puma en Sp 200. ee Rat Ce Sa br WATE "Sa feed” asi). A Review of the Flying Fish Genus Exocoetus (Exocoetidae) with Descriptions of Two New Species from the Southern Pacific Ocean N. V. Parin and I. Shirshow Institute of Oceanology (IORAN), Russian Academy B. Shakhovskoy of Sciences, Nakhimovskii pr: 36, Moscow, 117218 Russia Received April 6, 2000 ‘Abstract—Flying fishes of the genus Exocoetus are widely distributed in warm waters of the World Ocean ‘They inhabit predominantly the open seas making the bulk of biomass of the middle-sized epipelagic nekton. ‘The genus comprises five spevies—circumtropical E. volitans, Indo-Pacific E. monocirrhus, Allanic E. obtu- sirostris, and two new South-Pacific species described is 154 mm, ZIN 52026, type locality 20°12’ 8, 173°05’ W) in the present paper. E. gibbosus, n.sp. (holotype SL is distributed in the southern subtropical gyre between 10° and 32° 5, E. peruvianus, asp. (holotype SL 156 mm, ZIN $2029, 8°01" S, 82°30’ W) is distributed in a small site in the northern part of the Peruvian current between 3° and 22°30’ S. These species are related to E. obtusirostris but differ from it in the strong development of the posterolateral process of cleithrum, reaching beyond the posterior edge ofthe scapular orifice and in fewer, asa rule, rays in pectoral fins. E. gibbosus and E. peruvianue differ in body form and depth, and in some ‘morphic state of some characters, the hypothetic scheme ‘etus is elaborated, implying that E. gibbosus and E: peru ‘The flying fishes belonging to the genus Exocoetus undoubtedly dominate in the middle-sized nekton of the tropical epipelagial zone of the World Ocean out- side of neritic zones. In open waters, they make about 80-90% of the total numbers of diving fishes (Bruun, 1935; Nesterov and Grudtsev. 1980; Parin, 1983; Grudisev et al., 1986), Traditionally, three widely distributed species were recognized in this genus—cireumglobal E. volitans, Indo-West-Pacific E monocirrius. and E. obwusiros- tris, which live in the southem subtropical Pacific ‘Ocean (Parin, 1961a; Kovalevskaya. 1980). However, samples from different parts of their ranges were never compared. The goal of the present study was to com- pare the large-scale populations inhabiting different Oceans (superpopulations) of these three species to reveal their intraspecies structure, It was found that three closely related species were mixed under the name E. obtusirostris, two of which are described here as new species. MATERIAL AND METHODS The present study is based mainly on the material collected in cruises of the Institute of Oceanolozy of the Russian Academy of Sciences expedition vessels in 1953-1989 (some materials are deposited in the Zoo- logical Institute of the Russian Academy of Sciences~ ZIN and the Zoological Museum of Moscow Univer- sity—ZMMGU). Other collections were also used: S31 ‘other morphometric characters. Considering the apo- ‘of phylogenetic relationships within the genus Exoco- wianus are the most advanced species. those of the Institute of Biology of Southern Sea of the National Academy of Sciences of Ukraine (InBYuM); of the Atlantic Research Institute of Fisheries and Oceanosraphy. Kaliningrad (AtlantNIRO): and of ‘many other foreign museums—AMS, ANSP, BMNH. BSKU, CAS, CSIRO, FAKU, HUMZ, ISH, LACM, MCZ. MNHN, NMNZ. NMV, NSMT, SAM, SMF. SIO, USNM, RUSI, ZMB. ZMH, ZMCU, and ZMUU (acronyms explained in Eschmeyer, 1998). Most of these materials are examined and identified by the first. author in places of their permanent deposition ‘The number of fish of the genus Exocoenus identi- fied in the course of our multi-year work totaled many thousands. In many specimens, the length was mea- sured, and some main measurements and counts of some of the meristic characters were made (frequently the number of gill rakers and scales in the transverse row), but the information from labels (place of deposi- tion, registration number, coordinates, and the date of capture) are given only for specimens fully morpho- metrically studied. The same information is given in the text for specimens captured in marginal parts of species areas. Measurements were made mainly according to Parin (1960, 19614); the following abbreviations were used: aA. ab, and aV—the anteanal, antedorsal, and anteventral distances; €V and pY—the distances from the posterior edge of the operculum tothe beginning of the caudal fin base and from the latter to the beginning of the lower lobe of the caudal fin; c—head length$32 po—postorbital distance; o—horizontal diameter of the eye: ao—snout length; b—barbel length; He, H, and h—the greatest depth of head and trunk and the least caudal fin depth; p—body width above the bases of pectoral fins; DC—dorsocaudal distance: IP and IV length of pectoral and ventral fins; ID and !A—length of bases of dorsal and anal fins. In addition tothe great- cest head depth, its depth at the vertical of the posterior eye edge was measured—Ho; the greatest body depth (H)_was measured separately for its anterior part (before the vertical through the pectoral fin base end— HI) and posterior (behind this vertical—H2) part. The ‘width of the bony interorbital space (io) was measured at the level of the eye center instead of the usual method In the key and in comparisons of species, the values of morphometric characters were measured t0 0.5% of SL. The investigated meristic characters comprise the number of rays in the dorsal (D), anal (A), and pectoral @®) fins, the number of predorsal scales (Sq. pred), the number of scales in the anteriorly directed oblique transverse row between the beginning of the dorsal fin and the lateral line (Sq. tr) (presence of an additional row of smaller scales under the dorsal fin base is desig. nated by "+"), the number of rakers on the first gill arch (Sq, br), and the number of vertebrae (Vert), with sub- division into trunk vertebrae (Vert. pe) and caudal ver- tebrae (Vert. c) In descriptions of new species, the numerical values of meristic characters and proportional measurements are indicated at first for holotypes, then in parentheses for paratypes, and separately for other investigated specimens, ‘The synonymy comprises all publications contain- ing original descriptions of taxa and introducing new binomial combinations, as well as the articles pub- lished inthe last several decades. A more complete bib- Tiography is available in the publications by Abe (195: 1957), Parin (1960, 1973), and Parin and Gibbs (1990). ‘The maps of the geographical distribution of species are composed by the original materials and by the reli- able data from the cited publications. Some marks may designate several closely situated capture localities. All original drawings are made by the second author, He also made mass measurements and caleula- tions of meristie characters ‘THE GENUS EXOCOETUS LINNAEUS Exocoetus Linnaeus, 1758: 316 (the type species E, volitans Linnaeus, 1758: by monotypy). Halocypselus Weinland. 1858: 385 (the type species H. mesogaster Weinland, 1858 = E. volitans L., 1758; by monotypy) PARIN, SHAKHOVSKOY Diagnosis ‘Transverse section of anterior body part rounded, its ventral surface not flattened. Predorsal scales 16-23, scales in oblique transverse row between the beginning of dorsal fin and lateral line 6-84. Snout short, its length is smaller than eye diameter. Jaws equally long. ‘Teeth on jaws absent or rudimentary. D 12-15 (16), 12— 15, P1 (12) 13-16, Dorsal and anal fins low, with the longest rays in the anterior part, situated opposite each other, beginning approximately at the same vertical line. Pectoral fins very long, atiain or almost attain the caudal fin base, Ventral fins shor, in adults hardly reach the middle of ventroanal distance, their first or second ray longer than other rays. Ventral fins situated closer to the beginning of snout than to the caudal fin base and more than by three times closer to the posterior edge of operculum than to the beginning of the lower lobe of caudal fin ‘The osteological characteristic of the genus is pub- lished by Parin (1961b). COMPOSITION OF THE GENUS The genus Exocoetus comprises five species. The following key is suggested for their identification. 1(2). Gill rakers 28-37. Scales in the anteriorly directed oblique transverse row between the beginning of dorsal fin and lateral line 6-6+. Juveniles with elongated body (at SL less than 80 mm the body depth 16-20% SL) and short ventral fins (13-16% SL), without bar- bel. Cleithrum without posterolateral. pro- cess (see Fig. 5). Supracccipital processes wo. .E, volitans 2(1), Gill rakers 22-29 (30). Scales in the anteriorly directed oblique transverse row between the beginning of dorsal fin and lateral line 7-8+. Juveniles with high body (at SL fess than 80 mm the depth 19-29% SL), with elongated ventral fins (15-33% SL), with a barbel or without it. Cle- ithrum with a posterolateral process. Supraoccipi- tal process one, sometimes bifid posteri- Oth oeeeenre 3 34). Pectoral fins usually dark-brown. Teeth scarce, separately situated, as a rule on both jaws. Juveniles SL (13) 15-70 (90) mm with unpaired barbel, situated on lamellar out- growths of inner edges of demalia (see Fig. 9. = E-monocirrhus 4(3). Pectoral fins usually light-brown or gray. ‘Teeth, if any, very rarely spaced, often fully absent on one or both jaws. Juveniles without barbel, lamellar outgrowths of inner edges of dentalia absent 5 5(6). Pectoral fin with T 14-16, usually 1 15~ 16 rays. Posterolateral outgrowth of clei- theum head short, not attaining the posterior edge of scapular oritice (see Fig. 5). (Greatest JOURNAL OF ICHTHYOLOGY Vol. 40 Suppl. | 2000A REVIEW OF THE FLYING FISH GE body depth 16-20.5% SL. Head depth 16- 18.5% SL, interorbital distance _8-10% SDvcorens LE. obtusirostris 6(5). Pectoral fin with I 13-15, usually 113-14 rays, Posterolateral process of cleithrum head long, attaining beyond the posterior edge of scapular orifice (ee Fig. 5) feveeesnenee 7(8). At SL over 100 mm, the body reaches the ‘greatest depth atthe bases of pectoral fins. The greatest body depth in adults 19-22% SL, head depth 19-20.5% SL, interorbital di tance 9-10.5% SL; in juveniles SL under 80 mm, 23-29% SL, 22.5-38.5% SL, and I~ 15% SL E, gibbosus 8(7). At SL over 100 mm, the body reaches the ‘greatest depth as a ruie beyond the bases of pectoral fins. The greatest body depth in adultss15-19.5% SL, head depth 15-18% SL, and interorbital distance 7.5-9% SL; in juve~ niles SL under 80 mm 19.5-23.5% SL, 19.5— 23% SL, and 9.5-12.5% SL... peruvianus EXOCOETUS VOLITANS LINNAEUS Exocoetus volitans Linnaeus, 1758: 316 (holotype ZMUU Linn, Coll. 59: after Eschmeyer, 1998: 1775; “in alto pelagico Europaeo et Americano”). Exocoetus evolans Linnaeus, 1766: 521 (holotype BMNH 1853.11-12.181: after Eschmeyer, 1998: 553). Exocoetus volans Solander in Richardson, 1846: 264 (syntypes, BMNH: after Eschmeyer, 1998: 1774; “Seas of China and Polynesia”) Halocypselus mesogaster (non Bloch, 1795): We land, 1858: 385 (types unknown: after Eschmeyer, 1998: 1071; “coast of North America”). Exocoetus chilensis Abbott, 1861: 471 (syntypes ANSP 7498 and 7499: “Chile” Exocoetus volitans: Bruun, 1935: 28-34 (descrip- tion, the Atlantic Ocean). Breder, 1938: 31-33 (the Western Atlantic Ocean). Abe, 1957: 1141-1147 (syn- ‘onymy; description; the Westem Pacific Ocean). Parin, 1960: 215-218 (description, the North-Western Pacific Ocean). Evans & Sharma, 1963: 53-55 (the Central Atlantic Ocean). Kovalevskaya, 1964: 205-211 (early developmental stages). Munro, 1967: 119 (New Guinea). Kotthaus, 1969: 9-11 (western part of the Indian Ocean). Belyanina, 1975: 140 (the Caribbean Sea). Fahay, 1975: 16-17 Florida). Kovalevskaya, 1980: 217-218, 253-255 (early stages; distribution in the Pacific and Indian oceans). Yoshino, 1984: 80 (Japan). Grudtsev er al, 1986: 920-928 (morphometry, distribution in the Alaotic Ocean). Parin, 1986: 616— 617 (the North-Eastem Pacific Ocean). Heemstra & Parin, 1986: 394 (South Africa). Chen, 1987: 49-51 (carly developmental stages; the North-Western Pacific Ocean). Paxtom & Hanley, 1989: 333 (Australia) Parin & Gibbs, 1990: 587 (synonymy: the eastern tropical ‘Atlantic Ocean). Shiganova and Kovalevskaya. 1991 JOURNAL OF ICHTHYOLOGY Vol. 40 Suppl. 1 2000 SNUS EXOCOETUS (EXOCOETIDAE) $33 94-95 (larvae; the North-Eastern Atlantic Ocean). Belyanina, 1993: 110-111 (larvae; eastern Austra- lia). Parin, 1995: 1101 (the eastern tropical Pacific Ocean). Périn, 1996: 301 (the western central Pacific Ocean). Exocoetus volitans vagabundus Whitley, 1937: 216 (holotype AMS 9A.6854, “Tasman Sea between Syd- ney and Lord Howe Island”). Exocoetus obtusirostris chilensis: Fowler, 1944: 318-319 (Galapagos Islands). Fowler, 1945: 31 (the Grawing from the type specimen of E. ckilensis Abbott). TYPES E, volitans is the first of the named species of flying fishes. The identity of two species described by Lin- naeus (1758, 1766), E. volitans and E. evolans was shown by Braun (1935). E. volans Solander, without any explanations, was introduced into the synonymy of E. volitans by Giinther (1866); the type material obvi- ously deposited in BMNH (Eschmeyer, 1998) was never examined. The types of E. chilensis Abbott were investigated by Fowler (1944) who, after erroneously comparing the numberof gill rakers on the lower part ofthe first gill arch in them with the complete number of rakers in E. obtu- sirostrs (see Parin, 19612, p. 42), assigned them to the latter species as a subspecies E. obtusirostris chiliensis. ‘The syntypes of E. chiliensis (two specimens SL 156 and 158 mm) deposited in the collection of ANSP under the numbers 7498 and 7499 and reinvestigated by the first author have the following characters: D 13, 14; PT 13,1 13; Sq, pred 20, 19; Sq. tr 6+, 6+: Sp. br (let) 8 + 36, 10 + 24: measurements in % SL:'¢ 25.9. 26.2; 07.6, 7.1; 10 8.3, 8.4; H 16.9, 16.5:h 6.6, 6.7: IP 765, 16.6; 1V 12.0. 139. The holotype of E. volitans vagabundus Whitley (AMS IA 6854), a large juvenile SL. 92 mm, also rein- vestigated by the first author. All its characters are within the variation range of E. volitans: D 13, A 13, P 114, Sq. pred 20, Sq. tr6, Sp. br7 + 24; in % SL:¢ 27.7, 081, i0 9.5, H 18.1, h65, IP 71.7, IV 130. MATERIAL, Atlantic Ocean—in total, 69 specimens SL 29.5 176 mm: the collection of IORAN (without_num- bers)—four: 72-86 mm, 17°57’ N, 33°14” W, Febru- ary 15, 1976; one: 109.5 mm, 27°07’ S, 38°35” W, Feb- ruary 12, 1976; one: 134 mm, 15205" N, 21°51’ W, November 7, 1973; one: 120 mm, 6°00’ N, 20°00" W, January 4, 1965; one: 90 mm, 30°35’ N, 67°20" W’, July 17, 1978; two: 70.5 and 73 mm, 2°04" S, 35°57” W. February 14, 1976; two: 41.5 and 57 mm, 7°54’ N, 53°16 W, April 3, 1969; two: 40.5 and 43.5 mm. 12°22’ N, 59°58" W, March 27, 1969: one: 39 mm, 19°39" N, 67°34’ W, February 4, 1973; one 61 mm,S34 (16'S, 18°15’ W, January 18. 1967; one: 69 mm, S°10'N, 23°36" W, February 22, 1976; three: 53 61 mm, 10°21” N, 78°50” W, February 19, 1978; one: 45.5 mm, 1°21” S, 34°45’ W, February 24, 1964; one: 35 mm, 3°58” $, 21°59 w, November 11, 1971; five: 144-167, 12°47" N, 62°07 W, February 17, 1973; one: 142 mm, 1°58’ $, 10°21” W, September 7, 1970; one: 145 mm, 14°16" N, 19°28" W, March 18, 1971; one: 171 mm, 2°41" S, 11°23" W, September 6, 1970: three: 156-162 mm, 14°39" N, 74°16’ W. March 3, 1971; two: 150 and 155 mm, 14°10°N, 66°40 W, February 21, 1973; six: 106-149 mm, 0°20 S, 8°47" W, September 25, 1970; two: 142 and 146 mm, 16°35" N. 60°38" W, March 7, 1969; three: 165-176 mm, 6°59" S, 22°19’ W, Decem- ber 11, 1975; two: 167 and 173 mm, 9°33" S, 4°11’ W, ‘April 30, 1968; one: 152 mm, 4°25'S, 1°51’ W, April 16, 1968; one: 175 mm, 15°08", 28°16" W, February 14, 1976; one: 162 mm, 13°18" N, 61°59" W, February 18, 1973: one: 174 mm. 8°34" N, 22°28" W, February 23, 1976; one: 162 mm, 8°06" S, 28°40’ W, February 18, 1976: eight: 29.5-61.5 mm, the Caribbean Sea; five: 152-163 mm, Brazil basin, Indian Ocean—in total, 78 specimens SL 30. 163mm: the collection of IORAN (without num- bers)—one: 103 mm, 8°40” N, 61°54’ E, November 12, 1960; three: 58-115 mm, 10°03" S, 108°00" E, Novem- ber 4, 1959; two: 104 and 119.5 mm, 19°27" N, 59°24" E, November 4, 1960; two: 75.5 and 82 mm, 13°40" S, 104°49" E, March’ 20. 1961; two: 83 and 100 mm, 13°00" S, 91°15” E, March 1, 1973; one: 98.5 mm, 9°38’ S, 15°05’ E, October 24, 1959: two: 53 and 91 mm, 11°08" S, 91°07" E, March 20, 1973; two: 41 and 93 mm, 11°08" S, 104°32" E, March 3, 1973; one: 119 mm, 0°55’ N, 62°33" E, November 17, 1960; one: 61.5 mm, 14°34’ S, 70°57" E, December 19, 1960; one: 41'mm, 15°00’ S, 106*46” E, July 21, 1962; 1959; one: 77 mm, 12 5, cone: 62 mm, 11°35" S, 99°49’ E. March 5, 1973; one: 70 mm, 13°04" §, 96°23" E, March 22,” 1973; one: 43 mm, 16°05 S, 76°16’ E, July 18, 1960; one: 59 mm, 15°53”, 102°28" E, November 26," 1959: one: 35.5 mm, 12°49" S, 83°00" E, January 4, 1961; one: 30 mm, 13°03" N, 87°03" E, September 18, 1962: two: 135.5 and 139.5 mm, 3°37’ §, 39°54" E, March 16, 1960; one: 136 mm, 1°07” N, 56°35" E, March 17, 1983; one 127 mm, 9°19’ S, 73°12’ E, August 20, 1978; one: 136 mm, 9°12’ S, 68°37’ E, August 18, 1978: two: 136 and 147 mm, 11°36” S, 91°26' E, October 3, 1972; two: 128 and 154 mm, 9°19" S, 64°19’ E, August 17, 1978; wo: 149 and 152 mm, 8°24" N, 55°10" E, July 12, 1978: two: 139 and 153 mm, 10°35’ S. 50°03" E, April 10, 1983; one: 147 mm, 1°07" N, 36°34” E, March 15, 1983; one: 142 mm, 10°19" S, 36°56” E, March 29, 1983; one: 138 mm, 10°58” S, 50°35” E, April 4. 1983: two: 136 and 137 mm, 6°53" S, 53°38’ E, March 5, 1960; three: 120-127 mam, 10°19°N. 33°14” E, October 29, 1960; two: 135 and 142 mm. 3°04" S, 44°51” March 18, 1960; one: 141 mm, 11°48” N, 82°49’ E, JOURNAL OF ICHTHYOLOGY PARIN, SHAKHOVSKOY January 26, 1961; one: 144 mm, 5°00’ N, 90°52" E. February 11, 1961; three: 135-152 mm, 9°14’ N. 93°40’ E, March 6, 1961: one: 146 mm, 3°00" N. 83°59’ E, November 27, 1962; three: 134-150.5 mm. 14°45’ N, 88°08" E, May 11, 1957; one: 142 mm, 9°07’, 104°39" E, March I, 1973; four: 635-156 mm, 15°45" S, 99°56" E, March 24, 1973; four: 155-163 mm, 22°06’ S. 105°01"E, November 28, 1959; one: 159. mm, 16°01’ S, 11°35" E, November 28, 1959: two: 30 and 43 mm, Somali Basin, June 6, 1956; the collection of ZNCU—without numbers, two: 140 mm, 9°20" S, 49°27’ E, March 6, 1951; one: 149 mm, 6°38" N, 94°53’ E, May 8, 1951 Pacific Ocean— in total, 75 specimens SL 31— 170mm: the collection of TORAN (without num- bers)—two: 55 and 96 mm, 18°31” N, 175°00" W. ‘August 17, 1970; one: 115 mm, 12°13” N, 179°59" E. May 31, 1970; one: 123 mm, 11°25” N, 143°05" E, ‘April 18, 1975; wo: 95 and 100 mm, 9°01’ 8, 152°52'E, July 22, 1957; one: 107 mm, 5°00" N, 135°00" E, Janu" lary 31, 1966; two: 765 mm and 1i4 mm, 0°27" S. 84°58" W, August 28, 1968; one: 121 mm, 12°11’ N. 179°48" E, May 29, 1970; one: 81.5 mm, 21°35” S, 174°46' W, December 25, 1957; one 103.5 mm, 14°00" N, 137°31" E, May 29. 1958; one 98 mm, 9°59’ N. 134259’ E, January 29, 1966; three: 57-84 mm, 13°27" N. 143239" E, April 29. 1975; one: 40 mm, 20°03" 151948" W, January 30, 1959; one: 65_mm, 5°54” 100°12’ W, October 31, 1967; two: 67 and 73 mm. 16°08"_N, 157922" W, August 8, 1970; three: 42-51.5 mm, 19°03 N. 171°09" E, August 23, 1970: five: 48-74 mm, 6°03" N, 96°38" W, November 3, 1967: fone: 45 mm, 0°15" N, 179°37 W, June 5, 1967; one: 51.5 mm, 21°03" N, 167°0% B, September 7, 1969; one: 35 mm, 12°40’ S, 85°35" E, February 15,'1974; one: 35 mm. 0°01’ N, 140°03’ W, September 21, 1961; two: 31 and 38.5 mm, 6°04" S, 139°59" W. September 2. 1961: four: 142-152 mm. 32°33’N, 158°45" E, August 22. 1958: one: 150 mm, 8°01’ S, 82°30° W, February 10. 1982; one: 163 mm, 8°13" S, 83°58" W, February 9. 1982: one: 154 mm, 2°43" N, 128°04" E, March 13. 1966: one: 161 mm, 9°00’ N, 127935’ E, March 1, 1966: three: 164-169 mm, 0°00, 154943" W, February 6, 1974: fone: 155 mm, 4°46" N. 126914” E, March 15, 1966: two: 155 and 159 mm, 0°03’ N, 139°02' W, January 30. 1974; one: 139 mm, 13°26" N, 119°4¥’ E, March 18, 1966; one: 168 mm, 0°02" S, 122°04" W. January 25. 1974: three: 160-170 mm, 5°58” N, 139°57’ W, Sep- tember 16, 1961; two: 153 and 168 mm, 16°01" N. 158203’ W, June 25, 1965: three: 162-165 mm, 3°02" N. 140°08* W, September 18, 1961; two: 158.5 and 162 mm. 10°05" N, 157244” W. May 20, 1965; one: 132 mm. 28°37" N, 139°18' E, September 24, 1957; one: 147 mm, 34°48" N, 149°18” E. July 3, 1957: one: 154 mm. 27°25' N, 132°53’ E, October 30, 1955; two: 150 and 157 mm, 27°25" N, 143°20" E, October 20, 1955; five: 134-161 mm, 30°37" N. 142°17" E, October 2, 1955 ‘one: 162.5 mm, 26°04" N. 143904" E, October 11,19: fone: 160 mm, 29°16" N. 143°10" E, October 8, 1955: Vol. 40 Suppl. 1 2000835 A REVIEW OF THE FLYING FISH GENUS EXOCOETUS (EXOCOETIDAE) ~ Jo fie fel-}--=]e forjore fe }-|-lafufarls |= farfacte fe }-Isrfoela | - eta fee for] -f-]- Ju ferfsrfafarfe f-]-]e Jsclor fale fsele |-|-le feclel- ~|- J ferp-] fete fre/selsefarl-|-]-J6 lsele fa ]sr|sclorle a foelectoe|e vfejefufe fe }-]-]-]-]-]-]-le rf-fefe fs fa “Jo pelep-f-]-feds fede fe f-l-]-]o fee l- fede do |-fadule ds de ~|o fe fop- fafa ye fo focfocje J-|-|-le fiele fads fecly |-|-]a forfucls e ) leper} | - | ~ | [er fec} olor) eet ee ae ~@ sie fat pode [el ictonforte |~}-]-Jeclar]—|-]e foele |-|-]o foeler| mmeo ouseg so seg Vis ely =) Teds fe file fe |-|-}-]~ Jeter) c}-locler}-|-]-|z foclz | smoosueaso son Tjorfos jr {-|-)-]e ferjocfarfs }-|-|-le Joule] -|-forforlr }-]-|> fects 200 |r] o> fered =] ]er}es] 06] se foe }s luz}esjor}-!-}-loc}as]ar] - fer lector| - ow ~|- Jo fa for foe] -]-fe ferfselaelar{a forjmte |-f-]-{6 fel -}-lizja|-|- ra00 Jo} fofiefo}-]-]e futarfse}or]e Jorforfs |-]-|-Je fee }-lelarfe | - 090 el ale | ale tale eal Wee ele ele ele epleel pic foe tale lene ~|oyo]- fests ]o[-fe forfecfects |- fe Jor}-]-]-]-]e Isefs }-]o foels | - we230 9 sung +8 |e] c | 49] 9 [sz] zz | refer [at fer [or] oc | or] sr [er | or | ez] 9c] scl vc] or | ar] orl ew lew abs pardbg au ada, 2A sopads sete jo squmg esqaLIOA Jo JoquIM, smzosexg snva8 ayy Jo satzads ut moa asronsues) anbiygo ut s2|895 put soqeos ysiopaxd ‘eigen Jo s2qUNN “Taq, 2000 Vol. 40. Suppl. 1 JOURNAL OF ICHTHYOLOGYPARIN, SHAKHOVSKOY $36 ur229 dytoeg wrarseaqinos, - fz fac lo] - ee ee ee Wee iecy eee ) culec jee | co - jo jw fs | - -jo lw foe |i} - fe fo fe | - | - foe foe fe ve fiorja | - | - to jee fas | ele je ful} -|z je lea ls wir f- | - - fo for pr f- fu fst joc fu | - | - for fee | - oe frp - | - - | foo fe far j cle fe fe |- fa fate fo otf fe | - | - tofar foe fe | - | - far foe fe j- fa ise [se je z jis fouto fo fe Jest} oz] z | 1 foe feo} oc | - | - Jee just] se rfofis te fa - Jor jes fsr} ce} - fs fos far} - | - fs fas | oz b |ve jse fe | = -fe jets |-|- Jo lw ls |-|- |e lade ~ fie foe | =] - TSU | Coe le fat ee ce ger ee lol lice |e ~ [oz | we] se] a te }serfacr} a | - | 1 fest} asc} or | 1 | as a ee 4]: pce je fo} -]- | - fe jus | - foe = le oe | ufis fre te | - | - fe fs fe ja fae = oie tc |e s jor |sr fa f - “a e 5 eM oene |ee or fue fos fr | - jot fee ee eeelatl = a | ot | st] et | et z{rfofrjielisa|o za | on | st | a a va q sopads shes Jo s9quiny, J J9QUION "7 IGN, Suppl. | Vol. 40 JOURNAL OF ICHTHYOLOGY837 A REVIEW OF THE FLYING FISH GENUS EXOCOETUS (EXOCOETIDAE) el=ielieileii ig 6 |e fads . yong wsoseay smuniod - forte fe fe fede eo fu ja fe 209 ayioeg YpnOS, soso sp -}-7-]-]-]-]2 w | w | se | ow ease 104m S61 “wnmag sour Seale | -i|el | ms |e ioe] |. s fo fo fu -J-}-]-}-]- 4a 6 fu jn | IV MON, sunsansosqo o|-|.]-|/.4i./. st | sr | oe | ss vam Se ee |e e fo fo] a {1990 94H elas). a | 6 | w | si W290) 24y!9Fd JO SAL ole] e|-|4i|.'. 6 | o | w]e ‘W299 ueIpuy or | or | oe | 96 | x | ve Jo A e |e | iw le ]el a] s -];-]-]- e900 24ta0d rfe pele lwle fe fe || unang mp ce fo}w)efe fe te -|-]-]- S61 “unnag soe ‘omes s fs falejoafer | - -}-]-]- twe209 anueity sumo 3 se | ve | ec | ce | te | oc | oe aia] sl wz sarsads 4q'ds smizo20rg sna 24) 30 s2t90ds ut sroyer 18 Jo JOquINNY “— AIG, 2000 Suppl. 1 JOURNAL OF ICHTHYOLOGY Vol. 40PARIN, SHAKHOVSKOY $38 wwg (oze-90vWe IZ |(9'Ee-S 00 zz 97e-CODE Iz (upe-e 1s ze| (0 ze-¢oNR Oe | (972-1021 1Z|(H'Ez-FOZ)6 12] vi wee (oee-0 DoT (e'st-e yer es ez |(o'ee-t (see-woor ee (i pe-9 1zLz2) a stereo (srr-stneel|(esi-6 T-oeDoel| ('st-VeNo rl |OrI-vedovI|@I-TEDy'PL AL oese-r" (CLEEODTEL| (BLS 'SL-S'8)O'EL|O'LL-P ODT HL | B'EL-E'LSIS'RD|(7'RL-FOLIS'tL| Le PL (0'9L-¥'s9)¥ 04] al (see-o'6c)o Ie (Lee-roeds Ie (Lee Te-62)90¢|(e'ee-FODR TE, coor ielieeesevete] — oa (sr TDEPI esi-ieverilrer-cens tl bifevr ceDest a (e0969 | GETYED wesooe| rove werewe WL | OR-ONRD 4 (orespsxi (Wor-eonrsr|(esI-F OBI-roNTLI or-s ons (oI-esNext|(eor-oon9Cr cH (rsr-ysDrei|or-coneL|(eor-z" (ol-v9no8 (wsi-rs peo 1H Li-rsp99i (@xi-esnedi (e6t-z" roie leurrsps mn (sI-eeD0's! |(99I-PDzs1] (LI-6 lasr-o (or-vees. oH WLerurs | GoI-eol'6 | CeI-TooTl] Coca | Gores | OeI-vorii| Cross urzveonl o aesoor | (seorr| Gssvrs| @srHer| Wsoor] Ores | w's-eORr corres on CES | Grr | woos] rede] Crore] WeETOrE| (Es969 (66-9098 ® (rr-ezN6EI|O9I-LEDeFI|O9I-ErNESI|CSI-VEDY P| sI-9 LIP eDs sil eS-OED PL (sor-opsst od (892-9409 Se] (P'Be-S'SEIL'9T| (OOE-VLZIE'8z|(S'L2-7'SDE 9e|(O°6C-L-STIOYE| (PUE-L-LEN Z| (6 LE-U'SCIT9E|(1'6e-S'SENOLE| LOK-LLE)I-62| 2 ‘IETS ¥5|(OSS-TISILTS| (B'SS-G ISLES |(O'LS-ETSW PS (r'Ps-SosIS TS (F9S-TISW'ES|(C9S-r ISI ES|9rS-PeMtTs| Ad Koor-senysi{(eo-esneu (sor-wenooi{Ool-venso1|(eu-cenest a er|(o9r-ozrivrel ror-r ene ey] sLrvinser|(cs-eeeerl(TLy-veps ye] AR (919-s'¢9)0°99| (va9-879)6'99) (€19-S19)9's9|(r'89-er9)r99|(cL9-6eN)R'S9|(889-EPOr'99| ae woresmres| (-89-L P9899) (69-E 99179] (rRO-HH9IT 99 I PL-T'SHLRO|(Wo9-LODOH|(GOL-ESMT Eo] VE ww ve & & @ w @ (oout-0 oucor) Oort-ezw | Coxe-o08) | vLt-ozs0 | (ost-o18) | (oL-s62) 09 ses Tl ez Oreo eet sis uwe209 ayieg . 1ww299 wPipUL 1we099 anu} (As % uy suuuy woneues pur sunaw) nigo2oxg Jo saioee4> a1NAUHOAKLOW UE Suppl! Vol. 40 JOURNAL OF ICHTHYOLOGYA REVIEW OF THE FLYING FISH GENUS EXOCOETUS (EXOCOETIDAE) $39 the collection of ZNCU—S1 088, one: 137 mm, 10°05’N, DIAGNOSIS 96°06" W, December 7, 1960; the collection of SIO— —_Body oblong, its greatest depth 5.16.4 (in juveniles 58-215, one: 54 mm, 8°00’ N, 120°47’ W, October 9, SL under 80 mm 4.9-5.8) times in SL. Vertebrae (24— 1955. 26) + (18-20, usually 19-20) = 43-45, usually 4445, E.volitans @ E, monocirrhus Fig. 1. Skulls of Exceoetusvoltans and Exocoetas monocirrhus (a) dorsal and () ventral view), Designations of Sones: (bee) bas ‘occipitae, (bs) basisphenoideum, (coe) exooccipitle. (ep0)epiodeum. (eth! ethmoidale lateral. (0) frontal. im) nasal. (pro) prooticum. (ps) parasphenoideum. pt) pterodcum, (Soe) supraoceipitil.(spho) sphenoticum. Punctae show the opening in the Skull coo illed with cartilage, crossiny shows the emargination between protruding downwards chondral pas of Frontalia compris ing the anterior part of bran. JOURNAL OF ICHTHYOLOGY Vol. 40 Suppl. 1 2000sao PARIN, SH E.volitans E. obtusirostris |AKHOVSKOY E. monocierhus E. gibbosus Fig. 2. Ethmodiale laters (ventral view, lft side) in Bxocoetus voltans, , monocirus, , obtsirosis, and E. gibbosus Predorsal scales (16) 17-21, usually 17-20. Seales in the oblique transverse row’ 6-6+. Head 3.6-4.1 (in juveniles 3.3-3.7) times in SL. Gill rakers (28) 29-36 (37), usually 31-34. Teeth on jaws as a rule fully absent. D 13-15 (16), A 12-14 (15), P I (12) 13-15. Variational series of meristc characters are shown in Tables 1-3. Main measurements presented in Table 4. Pectoral fins light gray, with rather wide nonpigmented band along the posterior margin. Juveniles have no bar- bel. The greatest body length (SL) 176 mm In the orbital area of skull, the emargination between chondral parts of frontalia long and narrow; in adults its greatest width is 5.0-6.0 times in skull width at the same transect (Fig. 1). Supraoceipitale pentago- nal in dorsal view, with straight lateral sides, two pos- terior supraoccipital processes; pterotica with straight or convex outer margin; nasalia reach far beyond the anterior edge of frontalia (Fig. 1). Ethmoidalia lateralia with a smooth outer margin (Fig. 2). Operculum with a ‘convex lower margin; interoperculum narrow (its depth 2,5-2.8 times in its length); suboperculum without a deep emargination on the upper edge; praeoperculum with a rounded posterior edge (Fig. 3). Spinous pro- cesses of three anterior vertebrae approximately identi- cally wide or slightly increase posteriad (Fig. 4). Post- temporale bent, with a widened base; cle:thram without posterolateral process; basal plate of pelvic bone with three-four longitudinal outgrows directed anteriorly Fig. 5). DISTRIBUTION (Fig. 6) E. volitans is widely distributed in tropical waters of all oceans atthe temperature of 20,5-29.5°C (usually at 24-28°C) (Parin, 1962; Grudtsev et al., 1986), but its distributional range, strictly speaking, cannot be con- sidered circumglobal. In the Atlantic Ocean, this species is common between 20° N and 20° S, and only in the western meridional currents occurs beyond these limits. The north westernmost occurrence is noted at 35°57’ N, 73°13” W (a juvenile SL 63 mm, IORAN, no number, RV Vityaz-2, station 319, May '19, 1982), the south- westernmost occurrence is recorded in the Brazilian cur- rent at 30°02’ S, 36°18” W juveniles with SL 52-65 mm, ‘AtlantNIRO, no numbers, February 15, 1978). Catches of individual specimens may depend on the occasional penetration of “wandering” individuals and are recorded in the northeastem Atlantic Ocean off the Azores, Madeira, Morocco coast, and even in the west em part of the Mediterranean Sea (Tortonese. 937, 1958; Dollfus, 1955; Parin, 1973; Arruda, 1997). All or some of these occurrences may be based on erroneous identification. In the Gulf of Mexico and in the south- JOURNAL OF ICHTHYOLOGY Vol. 49 Suppl. 1 2000)‘A REVIEW OF THE FLYING FISH GENUS EXOCOETUS (EXOCOETIDAE) E. volitans sal a E, monocirrhus sop Fig. 3. Opercular hones of Exacoena volitans and E. monocirrhus, Designations of bones: \op) interoperculum. (op) operculum, (pop) praeoperculum. (sop) suboperculum eastern Atlantic Ocean, this species, obviously, is not present In the Indian Ocean, E. volitans is also widely dis- tributed from the African coasts to the Timor Sea. How- ever, itis absent in the Red Sea, the Persian Gulf, and in the northern part ofthe Bay of Bengal (the northemn- most occurrence in the Arabian Sea at 22°12’ N, 60°41’ E—one specimen, SL 79 mm, MCZ 136 378) JOURNAL OF ICHTHYOLOGY Vol. 40 Suppl. | ‘The southern boundary of distribution in open waters is situated at 20°-25° S, the southwestemmost occur- rence is in waters of southern Africa at 34°27" S, 25°57’ E (one specimen, SL 100 mm, SAM 23283, June 18, 1962), In open waters of the Pacific Ocean, the northern boundary of the ranze of E. volizabs is situated from 40° N in Kuroshio waters off Honshu (Kovalevskaya, 2000saz PARIN, SHAKHOVSKOY E.volitans E, monocirrhus Fig. 4. Anterior part of vertebral column of Exocoetus vltans and. monocirrhus. E.volitans Fig. 5 Skeleton of pectoral and pelvic gnlles of Exocoens volians, E, monocirrhus, and E. gibbosas lateral view, right side; com bination sek-p in dorsal view). Designations of bones and of ther elements: (el) ciitrur, (cor coracoideum, (peel) postlteral process of cleithrum, (p) postemporale, (cr) radialia, (se) scapula, (sel supralektrum. 1980), to the north of the Hawaiian Islands (one speci- New South Wales (AMS 1.4011 and I.4012, SL 139 and men, SL 163 mm, CAS 81 822, 28°17’ N, 16134’ W), 38 mm) to Kermadec Islands and via southern archipe!- and proceeds along 20° N to the extremity of Lower agoes of Polynesia—the southernmost occurrence at California (a specimen SL. 160 mm, CAS 81849, 23°30" S. 1118°12" W «two juveniles. SL 53 and between the San Lucas Cape and San Benedicto 70mm, SIO 38-303)—to the coast of Peru at 10° $ (in Island), the southern boundary—irom Wallongong. the southern hemisphere in the winter, the boundary JOURNAL OF ICHTHYOLOGY Val. 40 Suppl. 1 2000‘A REVIEW OF THE FLYING FISH GENUS EXOCOETUS (EXOCOETIDAE) veniles SL under 100 mm, open signs capture 30 6040 JOURNAL OF ICHTHYOLOGY Vol. 40 Suppl. 1 2000saa shifts to the equator). The southwesternmost capture is recorded at 16°11’ S, 87°05" W (female, SL 175 mm, from the collection of AtlantNIRO, SRTM 8459, March 27, 1968). In epicontinental seas of the western Pacific Ocean, E. volitans is recorded only in the north- cern (deep-sea) part of the South China Sea—the south- cemmost occurrences at 7°40 N, 108°12’ E (a juvenile CAS 81924) and 10°26’ N, 12°44" E (SL 129 mm, CAS 81.069), in inner seas ofthe Philippine Istands and in the East China Sea. Occasional records are known for the Yellow Sea (ZIN 14760) and the Tsushima Strait (Mori, 1952; Lindberg and Legeza, 1965), but the record from Peter the Great Bay (Parin, 1962a) is actu~ ally E. monocirrhus (see below). The species under ‘consideration was never recorded in the Strait of Mal- acca, the Java Sea, the Bali Sea, the Flores Sea, the Banda Sea, the Molucca Sea, the Ceram Sea, of the ‘Arafura Sea farther than 9°23" 8, 127%48" E (SL 149 mm, SIO 61-723), and the Gulf of Carpen- tari, ie, in'the chain of water basins separating the Pacific and Indian oceans. ‘Thus, the area of E. volitans consists of three sepa~ rate units—the Atlantic Ocean, Indian Ocean, and Pacific Ocean parts, and almost completely repeats the range structure of another oceanic fish—Cheilopogon ‘furcatus (Parin and Belyanina, 1998). In the latter spe- cies, the differences between populations from differ- ent oceans are more significant (sufficient for separa- tion of subspecies) than in E. volizans (see below). AGE AND GEOGRAPHIC VARIATION In principal body proportions, the juveniles SL. 30- 80 mm do not radically differ from adults (Fig. 7, also “Table 4), though some indices change with growth— the head length, eye diameter, interorbital distance, all indices of body depth, and length of ventral fins decrease, the length of pectoral fins increases. Small juveniles, SL under 50 mm, are characterized by the presence of two transverse dark bands situated between the third quarters ofthe dorsal and anal fins (including their proximal parts) and before the beginning of the caudal fin, No significant morphometric differences between populations inhabiting different oceans are revealed (ee Tables 1-4), There are minor differences in the ‘number of gill rakers (in the Pacific Ocean 31.5 +0.14, in the Indian Ocean 32.0 + 0.13. and in the Atlantic Ocean 32.7 + 0.12), in the number of scales in the oblique transverse row (in the Pacific Ocean frequently 6, in two other oceans 6+), and in the number of verte- brae (in the Pacific Ocean usually 45, on an average 44.6 + 0.08), in other oceans usually 44, on an average 44.3 £0.10 in the Indian Ocean and 44.1 + 0.07 in the ‘Atlantic Ocean). Morphometrical characters in samples from different oceans almost do not differ. The differ- ences in the length of ventral fins in juveniles from the northwestern Pacific Ocean noted by Parin (1960) in small material (original data) and from the western PARIN, SHAKHOVSKOY Avtlantic Ocean (after the data by Breder, 1938) are not confirmed by our material. COMPARATIVE REMARKS E. volitans differs from all other species of the genus Exocoetus in that it has @ somewhat more elongate body (this difference is well expressed in juveniles with SL under 100 mm), a smaller number of scales in the oblique transverse row (6-6+ versus 7-8+), more ‘numerous gill rakers, and some osteological characters. Of the latter, more important are some traits of the structure of the skull roof, of operculum, skeleton of paired fins, and of the anterior part of the vertebral col- umn, Two traits of the skeleton of E, volitans—the absence of the posterolateral process of cleithrum and. the presence of two posterior supraoccipital pro- cesses—are unique in the genus Exocoetus, but coin- cide with those in all other genera of the Exocoetidae. Exocoetus monocirrhus Richardson Ecocoetus monocirrhus Richardson, 1846: 265 (the holotype is unknown: after Eschmeyer, 1998: 1115; Sea of China), Exocoetus georgianus Valenciennes in Cuvier & Valenciennes, "1847: 136 (lectotype MNHN 601; “5° latitude ‘and 90° longitude orientale;” three paratypes: two specimens MNHN 1996-141 are captured together with the lectotype, the third specimen—MNHN B.84I is labelled “Atlantique, non loin du Brasil”), Exocoetus melanopus Giinther, 1868: 459 (holotype BMNH 1868.2.29.54: after Eschmeyer, 1998: 1056; Zanzibar) Exocoetus obtusirostris (non Giinther, 1866): Liit- ken, 1876: 294 (between Japan and the Hawaii. Exocoetus volitans (non Linnaeus, 1758): Weber & de Beaufort, 1922: 177-178 (the Java Sea). Halocypselus obtusirostris (non Giinther, 1866) Breder, 1928: 17 (waters of Central America, differ- ences from E. volitans). Holocypselus borodini Nichols & Breder, 1932: 1 (holotype MCZ. 6216; paratype AMNH 6998; Maur tis). Exocoetus monocirrhus: Bruun, 1935: 37 (men tioned). Nichols & Murphy, 1944: 237 (the Gulf of Panama). Abe, 1955: 1013-1020 (synonymy: descrip- tion; waters of southem Japan). Parin, 1960: 218-220 (synonymy, description; the northwestem Pacific Ocean). Kovalevskaya, 1964: 211-217 (early develop- mental stages). Parin & Besednov, 1966 (the Gulf of Tonkin). Munro, 1967: 119 (New Guinea). Kotthaus, 1969: 8-9 (western part of the Indian Ocean). Mai & Bui, 1978: 251-253 (Vietnam). Kovalevskaya, 1980: 219-220, 255-258 (early stages, distribution in the Pacific and indian oceans). Yoshino, 1984: 80 (Japan). Heemstra & Parin, 1986 (southern Africa). Chen, 1987: 48-49 (early developmental stages; the northwestern JOURNAL OF ICHTHYOLOGY Vol. 40 Suppl. 1 2000A REVIEW OF THE FLYING FISH GENUS EXOCOETUS (EXOCOETIDAB) sas HI 20) ae. BS Seog Moree gs os 8 13 +40 q ° a oa] 9° 2 silo 20 40 60 80 100 120 140 160 180 20 40 60 80 100 120 140 160 180 SL,mm Fig. 7. Size variation of some morphometric characters of Exocoerusvolitans (% SL) from the Alantic Ocean (thombs). Indian ‘Ocean (squares) and Pacific Ocean crosses) a) Head length, cb the greatest body depth, (c) width of inerorbital space, (head dpi, () veneal in length (pectoral fn lenath JOURNAL OF ICHTHYOLOGY Vol. 40 Suppl. 12000