23.3% SL), larger eye (in fish> 115 mm SL eye diameter 7.1–9.4 vs. 6.0–7.3% SL), deeper body and head (H 16.6–21.6, usually> 18.0 vs. 13.9–19.4, usually <18.0% SL; Hc 16.9–19.0 vs. 13.9–16.8% SL), and some other characters (see Tables 7, 14); juveniles also in chin barbel morphology and pigmentation ( Figs. 2 , 4f , 46 ) and pigmentation of body and caudal fin (see descriptions above).

Cypselurus comatus differs from C. opisthopus in fewer predorsal scales and precaudal vertebrae ( Tables 3, 6), more anterior position of pelvic-fin base (index cV/pV 0.88–1.03 vs. 1.04–1.46, see Fig. 4a ), and some other characters (see Tables 7, 14); juveniles also in larger head, longer pectoral fins and lower dorsal fin ( Fig. 4c–f ), as well as morphology and pigmentation of chin barbel and pigmentation of body and fins; adults in pigmentation of head and pectoral fins and jaw teeth morphology (see descriptions above).

Cypselurus comatus differs from C. nossibe in slightly fewer pectoral-fin rays (13–16, usually ≤ 15 vs. 15–17, usually 16) and transverse scales (6–8½, usually ≤ 7½ vs. 8–9), smaller body width (12.5–15.3, usually <15.1 vs. 14.8–19.0, usually> 15.1% SL), shorter dorsal fin (16.9–19.3, usually <18.5 vs. 18.0–21.2, usually> 18.5% SL); adults also differ in jaw and palatine teeth morphology (jaw teeth mainly conical and with additional cusps, palatine teeth sparse or absent vs. jaw teeth mainly tricuspid, palatine teeth numerous) and pelvic-fin pigmentation (pale vs. pigmented distally), and juveniles in chin barbel morphology (long with a median keel vs. diminutive without a keel), dorsal-fin morphology (middle and posterior rays elongated in C. nossibe ), longer snout, smaller interorbital distance, slimmer caudal peduncle, as well as in body and pectoral-fin pigmentation (see description of C. nossibe in Shakhovskoy & Parin 2022).

Cypselurus comatus is most similar to C. naresii (especially to C. n. naresii ). It differs from that species in fewer predorsal scales ( Table 3), more anterior position of pelvic-fin base (index cV/pV 0.88–1.03, usually <1.00 vs. 0.88– 1.22, usually> 1.00 (except C. n. septentrionalis), see Fig. 4a ), larger dorso-caudal distance (25.1–28.2, usually> 26.0 vs. 22.0–27.5, usually <26.0% SL, Fig. 4b ); juveniles also have a longer head and postorbital distance (in fish 25–135 mm SL, c 24.8–27.7, usually> 25.3 vs. 22.8–26.8, usually <25.3% SL; in fish 65–135 mm SL, po 11.8– 12.7 vs. 10.6–12.4, usually <11.8% SL), deeper body and head (in fish 30–100 mm SL, H 17.9–20.7 vs. 14.6–19.9, usually <17.9% SL; in fish 65–145 mm SL, HC 17.9–19.0 vs. 14.6–18.1% SL) and differing pigmentation of body, head and caudal fin (see descriptions above); adults also differ in pigmentation of pelvic (pale vs. dark distally), pectoral (mainly pale brown to grayish, sometimes nearly transparent vs. mainly brown to dark brown) and anal (unpigmented vs. usually with dark dots [except C. n. socotranus ]) fins.

Cypselurus comatus differs from C. persimilis in having more dorsal-fin rays and fewer predorsal scales ( Tables 1, 3), a more anterior position of pelvic-fin base (index cV/pV 0.88–1.03, usually <1.00 vs. 0.97–1.24, usually> 1.00, see Fig. 4a ), a longer dorsal fin originating more anteriorly (aD 70.3–73.9, usually <73.0 vs. 72.3–75.9, usually> 73.0% SL; lD 16.9–19.3 vs. 14.3–17.7, usually <16.9% SL; Dc 25.1–28.2 vs. 21.8–25.5, usually <25.0% SL) and other characters ( Tables 7, 14), as well as in pigmentation of the head (dark specks sparse or absent vs. usually numerous) and pectoral fins (mainly pale brown to grayish, sometimes nearly transparent vs. mainly brown to dark brown).

Cypselurus comatus differs from C. angusticeps in having fewer dorsal-fin rays and more vertebrae ( Tables 1, 6), shorter dorsal fin originating more posteriorly (aD 70.3–73.9, usually> 71.0 vs. 67.2–72.3, usually <71.0% SL; lD 16.9–19.3, usually <19.0 vs. 18.1–24.1, usually> 19.0% SL; Dc 25.1–28.2, usually <27.5 vs. 25.5–31.3, usually> 27.5% SL, see Fig. 4b ), shorter pectoral fins ( Fig. 4c ); juveniles also differ in chin barbel morphology and pigmentation ( Figs. 4f , 39 , 46 ), a more posterior position of pelvic-fin base, shorter anal fin, narrower body and other characters (see Table 7).

Cypselurus comatus differs from all species of the subgenus Poecilocypselurus in the presence of a chin barbel in juveniles, more posterior position of pelvic-fin base (index cV/pV 0.88–1.03, usually> 0.90 vs. 0.48–1.05, usually <0.90 (except C. neglectus shcherbachevi and C. callopterus ), and jaw teeth morphology (mainly conical and with additional cusps vs. mainly tricuspid [except C. bosha and C. olpar ]).

Adults of C. comatus may easily be confused with Atlantic species Cheilopogon melanurus . Cheilopogon melanurus has more vertebrae, longer pelvic fins that are more anteriorly placed, and lacks palatine teeth (see Parin & Belyanina 2000).

Biology. Females and males are mature starting at 188 and 167 mm SL, respectively. Mature and close to mature adults were captured in the south-western Atlantic in November (IORAS 04363–04365) and December [ZMH (ISH 238/51)], and in the north-western Atlantic in March (ZMUC P34718). Bruun (1935) reported mature adults of C. comatus from the north-western Atlantic for January and February, and Breder (1938) for January–April. In the north-western Atlantic, two early juveniles 12.5 and 13.2 mm were captured in March (IORAS uncat., 19°50’N 68°14’W) and small juveniles ~ 30 mm SL in September and January (UMML uncat.). Thus, the spawning season is protracted at least from November to April and perhaps extending later into summer, with spawning in the south-western Atlantic probably starting earlier than in the north-western Atlantic.

Distribution. The distribution of C. comatus is presented in Figure 49 . This neritic species is found in two isolated areas: in the north-western Atlantic from New York ( Mitchill 1815) to southern Lesser Antilles (USNM 294681, 13°35’N 59°54’W), and in the south-western Atlantic from north-eastern Brazil ( Kovalevskaya 1982) to Rio de Janeiro (BMNH 79.5.14.674). Cypselurus comatus is absent in waters influenced by Amazon River outflow (though Grudtsev (1987) reported one occurrence in this area). It is probable that C. comatus , like its Indo-Pacific consubgeners, may avoid areas with wide continental shelves between northern Brazil and Trinidad and Tobago. Vega-Cendejas & de Santillana (2019: 5) listed C. comatus from Gulf of Mexico (Yucatan shelf), however did not provide any data in support of species identification. Therefore, the occurrence of the species in the Gulf of Mexico needs confirmation, especially because all previous reports of C. comatus in the region were based on a misidentification (see Parin et al. 2019: 190). According to Grudtsev (1987), this species was found in the western Atlantic at temperatures 23.4–28.2°C (optimum 25–27°C).

Concluding remarks on the flying fish genus Cypselurus

Parin (1961) devided the genus Cypselurus into two genera, Cypselurus and Cheilopogon , but his classification was not accepted by many researchers (e.g. Gibbs & Staiger 1970, Dasilao & Sasaki 1998), who reasonably argued that morphological distinctions between these two genera are not obvious. Chen (1987) based on morphology of early life history stages concluded that the subgenus Poecilocypselurus is most closely related to the genus Prognichthys Breder , instead of to Cypselurus s.str. However, monophyly of the genus Cypselurus is supported by data of molecular systematics ( Lewallen et al. 2011, Gordeeva & Shakhovskoy 2017).

After morphological study of the genus Cypselurus , we conclude that the only diagnostic character in adults is the morphology of jaws: in representatives of Cypselurus s. latu. ( Fig. 50a–g) the lower jaw is usually shorter or equal to the upper one and the upper jaw is usually not pointed at the anterior tip, while in representatives of Cheilopogon ( Fig. 50h–n) the lower jaw is usually longer than upper one and the upper jaw is usually slightly pointed at the tip (thus jaws form a sort of “beak”). However, the morphology of jaws is subject to variation both within genera ( Fig. 50) and within species. Despite the paucity of reliable diagnostic characters, we regard the genus Cypselurus as a valid monophyletic taxon. This assessment is made with the understanding that the genus Cheilopogon needs taxonomic revision in light of evidence suggesting that it is paraphyletic ( Lewallen 2011; Lee 2016; Gordeeva & Shakhovskoy 2017). As to differences between two subgenera of Cypselurus s.latu ( Cypselurus s.str. and Poecilocypselurus ), our study showed that position of pelvic-fin base is not a good character for distinguishing the two subgenera because of overlap, and the only reliable diagnostic character is presence or absence of a chin barbel in juveniles. Therefore, further quest for diagnostic characters is needed.

As a result of our three part review, the genus Cypselurus includes two subgenera, seventeen species and eighteen subspecies. The list of valid species and subspecies of the subgenus Poecilocypselurus was provided in the second part of the review ( Shakhovskoy & Parin 2022: 99). Species and subspecies of the subgenus Cypselurus s.str. that we regard as valid are listed below. Although some subspecies (such as Cypselurus naresii septentrionalis and C. n. socotranus ) might merit full species rank, such a decision should not be done without solid supporting evidence based on representative material including both adults and juveniles.

......Figure legend found on the next page

Subgenus Cypselurus s. str.

Cypselurus nossibe

Cypselurus hiraii

Cypselurus opisthopus

C. opisthopus opisthopus

C. opisthopus crockeri

Cypselurus persimilis

Cypselurus naresii

C. naresii naresii

C. naresii septentrionalis

C. naresii ordinarius

C. naresii albitaenia

C. naresii socotranus

Cypselurus comatus

Cypselurus angusticeps

C. angusticeps angusticeps

C. angusticeps folletti

The following provisional key is provided for identification of species and subspecies of Cypselurus s.str. We are aware of imperfections in the key. For example, the subspecies in couplet 3 may be distinguished only by chin barbel in juveniles, adults cannot be confidently identified at the present state of knowledge. We recommend users verify their identification with the full-length description provided in the species/subspecies accounts.

1. Vertebrae 45–47, usually 46–47; gill rakers 23–32, usually ≥ 26. Juveniles with a short, pear-shaped or cordiform, chin barbel not protruding beyond middle of eye..................................... Cypselurus hiraii (north-western Pacific)

- Vertebrae 39–45, usually ≤ 44; gill rakers 19–28, usually ≤ 25. Juveniles with a long (usually protruding far beyond middle of eye) or short chin barbel, if short, not pear-shaped or cordiform................................................. 2

2. In adults, pelvic fins with a large dark spot distally, anal fin usually with dark dots................ Cypselurus naresii (3)

- In adults, pelvic fins usually without a large dark spot distally (except C. o. crockeri ), anal fin usually unpigmented....... 7

3. Chin barbel in juveniles whip-like, very long, usually as long or longer than standard length........................................................................................ C. n. naresii (eastern Australia and Polynesia)

- Chin barbel in juveniles not whip-like, moderately long, usually shorter than standard length......................... 4

4. Dorsal-fin rays 12–13; predorsal length 69.1–70.6% SL; pelvic-fin base anteriorly placed (index cV/pV 0.81–0.89, cV 32.2– 33.1% SL, aV 56.3–58.0% SL); chin barbel in juveniles dark with a dark median keel............................................................................................... C. n. septentrionalis (north-western Pacific)

- Dorsal-fin rays 9–13, usually 11 or fewer; predorsal length 68.8–75.7, usually> 71.0% SL; pelvic-fin base posteriorly placed (index cV/pV 0.88–1.22, cV 34.1–41.8% SL, aV 58.8–65.4% SL); chin barbel in juveniles dark with a pale median keel.... 5

5. Vertebrae 40–41, precaudal vertebrae 25–27; in adults 145–185 mm SL pelvic-fin length 24.7–27.9% SL, anal fin of adults unpigmented; chin barbel of juveniles mostly pale....................... C. n. socotranus (Gulf of Aden and Socotra)

- Vertebrae 40–45, usually ≥ 42, precaudal vertebrae 26–30, usually ≥ 27; in adults 145–185 mm SL pelvic-fin length 27.4–35.1, usually> 28.0% SL; anal fin of adults usually with dark dots; chin barbel of juveniles mainly dark.................... 6

6. Predorsal scales 29–37, usually ≥ 32; vertebrae 42–45, usually ≥ 43; in juveniles ≤ 90 mm SL head lower surface mostly pale, chin barbel with a pale longitudinal streak distally................. C. n. albitaenia (western and central Indian Ocean)

- Predorsal scales 27–35, usually ≤ 31; vertebrae 40–44, usually ≤ 42; in juveniles ≤ 90 mm SL head lower surface mostly dark, chin barbel usually without a pale longitudinal streak distally.................................................................................................... C. n. ordinarius (eastern Indian and western Pacific oceans)

7. Pelvic-fin base nearer to origin of caudal-fin lower lobe than to head posterior margin (index cV/pV 0.97–1.46, usually> 1.00)............................................................................................... 8

- Pelvic-fin base nearer to head posterior margin than to origin of caudal-fin lower lobe (index cV/pV 0.78–1.06, usually <1.00).............................................................................................. 10

8. Vertebrae 42–45, usually ≥ 43, precaudal vertebrae 28–30, usually ≥ 29; index cV/pV 1.04–1.46, usually> 1.17; in adults pectoral fins usually not protruding beyond end of dorsal-fin base, pelvic fins usually pigmented......... Cypselurus opisthopus (9)

- Vertebrae 40–43, usually ≤ 42, precaudal vertebrae 26–29, usually ≤ 28; index cV/pV 0.97–1.24, usually <1.17; in adults pectoral fins usually protruding beyond end of dorsal-fin base, pelvic fins usually pale............................................................................ Cypselurus persimilis (eastern Indian and western Pacific oceans)

9. In adults snout length 4.2–5.5, usually> 4.6% SL, pelvic fins usually with a weak dotted pigmentation................................................................... C. o. opisthopus (eastern Indian and western Pacific oceans)

- In adults snout length 4.0–4.6% SL, pelvic fins usually densely pigmented................... C. o. crockeri (Melanesia)

10. In adults jaw teeth mainly tricuspid, palatine teeth numerous; juveniles with a tiny chin barbel never protruding beyond nostril........................................................... Cypselurus nossibe (western Indian Ocean)

- In adults jaw teeth mainly conical and with additional cusps, palatine teeth sparse or absent; juveniles with a long chin barbel always protruding beyond nostril........................................................................ 11

11. Vertebrae 41–44, usually ≥ 42; dorsal fin length 16.9–19.3, usually <19.0% SL; juvenile chin barbel ribbon-like, dark with a pale median keel...................................................... Cypselurus comatus (western Atlantic)

- Vertebrae 39–42, usually ≤ 41; dorsal fin length 18.1–24.1, usually> 19.0% SL; juvenile chin barbel necktie-like, pale with two dark zones..................................................................... Cypselurus angusticeps (12)

12. Predorsal scales 23–31, usually ≤ 27, transverse scales 7–8½, usually ≤ 7½; in juveniles <150 mm SL body width 13.2–17.4% SL, head length 23.7–28.4, usually> 24.0% SL, head depth 17.1–20.0, usually ≥ 17.5% SL.............................................................................. C. a. angusticeps (Indian Ocean and West Central Pacific)

- Predorsal scales 25–31, usually ≥ 27, transverse scales 7–9½, usually ≥ 8; in juveniles <150 mm SL body width 12.9–13.0% SL, head length 23.8–23.9% SL, head depth 16.9–17.3% SL...................... C. a. folletti (eastern Pacific Ocean)

Figures

FIGURE 2. Chin barbel of Cypselurus hiraii (ventral view of head). (A)—47 mm SL, IORAS 04133, 41°35–58’N 138°28’E; (B)—50 mm SL, IORAS 04132, 41°31’N 142°17’E; (C)—52 mm SL, IORAS 04132, same place; (D)—61.5 mm SL, IORAS 04127, 37°N 160°E; (E)—71 mm SL, IORAS 04133, 41°35–58’N 138°28’E

FIGURE 4. Size variation of morphometric characters (in % SL) in Cypselurus hiraii (◊), C. opisthopus (∆), C. naresii (○), C. persimilis (+), C. angusticeps (□) and C. comatus (x): (A)—cephalo-ventral distance divided by postventral distance (index cV/ pV); (B)—dorso-caudal distance (Dc); (C)—pectoral-fin length (lP); (D)—length of the first pectoral-fin ray (lP1); (E)—height of dorsal fin (HD); (F)—length of chin barbel (Lcir).

FIGURE 39. Chin barbel of Cypselurus angusticeps angusticeps (ventral view of head). (A)—36.5 mm SL, IORAS 04347, ~ 10°N 142°E; (B)—66 mm SL, IORAS 04330, 2°03’N 54°28’E; (C)—93 mm SL, IORAS 04341, 10°00’N 126°30’E; (D)—109 mm SL, IORAS 04340, 15°53’S 102°28’E; (E)—136 mm SL, IORAS 04320, 6°58’S 154°00’W; (F)—143.5 mm SL, IORAS 04335, ~ 7°N 135°31’E.

FIGURE 45. Cypselurus comatus. (A)—66 mm SL, IORAS 04368, 13°32’S 32°57’W; (B)—80 mm SL, IORAS 04366, 19°40’N 76°42’W; (C)—99 mm SL, IORAS 04367, 17°46’S 35°30’W; (D)—134 mm SL, IORAS 04359, 19°14’S 34°02’W; (E)—196 mm SL, IORAS 04365, 20°41’–21°00’S 34°46’–34°48’W.

FIGURE 46. Chin barbel of Cypselurus comatus (ventral view of head). (A)—66 mm SL, IORAS 04368, 13°32’S 32°57’W; (B)—99 mm SL, IORAS 04367, 17°46’S 35°30’W; (C)—144 mm SL, IORAS 04369, Atlantic Ocean.

FIGURE 47. Size variation of morphometric characters (in % SL) in Cypselurus comatus (x—north-western Atlantic; +— south-western Atlantic; ж—Atlantic Ocean, precise locality unknown). (A)—preanal distance (aA); (B)—head length (c); (C)— eye diameter (o); (D)—pelvic-fin length (lV); (E)—dorsal-fin height (HD).

FIGURE 48. Pectoral (A–C) and pelvic (D–G) fins of Cypselurus comatus. (A)—80 mm SL, IORAS 04366, 19°40’N 76°42’W; (B & E)—99 mm SL, IORAS 04367, 17°46’S 35°30’W; (C)—193 mm SL, IORAS 04363, 20°41’–21°00’S 34°46’–34°48’W; (D)—66 mm SL, IORAS 04368, 13°32’S 32°57’W; (F)—144 mm SL, IORAS 04369, Atlantic Ocean; (G)—196 mm SL, IORAS 04365, 20°41’–21°00’S 34°46’–34°48’W.

FIGURE 49. Distribution of Cypselurus comatus (○—our data; ◊—data from literature: Mitchill 1815; Bruun 1935; Breder 1938; Staiger 1965; Kovalevskaya 1982; Monteiro et al. 1998; Quattrini et al. 2004; Schwartz 2006). Filled symbols indicate capture location for fishes ≤ 100 mm SL, open symbols indicate capture location for fishes> 100 mm SL.