BULLETIN OF MARINE SCIENCE, 26(1): 72-98, 1976
INDO-WEST PACIFIC HALFBEAKS (HEMIRAMPHIDAE) OF THE GENUS RHYNCHORHAMPHUS WITH DESCRIPTIONS OF TWO NEW SPECIES
Bruce B. Collette
ABSTRACT The genus Rhynchorhamphus Fowler, 1928 differs from other genera of Hemiram- phidae in having a fimbriate nasal papilla, a large domed upper jaw the length of which is equal to or greater than the upper jaw width, many gill rakers (47-78 on the first arch and 40-68 on the second arch), and two dorsally-directed branches of the lateral line. Loligorhamphus Whitley, 1931, is placed in the synonymy of Rhynchorlwmpl/lls. Four species are recognized using characters such as the relative length and width of the upper jaw, numbers of gill rakers on the first and second gill arches, and numbers of fin rays in the dorsal and anal fins. R. georgii (Valenciennes, 1846) is the most widely distributed species, occurring from the Persian Gulf to the East Indies, Australia, New Guinea, the Philippine Islands, and the South China Sea. Six nominal species are placed in the synonymy of R. georgii: Hemiramphus russeli Valenciennes; He. leucop/ems Val- enciennes; He. eclancheri Valenciennes; He. plumatlls Blyth; He. can/ori Bleeker; and Loligorhamphus normani Whitley. R. georgii and R. arabicus Parin and Shcherbachev, 1972 are redescribed. Two new species are described: R. maiabariclis from southern India and Ceylon and R. naga from the Gulf of Thailand and East Indies.
The purpose of this paper is to diagnose characters studied were numbers of rays in the genus Rhynchorhamphus, describe two the dorsal, anal, and pectoral fins (all rays unrecognized species, and redescribe the two counted separately); numbers of gill rakers known species. Although eight nominal spe- on the first and second gill arches; numbers cies attributable to the genus Rhynchorham- of precaudal, caudal, and total vertebrae; phus Fowler have been described, only one and numbers of predorsal scales. Radio- valid species has been recognized until re- graphs were used to obtain vertebral num- cently when Parin and Shcherbachev (1972) bers and to check numbers of dorsal and anal described R. arabicus from Aden. The other fin rays. Morphometric characters included: six nominal species are synonyms of the lower jaw length, head length, distance from widely distributed Rhynchorhamphus georgii pectoral fin origin to pelvic fin origin (Pr (Valenciennes). While examining material P~), distance from pelvic fin origin to cau- of Rhynchorhamphus in various museums, dal fin base (P~-C), snout length, orbit it become clear that two unrecognized spe- length, bony interorbital distance, length of cies exist: one from the Malabar and Coro- upper jaw, and width of upper jaw. Talwar mandel coasts of southern India and the (1964b) did not find any morphometric dif- waters of Ceylon (now Sri Lanka); the other ferences between males and females in four in the Gulf of Thailand and Java Sea. The characters (head plus lower jaw length; head southern India-Ceylon species is the basis of length; PrP~; and pectoral fin length) in a an important fishery in the Gulf of Mannar sample of R. malabaricus from Rameswaram and Palk Bay between Ceylon and India. Island in the Gulf of Mannar; therefore, males and females were not treated sepa- METHODS AND MATERIALS rately in my study. The shape of the pre- The methods used are similar to those in orbital canal and location and number of previous papers in this continuing series (see pores in it were examined on representative Collette, 1974a and b) of studies on the sys- specimens from each collection. Squamation tematics of the Synentognathi. Meristic of the upper jaw was studied in a sample of
72 COLLETTE: REVIEW OF THE GENUS RHYNCHORHAMPHUS 73 specimens on which scales were still present. Teeth on the upper and lower jaws and on the upper and lower pharyngeal bones were examined and the pharyngeal teeth are fig- a ured. A developmental series is included to show the increase in fimbriation of the nasal papilla with fish size. Specimens of all the species were cleared and stained to compare selected osteological characters with other genera of Synentognathi (see Collette, ]966). Figure 1. Increase in complexity of the fimbriate Abbreviations used for the institutions nasal papilla with size in Rhynchorlzamphus naga. cited herein are as follows: Left nasal fossa, one fold drawn to right. USNM 214083 Gulf of Thailand. a. 64 mm SL, b. 85.1 AMNH-American Museum of Natural History, mm, c. ]04 mm, d. 144 mm. New York. AMS-Australian Museum, Sydney. ANSP-Academy of Natural Sciences, Philadel- Rhynchorhamphus Fowler phia. Hemiramphus (Rhynchorhamphus) Fowler, 1928: BMNH-British Museum (Natural History), Lon- 75 (type-species Hemiramphus georgii Val- don. enciennes, by original designation). BPBM-Bernice P. Bishop Museum, Honolulu. Loligorlwmphus Whitley, 1931: 105 (type-spe- CAS-California Academy of Sciences, San Fran- cies Loligorhamphus normani Whitley [= R. cisco. geOl'gii], by original designation). CMFRI-Central Marine Fisheries Research In- stitute, Mandapam Camp, India. Diagnosis.-Rhynchorhamphus differs from CSIRO-C.S.I.R.O., Division of Fisheries and Oceanography, Cronulla, Australia. all other genera of Hemiramphidae in hav- FBQ-Fisheries Branch, Department of Primary ing a fimbriate nasal papilla (Fig. 1), a large Industries, Brisbane, Australia. domed upper jaw, and many gill rakers (47- FRSHK-Fisheries Research Station, Aberdeen, 78 on the first arch and 40-68 on the second Hong Kong. arch, Tables 1-2). Rhynchorhamphus dif- GVF-George Vanderbilt Foundation, specimens at CAS. fers from the other genera, except Hemi- KUB-Kasetsart University, Bangkok. ramphus, in having two dorsally directed MCZ-Museum of Comparative Zoology, Harvard branches of the lateral line instead of one University, Cambridge, Mass. (none in Dermogenys). The additional MFLB-Marine Fisheries Laboratory, Bangkok. MSNG-Museo Storia Naturale, Genoa. branch parallels and is anterior to the branch MNHN-Museum National d'Histoire Naturelle, to the pectoral fin base, slightly posterior to Paris. the cleithrum (see illustrations of each spe- NHMV-Naturhistorisches Museum, Vienna. cies). Rhynchorhamphus differs from Hemi- NMC-National Museum of Canada, Ottawa. ramphus in having scales on the upper jaw RMNH-Rijksmuseum van Natuurlijke Historie, and having the swimbladder entire, as in Leiden. Hyporhamphus, not divided into separate SIO-Scripps Institution of Oceanography, La Jolla, Calif. chambers. As shown by Collette (1974a: SU-Stanford University, specimens now at CAS. figs. 1-2), Rhynchorhamphus belongs to the UMMZ-University of Michigan Museum of group of halfbeaks that lack a posterior Zoology, Ann Arbor. branch to the preorbital canal. This group USNM-National Museum of Natural History, includes Arrhamphus and one species group Washington, D. C. of Hyporhamphus. The canal is a long nar- VMM-Vanderbilt Marine Museum, Centerport, row tube that has one or two central pores New York. ZMK-Zoological Museum, University of Copen- in addition to the openings at the ends of the hagen. dorsal and ventral arms of the canal (Fig. 2). ZSI-Zoological Survey of India, Calcutta. The upper jaw length is about equal to or 74 BULLETIN OF MARINE SCIENCE, VOL. 26, NO. I, 1976
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Dorsal views (diagrammatic) of upper jaws of the four species of Rhynchorhamphus. Up- per specimen of each pair shows a low number of upper jaw scales, lower specimen a high number. a. R. arabicus, Aden, MCZ 41922, 179 mm SL. b. R. arabicus, Aden, USNM 214092, 182 mm SL. c. R. malabaricus holotype, Ceylon, USNM 214081, 167 mm SL. d. R. malabaricus, Ceylon, USNM 214937, 160 mm SL. e. R. naga, Gulf of Thailand, USNM 214083, 152 mm SL. f. R. naga, Singa- pore, ZMK 341747, 169 mm SL. g. R. georgii, Philippine Is., USNM 138678,205 mm SL. h. R. georgii, N. Borneo, USNM 138688, 161 mm SL. 11 + 23 = 34) clearly do not apply to R. and uniformity of nomenclature. In this georgii. Examination of the two specimens case, the choice is between acutus Gunther in the Agassiz Museum (see Collette, and georgii Valenciennes. Few authors have 1974b) from the Kingsmill Islands (MCZ misidentified R. georgii although a number 8777) on which Fowler based his descrip- of authors have misidentified other species tion of georgii showed that they are Hypo- as R. georgii. It is in the best interests of rhamphus a. acutus (Gunther). In fact, re- nomenclatural stability that the nominal spe- examination of most of Fowler's central cies actually designated, i.e., He. georgii, Pacific material shows that he almost consis- stand as the type-species, thereby preserving tently misidentified Hy. acutus as R. georgii usual and current usage of the generic name (see misidentifications under R. georgil). He Rhynchorhamphus. applied the name Hy. acutus to other central Pacific species of Hyporhamphus such as Parasites.--Copepods and isopods were re- Hy. affinis (see Collette, 1974b). Previ- moved from the study material. Four species ously, however, Fowler (1927) correctly of copepods were identified by Dr. Roger F. identified a fine series of georgii from the Cressey (USNM): Bomolochidae: Bomo- Philippines (ANSP 55963-6). lochus bellones Burmeister and B. sinensis According to article 70 of the Interna- (Cressey) ; Anthosomatidae: Lernanthropus tional Code of Zoological Nomenclature, if belones Kr¢yer; and Lernaeidae: Penicu/us a zoologist considers that a type-species was sp. A comprehensive review of the parasitic misidentified, he is to refer the case to the copepod fauna of the Hemiramphidae is Commission to designate as type-species planned by Cressey and Collette; a similar whichever species will best serve stability study on the Belonidae has been published, 80 BULLETIN OF MARINE SCIENCE, VOL. 26, NO.1, 1976 Table 5. Morphometric comparison of the four species of Rhynchorhamphus R. arab/cus R. malabar/crlS Character Min. Max. x N SO Min. Max. x N SO Standard length (mm) 158 190 177.3 4 13.65 147 226 180.4 54 21.16 0/00 SL Lower jaw length 327 374 353.7 4 23.04 283 368 330.6 44 20.78 Head length 222 238 228.9 4 7.52 209 238 223.8 54 5.78 Pectoral-Pelvic distance 431 462 445.4 4 12.93 421 464 438.8 36 9.94 Pelvic-Caudal base 324 339 330.1 4 7.25 322 347 333.7 36 6.65 Snout 85 91 87.1 4 2.65 77 93 85.1 54 3.46 Orbit 51 56 53.5 4 3.25 47 60 53.4 54 3.20 Interorbit 55 63 59.2 4 3.15 54 61 57.2 54 1.90 Upper jaw length 48 54 51.1 4 2.53 45 59 50.9 54 2.81 Upper jaw width 52 58 53.9 4 3.04 44 54 49.3 54 2.20 0/00 Head L Snout length 370 390 380.8 4 8.38 358 415 381.0 65 12.04 Orbit 227 243 233.6 4 7.41 213 266 238.1 65 11.40 Interorbit 249 272 258.7 4 11.26 240 278 255.7 65 8.11 Upper jaw length 210 234 223.5 4 11.97 203 256 229.0 65 11.52 Upper jaw width 218 252 235.6 4 13.75 195 241 220.4 65 9.48 Proportions Lower jaw length/head length 1.41 1.68 1.55 4 0.11 1.23 1.63 1.48 52 0.08 Upper jaw length/width 0.92 0.99 0.95 4 0.03 0.90 1.17 1.04 65 0.06 lnterorbit/ orbit 1.06 1.15 1.11 4 0.04 0.93 1.24 1.08 65 0.07 see Cressey and Collette (1970). Cymo- nithan (1957) described Oligapta oligapta thoid isopods were given to Dr. Thomas E. (Axinidae) from the gills of a specimen Bowman (USNM) for subsequent study. from Mandapam Camp. Tripathi (1959) There appear to be no previous records of described Tribaculocauda georgii (Dactylo- copepods from speCIes of Rhynchorham- gyridae) from Port Canning, Matla R.; In- phus; although several parasitological papers docotyle hemiramphi (Discocotylidae) from use the unverified name Hemiramphus Puri, Orissa, Bay of Bengal; and Axine hemi- georgii for hosts of monogenetic trematodes rhamphae (Axinidae) from Puri and from the Philippines or India. Vazquez- Hooghly. Price (1962) transferred Indo- Colet and Africa (1938, 1939, 1940) re- cotyle Tripathi to the Axinidae. Young ported on the metacercariae of several spe- (1968: 415) noted that the genera ofAxin- cies of heterophyid trematodes found in the idae are found on three related families of flesh of several species of Philippine fishes, fishes: Hemiramphidae, Exocoetidae, and including He. georgii. There is a photograph Belonidae. in one of the papers (1939: pI. 1, fig. 2) The copepods collected during this study which appears to be of a species of Hypo- are listed by host species in the following rhamphus rather than R. georgii. paragraphs. All are new records for Rhyn- Two papers describe species of mono- chorhamphus but three have been reported genetic trematodes from Indian halfbeaks from species of Belonidae by Cressey and identified as Hemirhamphus georgii. Un- Collette (1970): Bomolochus bellones from COLLE1TE: REVIEW OF THE GENUS RHYNCHORHAMPHUS 81 Table 5. (Continued) R. naga R. georgii Character Min. Max. i< N SD Min. Max. i< N SD Standard length (mm) 62 177 130.9 78 30.10 85 231 157.9 70 36.71 0/00 SL Lower jaw length 318 721 476.4 53 90.38 301 605 403.9 35 70.10 Head length 161 245 229.7 71 10.36 211 249 228.9 64 7.86 Pectoral-Pelvic distance 309 462 442.3 76 17.70 400 472 448.7 61 12.15 Pelvic-Caudal base 230 346 322.1 76 13.81 301 339 320.0 61 9.56 Snout 65 101 90.9 62 5.18 83 106 94.2 41 5.18 Orbit 40 65 56.7 74 3.54 43 62 53.2 64 4.80 Interorbit 39 62 55.2 74 3.28 43 65 58.2 59 3.71 Upper jaw length 40 69 57.0 77 4.59 46 75 63.0 66 6.07 Upper jaw width 28 54 44.6 77 3.41 41 57 48.8 66 3.91 0/00 Head L Snout length 351 424 395.8 56 12.77 365 440 412.2 40 19.34 Orbit 216 281 246.6 68 12.61 193 276 231.6 62 19.55 Interorbit 217 263 240.1 68 10.01 195 294 254.8 57 14.39 Upper jaw length 209 307 248.4 71 15.80 216 323 275.9 63 23.27 Upper jaw width 160 237 194.6 71 12.68 182 246 213.9 63 15.22 Proportions Lower jaw length/head length 1.52 3.29 2.08 53 0.40 1.36 2.64 1.75 36 0.30 Upper jaw length/width 1.04 1.50 1.28 78 0.08 1.04 1.50 1.30 67 0.09 Interorbit/orbit 0.84 1.21 0.98 75 0.06 0.91 1.38 1.11 60 0.11 ------ 16 species; B. sinensis from two and Lern- lones from Batavia, the Java Sea, the East anthropus be/ones from 11; and two have Indies, and the Gulf of Thailand (12 females been reported from Australian Hemiram- and two males from under the gill covers, on phidae by Collette (1974a): B. bellones the gill arches, and under the lower jaw oral from three species of Hyporhamphus and L. valve of RMNH 6953, 12205, 12141a, be/ones from two species of Hyporhamphus 12144, 12197, and 92138; UMMZ 171809; and Arrhamphus sclero/epis. ZMK 341744); B. sinensis from the East In- Rhynchorhamphus arabicus: Lernanthro- dies and Java (11 females under the gill pus sp. from the Gulf of Aden (male from covers and oral valves and on the gill arches the gill arch of MCZ 41922). of RMNH 6953, 12201, and 12204); Lern- Rhynchorhamphus ma/abaricus: Bomo- anthropus be/ones from the East Indies /ochus bellones from Ceylon and the Mala- (four females from the gill arches of RMNH bar Coast of India (four females under the 6953 and 12201); Lernanthropus sp. from gill covers and on the gill arches of USNM the Gulf of Thailand (male, probably also 213566 and BMNH 1935.7.2.1-7); Bomo- L. be/ones from the gill arch of USNM /ochus sp. from the Coromandel Coast of In- 206077); and Peniculus sp. from the East dia (two juvenile females too small to iden- Indies (female attached to left pectoral fin tify but probably also B. bellones from under of RMNH 12201). the lower jaw oral valve of USNM 21836). Rhynchorhamphus georgii: Bomolochus Rhynchorhamphus naga: Bom%chus be/- bel/ones from the East Indies (four females 82 BULLETIN OF MARINE SCIENCE, VOL. 26, NO.1, 1976 12.0 o R. malabaricus 11.0 o ~ * R. arabicus 000 10.0 o 000 0 o 0 0 o 9.0 <1Jb a 0 ~o o .00 00 E 0 !.8.0 " 3.0 2.0 2.0 3.0 4.0 5.0 6.0 7.0 8.0 9.0 10.0 11.0 12.0 13.0 14.0 15.0 Upper law length (mm) Figure 4. Relationship of upper jaw length to upper jaw width in the four species of Rhynchorham- phl/s. and two males from under the oral valves branches of the lateral line ventral to the and gill covers of RMNH 6953 and 92138); pectoral fin origin, Rhynchorhamphus ap- B. sinensis from Batavia (one female and pears most closely related to Hemiramphus. one male from RMNH 12202); Lernanthro- Specialized characters such as the fimbriate pus belones from the East Indies (one fe- nasal papilla, large number of gill rakers, male from the gill arch of RMNH 92138); and large domed upper jaw indicate that and Lernanthropus sp. from Java and the Rhynchorhamphus is a specialized deriva- Gulf of Thailand (two males, also probably tive of Hemiramphus. There are two species L. belones, from the gill arches of RMNH groups in the genus. R. arabicus and R. 12200 and UMMZ 101266). malabaricus have more gill rakers, fewer Range.-Rhynchorhamphus is a tropical dorsal and anal fin rays, fewer vertebrae, a Indo-West Pacific genus that occurs close to shorter and wider upper jaw, and smaller shore (Fig. 7). Its distribution is comple- teeth on the pharyngeal bones than do R. mentary to that of the Central Pacific Hypo- georgii and R. naga. rhamphus acutus, a species which occurs Rhynchorhamphus arabicus south or west of the andesite line, which Parin and Shcherbachev separates continental- from non-continental- Figure 8a type rocks (see Collette, 1974b: fig. 2). Rhynchorhamphus arabicus Parin and Shcher- Relationships.-Based on the shared special- hachey, 1972: 569-571 (original description; ization of possessing two dorsally··directed southern Yemen). Collette, 1974a: 91 (valid- COLLETTE: REVIEW OF THE GENUS RHYNCHORHAMPHUS 83 a n c d Figure 5. Upper and lower pharyngeal bones in four species of Rhynchorhamphus. a. R. arabicus, Aden, USNM 214092, 182 mm SL. b. R. malabaricus, USNM 214937, Ceylon, 173 mm SL. c. R. naga, USNM 206080, Gulf of Thailand, 158 mm SL. d. R. georgii, USNM 214093, Gulf of Thailand, 170 mmSL. 84 BULLETIN OF MARINE SCIENCE, VOL. 26, NO. I, ]976 malabaricus which has 57-71 (x 64.7) gill rakers but only six out of 66 specimens that I examined overlapped and none of the 355 reported by Talwar (1964b) did so. R. arabicus is similar to R. malabaricus in aver- aging fewer dorsal and anal rays than R. georgii and R. naga (Table 3). Both R. arabicus and R. malabaricus average fewer vertebrae (usually 54 or 55) than R. naga and R. georgii, which usually have 55-57 (Table 4). R. arabicus has the widest upper jaw in the genus (Table 5, Fig. 5), 52-58 thousandths of standard length (x 53.9) compared to 44-54 (x 49.3) in R. malabar- icus, which has the next widest upper jaw. R. arabicus and R. malabaricus have smaller pharyngeal teeth (Fig. 5). Description.-Meristic data are presented in Tables 1-4. Pectoral fin rays 11. Predorsal scales 40 or 41 in my material, 39 reported by Parin and Shcherbachev (1972). Mor- phometric data are summarized in Table 5. Maximum known size.-Female 190 mm SL (USNM 214092, Aden). Range.-Known only from the Gulf of Aden (Fig. 6). Parin and Shcherbachev (1972) provisionally assumed that the range might extend to the Red Sea and Persian Gulf based on records of Borodin (1932) and Blegvad (1944) of He. georgii. Borodin's record is based on a misdentified specimen (VMM 863) of a species of Hyporhamphus similar to Hy. dussumieri (Valenciennes). Two of Blegvad's specimens (ZMK eN 5-6) are R. georgii (the third is a specimen of Figure 6. Otolith (sagitta) of Rhynchorham- Hyporhamphus sp.) and two other collec- phlls malabaricl/s, Gulf of Mannar, USNM 214082, 220 mm SL. Outer surface of left sagitta on left, tions from the Persian Gulf (FMNH 51254 ) inner surface on right. and Gulf of Oman (MNHN 65-558) are also R. georgii. Material Examined.-Aden: MCZ 41922 (2, 158- ity of species confirmed; additional material ]79) and USNM 2]4092 (2, 182-190); Aden Har- from Aden). bor; R. W. Risebrough; May 22, 1958. Diagnosis: R. arabicus has more gill rakers than do any of the other species of Rhyncho- Rhynchorhamphus malabaricus rhamphus: 69-78 (x 73.2) on the first arch new species and 63-68 (x 65.3) on the second arch Figure 8b (Tables 1-2). The range of number of gill Hemirhamphlls georgii (not of Valenciennes). rakers on the first arch overlaps with R. Day, 1878: 515-516 (description, in part; COLLETTE: REVIEW OF THE GENUS RHYNCHORHAMI'HUS 85 '0' 90' zli' zoo • MARl .•••"''' : I~ .':;"1""" I .:r_ j:."'.'~'".,': .:~.~~~. ~,.. " IS. o' . O' \ c~g?S :; ~ '-~ . '~~~~~:""~. b'.,' ~ ___~ r-R~-y-nC-h-Or-h-O-m-Ph-U-S--' zoo @ arabicus @ maloboricus • nago * georgii I 20' ..,. o' 80' 100" I O' Figure 7. Distribution of the four species of Rl1yncllOrl1ampl1us based on specimens personally exam- ined. India; not pI. 120, fig. 2 which is He. mar- overlapping with R. georgii at the lower end ginatus). Day, 1889: (description in part, of the range (52-67, x 59.4) and with R. after Day, 1878). Devanesan, 1937: 2175- 2181 (early life history, figs. 2-4; important arabicus at the upper end of the range (69- fishery in Palk Strait between India and Cey- 78, x 73.2, Table 1). R. malabaricus is lon). Chidambaram and Menon, 1948: 759- completely separated from R. arabicus in 762 (development, figs. 6-9; western India). number of gill rakers on the second arch Devanesan and Chidambaram, 1953: 25-26 (52-62 vs. 63-68, Table 2) but overlaps (common names, bionomics; important fish- ery in Palk Bay). Krishnamurthi, 1957: 236, broadly with R. georgii (45-63, x 55.2). R. 239, 251 (9th-10th most important fishery, malabaricus is similar to R. arabicus in hav- Rameswaram Is., Gulf of Mannar). ing fewer dorsal and anal fin rays than R. Hemiramplllls geargii (not of Valenciennes). georgii and R. naga (Table 3). Most R. Chacko, 1949: 87 (plankton feeder; Gulf of malabaricus can be distinguished from most Mannm"). De Silva, 1956: 50 (description; Ceylon). R. georgii by having the sum of dorsal and R!JYllc!Jarl1amplllls geargii (not of Valenciennes). anal fin rays equal to 28 or less (25-30, Munro, 1955: 74 (description, in part; Cey- x 27.2) compared to usually 29 or more Ion; pI. 13, fig. 206, after Day, 1878, pI. 120, (26-32, x 29.2). R. malabaricus and R. fig. 2). arabicus average fewer vertebrae (54 or 55) Hyporl1amplllls geargii (not of Valenciennes). than R. naga and R. georgii (55-57, Table Talwar, 1964a: 3 (stomach contents of 1042 specimens; Gulf of Mannar and Palk Bay). 4). R. malabaricus is also similar to R. Talwar, 1964b: 168-196 (racial analysis, biol- arabicus in having a relatively wide upper ogy; Gu]f of MannaI' and Palk Bay), Tal- jaw (Fig. 5); length divided by width 0.90- war, 1968: 67-68 (54 myotomes). Quasim, 1.17, x 1.04 in R. malabaricus compared to 1973: 18 (feeding habits, after Talwar). 0.92-0.99, x 0.95 in R. arabicus (Table 5). Diagnosis: R. malabaricus has many gill R. malabaricus has the shortest lower jaw rakers on the first arch (57-71, x 64.7), (283-368 thousandths of SL, x 331), differing 86 BULLETIN OF MARINE SCIENCE, VOL. 26. NO.1, 1976 Figure 8. a. RhYllchorhamphus arabicus, Aden, MCZ 41922, 190 mm SL. b. R. malabaricus n. sp. holotype, Ceylon, USNM 214081, 167 mm SL. from R. georgii and R. naga (301-721, quency data to determine age because x 404 and 476 respectively), but broadly neither scales nor otoliths were useful. Three overlapping with R. arabicus (327-374, year classes were present in March, with x 354). R. malabaricus and R. arabicus modal lengths of 135, 175, and 235 mm FL have smaller pharyngeal teeth (Fig. 5) than (123, 161, and 218 mm SL). Talwar also do R. naga and R. georgii. ran regressions of weight (W) against fork length (L) for a sample of males and one of Description: Meristic data are presented females. He presented the following equa- in Tables 1-4. Pectoral fin rays 11-12. Pre- tions, for females: dorsal scales 37-41, usually 39 or 40 in my material; 35-41, usually 38 or 39 according log W = 2.8000 log L - 4.8156 to Talwar (1964b). Morphometric data are and for males: summarized in Table 5. log W = 1.6667 log L - 2.2437. Maximum known size: In his study on the food of R. malabaricus (as Hyporhamphus Types: Holotype USNM 214081; Ceylon, georgii) from the Gulf of Mannar and Palk Myliddy near Kankesanturai; T. Roberts; 18 Bay, Talwar (1964a) examined 1042 speci- March 1970; male, 167 mm. D 14; A 13; Pl mens that ranged in size from 155-275 mm 11-11; RGRl 19 + 47 = 66; RGR2 10 + 50 fork length. Converting fork length to stan- = 60; predorsal scales 38; vertebrae 38 + 17 dard length, based on a sample of the material = 55; preorbital canal shown in Fig. 2b; up- that I examined [SL = (FL - 6.26) /1.05J per jaw length 8.6 mm, width 8.0 mm. Para- shows that Talwar had specimens up to about types, all the other specimens listed under 256 mm SL. The largest specimen examined material examined. in my study was a 226 mm SL female (244 mm FL) from the same area (USNM Etymology: Named malabaricus because the 214082, Pambam); 12 specimens 180 mm first specimens of this species that I exam- SL or larger were females. ined (BMNH 1935.7.2.1-7) were from the Talwar (1964b) analyzed length fre- Malabar Coast of India. COLLETTE: REVIEW OF THE GENUS RHYNCHORHAMPHUS 87 Biology: Talwar (1964a and b) has pre- Table 1 (gill rakers on the first gill arch), sented one of the few reasonably detailed Table 3 (dorsal and anal rays), and Table studies on the biology of a species of half- 4 (total number of vertebrae). Counts from beak. His data on R. malabaricus (as Hy- my Coromandel sample are very similar to [Jorhamphus georgii) from the Gulf of Man- Talwar's data. nar and Palk Bay are summarized here. Based on examination of the gut contents of Range.-R. malabaricus is confined to the 1042 specimens ] 55-275 mm fork length southern third of the Indian peninsula (Fig. (]42-256 mm SL), Talwar (1964a) found 7), along the Malabar Coast of the eastern that the main diet of the maturing fish was Arabian Sea, the Coromandel Coast of the the sea grass Cymodocea and green algae. western Bay of Bengal, and in the Gulf of Several genera of diatoms and also the re- Mannar and Palk Bay, between Ceylon and mains of polychaetes were present. During India. R. georgii also occurs throughout the the spawning season, the diet changed to range of R. malabaricus but appears to be 80% pteropods (Creseis acicula) plus bi- much less abundant, judging from museum valve larvae, copepods, decapod larvae, and material and Talwar's study of the halfbeaks diatoms. Spent fish fed largely on copepods of the Gulf of Mannar and Palk Bay (Tal- (primari]y Acartia erythraea) and foraminif- war, 1964a and b). era. Three other species of halfbeaks in the nrea-Hemiramphus far, He. marginatus, Material Examined.--India, Malabar Coast: BMNH 1935.7.2.1-7 (12, 162-179); S. Malabar; Devane- and Hy. quoyi-fed largely on Cymodocea sen. nnd green algae all year. Ceylon: 29 specimens (152-215 mm SL) from From macroscopic examination of the 6 collections. USNM 213563 (13, 166-215) and ovaries over several years, Talwar (1964b) CAS 33297 (2, 178-180); Mannar 1., 1 mi. west of Pesalai; M/V BERuwALLA, beach seine; 17 concluded that R. malabaricus spawns from March 1969; F. 1. Schwartz 69-52. USNM 213566 mid Mnrch to mid April. At maturity, the (I, 195); Mannar 1., near Pesalai, Colombo mar- eggs vary from 1.2 to 1.6 mm in diameter ket; 4 March 1969; F. J. Schwartz 69-26. USNM (mode 1.4 mm). The smallest ripe male in 213561 (1, 156); Hikkaduwa; 13 Jan. 1970; C. C. TaLwar's study was 153 mm fork length Koenig 69-54. USNM 214937 (7, 160-173); My- liddy near Kankesanturai, N. tip of Ceylon; 18 ( 139 mm SL) and the smallest ripe female March 1970; T. Roberts. MCZ 50748 (1, 201); 149 mm FL (136 mm SL). All individuals Mannar fish market; 8 Feb. 1970; C. C. Koenig over 149 mm FL (136 mm SL) were ma- 69-97. USNM 213837 (4, 152-177); Kalupittiya ture. Talwar (1964b) found that the rela- fish. market; 25 Jan. 1970; C. C. Koenig 69-80. tionships of numbers of ripe eggs (F) to India, Coromandel Coast: 26 specimens (147- 226 mm SL) from 4 collections. USNM 214082 fork length (L) could be expressed as: (15, 149-226) and ZSI F 7222/2 (1, 199); Gulf of Mannar, Pam bam, Mandapam Camp; 7 Feb. log F = 3.8140 log L- 5.5047 1975; M. Devaraj. CMFRI un cat. (1, 180); Pam- bam; 16 April 1958. USNM 213834 (I, 161); His fig. 4 shows a parabolic curve increasing Pondichery State, Vaithikuppam fishing village, from about 1050 eggs in a female 180 mm purchased in Pondichery market; 22 Sept. 1966; FL to 4550 eggs at 240 mm FL. F. H. Berry 66-22. USNM 213836 (8, 147-178); Madras State, Enore area near Madras, purchased from Madras State Fish. Dept. Mar. BioI. Sta.; 14 Geographic variation.-R. malabaricus was Sept. 1966; F. H. Berry 66-62. divided into three populations for compar- ison of meristic characters: Malabar Coast, Rhynchorhamphus naga new species Coromandel Coast, and Ceylon (Tables 1-4). Figure 9a All populations have similar ranges and means. Data from Talwar (1964b) for a H emiramphus geOl·gU. Cantor, 1849: 1230-1231 large snmple from the Gulf of Mannar and (description, in part; Penang, Malaysia [BMNH unreg.]). Bleeker, 1852: 4 (Java), Palk Bay are included, for comparisons, in 10 (in key). 88 BULLETIN OF MARINE SCIENCE, VOL. 26, NO, I, 1976 a b Figure 9. a. Rhynchorhamphus naga n. sp., holotype, Gulf of Thailand, USNM 206077, 165 mm SL. b. R. georgii, North Borneo, USNM 138688, 161 mm SL. Hemirhamphus CantO/'i. Bleeker, 1866: 145- R. arabicus (327-374, x 354), and R. georgii 146 (original description, in part; East In- (301-605, x 404). This is also shown by dies). Bleeker, 1871: 53 (description, in part; pI. 252, fig. 2; East Indies). the ratio lower jaw length/head length which Hemirhamphus georgii. GUnther, 1866: 264 is greater in naga, 1.52-3.29 (x 2.08) com- (description, in part; Penang [BMNH unreg.]). pared to 1.23-2.64 (x's 1.48-1.75) in the Weber and de Beaufort, 1922: 147-149 (de- other three species. R. naga has the narrow- scription, in part; East Indies). Hemirhamphus cantoris. GUnther, 1866: 264- est upper jaw, 28-54 thousandths of standard 265 (description, in part; China seas [BMNH length (x 44.6) compared to 41-58 (x's 48.8- unreg.]). 53.9) in the other three species; 160-237 Diagnosis.-R. naga has the fewest gill rak- thousandths of head length (x 194.6) com- ers of any of the species of Rhynchorham- pared to 182-252 (x's 213.9-235.6); and phus, 47-59 (x 52.4) on the first gill arch, also the narrowest interorbital distance, the 40-53 (x 45.1) on the second gill arch but ratio interorbital distance/orbit length equal overlaps with R. georgii in counts on both to 0.84-1.21 (xO.98) compared to 0.91- arches (Tables 1-2). R. naga is interme- 1.38 (x's 1.11-1.24). R. naga has the largest diate between R. arabicus and R. malabar- pharyngeal teeth in the genus (Fig. 5). icus on the one hand and R. georgii on the Description.-Meristic data are presented in other in numbers of dorsal and anal fin rays Tables 1-4. Pectoral fin rays 10- 12, usually (Table 3). R. naga usually has 14 or 15 11. Predorsal scales 37-43, usually 39 or dorsal and anal fin rays for a total of 28 or 40. Morphometric data are summarized in 29, broadly overlapping with R. malabaricus Table 5. at 28 and R. georgii at 29. R. naga has the longest lower jaw (Table 5) in the genus Maximum Known Size.-I77 mm SL (318-721 thousandths of SL, x 476) com- (MFLB uncal., Bangkok and BMNH unreg., pared to R. malabaricus (283-368, x 331), Penang, Malaysia). COLLETTE: REVIEW OF THE GENUS RHYNCHORHAMPHUS 89 Types.-Holotype USNM 214085; Gulf of material, most of which was collected on Thailand, 70 46' N, 1030 40' E; Naga Ex- the Naga Expedition. However, shore-based pedition; 22 Oct. 1959; female, 165 mm. D collecting by Karl Lagler and Supap Pun- 14; A 14; P1 11-11; RGR1 14 + 39 = 53; poka Monkolprasit in Prachuab Khiri Khan RGR~ 8 + 38 = 46; predorsal scales 39; ver- Province on the eastern side of the Isthmus tebrae 39 + 17 = 56; preorbital canal shown of Kra in the Gulf of Thailand resulted in in Fig. 2c. Paratypes are the 111 specimens large numbers of R. georgii (more than 84 from the Gulf of Thailand and South China specimens saved, 114-200 mm SL) and only Sea listed under material examined. The one specimen of R. naga. specimens from the Java Sea are not des- ignated paratypes because they are mostly Material Examined.-Gulf of Thailand-South China old specimens in relatively poor condition. Sea: 112 specimens (62.4-177 mm SL), from 26 The collection from Borneo is not included collections. USNM 214084 (15, 91.2-161) and MCZ 50749 (2, 144-160); Naga 59-61; sta. 60- as paratypes because they come from the 427, 1.4 mi. from Songkhla, 7-15 N, 100-35 E; 18 limits of the range. Nov. 1960. CAS 32348 (3, 63.7-164); Naga 60- 203; reg. 2071, 9-57-54 N, 103-33-42 E; sta. 60- Etymology.-Named after the Naga Expedi- 55c; T. Matsui; 16 Feb. 1960. CAS 32350 (2, 138- tion to the Gulf of Thailand and South China 141); Naga 60-380; sm locality 60-699, reg. 2397; Sea during which most of the fresh material R/V STRANGER;T. Matsui et al.; 14-15 Aug. 1960. CAS 32363 (3, 141-167); sm locality S-I, sta. 3; was collected from 1959-1961. Swamg and Kosol; 20 Oct. 1959. CAS 32351 (5, 70-150); Naga 60-381; reg. 2398, Songkhla an- Geographic variation.-No important geo- chorage, 7-22-00 N, 100-43-30 E; R/V STRANGER; graphic variation was found in R. naga. T. Matsui et al.; 9 Aug. 1960. CAS 32357 (4, Morphometric comparisons were made be- 145-165); Naga 60-464; reg. 2739, sm locality 61- 156; R/V STRANGER;E. Brinton et al.; 12 Feb. tween a sample from the Gulf of Thailand 1961. CAS 32347 (2,62.4-167); Naga 60-27; reg. and one from the East Indies. No differ- 2038, off Ko Kra, 8-25-30 N, 100-44-40 E; R/V ences were found in ranges or means so the STRANGER;R. Bolin; 25 Jan. 1960. CAS 32360 (1, morphometric data for the two populations 68.8); Naga 60-512; reg. 2787, 11-59-30 N, 101- were combined for comparison with the 26-36 E; R/V STRANGER;21 Nov. 1960. CAS 32352 (I, 140); Naga 60-387; reg. 2404, 8-39-00 N, 100- other three species of Rhynchorhamphus 15-00 E; R/V STRANGER;T. Matsui; 6 Aug. 1960. (Table 5). Three populations were com- CAS 32356 (1,99.7); Naga 60-445; reg. 2720, 11- pared meristically, Gulf of Thailand, East 30-00 N, 99-47-12 E; R/V STRANGER;T. Matsui; Indies, and Borneo. The large samples from 12 Dec. 1960. CAS 32353 (1,155); Naga 60-415; reg. 2690, 7-42-05 N, 104-58-20 E; R/V STRANGER; the Gulf of Thailand and East Indies are T. Matsui; 6 Oct. 1960. CAS 32349 (1, 87.8); virtually identical (Tables 1-4). There are Naga 60-242; reg. 2259, 11-41-30 N, 100-12-10 E; slightly fewer gill rakers on the first and sec- R/V STRANGER;J. Faughn; 1 May 1960. CAS ond gill arches in the small sample of small 32361 (1, 136); Naga 60-541; reg. 2816, sm 59- specimens from Borneo than in the other two 62, 10-28 N, 102-40 E; R/V STRANGER;Swamg; 26 Oct. 1959. CAS 32354 (1,77.0); Naga 60-423; samples; however, this slight difference reg. 2698, 8-57 N, 102-53 E, 40 mi. from Paulo seems insignificant. Panjung; R/V STRANGER;T. Matsui; 14-15 Nov. 1960. CAS 32355 (1, 76.8); Naga 60-432; reg. Range.-R. naga occurs throughout the 2707, 11-37.1 N, 101-46.6 E, 38 mi. from Goh Gulf of Thailand south into the Java Sea, Chang; R/V STRANGER;T. Matsui; 22 Nov. 1960. USNM 214085 (1, 165), holotype and USNM north into the South China Sea and east as 206077 (8, 147-172); reg. 2815; 7-46 N, 103-40 E; far as Sandakan, Borneo (Fig. 7). There is Naga Exped.; 22 Oct. 1959. USNM 206080 (9, an early record of Cantor (1849) from Pin- 83.8-158); Naga 60-263; reg. 2280, 7-48-30 N, ang [sic], Malaysia, at the northern end of 105-08 E; 23 June 1960. USNM 214083 (26,64.0- the Straits of Malacca (BMNH unreg.). R. 163), BMNH 1975.7.2.1-2 (2, 113-148) and ZSI F 7223/2 (1, 149); Naga 60-426; 7-15 N, 100-35 georgii also occurs throughout this area but E; 17 Nov. 1960. CAS 32346 (J, 155); reg. 1592, is much less abundant, judging from museum Mae Nam Chantaburi R.; Tha Chalaep Harbor; R. 90 BULLETIN OF MARINE SCIENCE, YOLo 26, NO. I, 1976 Rofen; 24 Dec. 1957. MFLB uncat. (1, 177); 1860: 25 (listed; Borneo). Bleeker, 1861a: Bangkok mkt.; 24 Dec. 1965. KUB uncat. (1, 32, 60 (listed; Singapore). Bleeker, 1861b: 172); Klong Wan; S. Punpoka. ZMK uncat. (1, 78 (listed; Penang). Day, 1865: 170 (de- 123); Thailand; K. Rordan. BMNH 1848.3.16.11 (1, scription after Cantor; seas of Malabar, In- dia). Day, ]873: 292 (description; seas of 132); "China Seas"; E. Belcher. CAS 32359 (1, India). Kner, 1887: 323 (description; Nov- 80.7); Naga 60-489; reg. 2764, South Viet Nam, ara Exped.; Madras [NHMV 5585]). Fowler, Nhatrang Bay, 0.5 mi. from Cau Da, 12-12 N, 109- 1927: 161 (Orion, Luzon, Philippine Is. 13 E; R/V STRANGER;20 Sept. 1960. CAS 32358 [ANSP 55963-66]). Fowler, 1929: 603 (1, 67.9); Naga 60-480; reg. 2755, South Viet (listed; Hong Kong markets). Fowler, 1932: Nam, Nhatrang Bay, off Cau Da, 12-12 N, 109-13 269-270 (description; fig. 14; Amoy and Hong E; R/V STRANGER;T. Matsui; 10-11 Sept. 1960. Kong). Herre, 1933a: 3 (Sandakan, British CAS 32362 (1, 95.8); Naga 60-545; reg. 2820, North Borneo [SU 27764]). Umali, 1936: 25 South Viet Nam, Nhatrang Bay; R/V STRANGER; (listed), 77 (description, fig. 39, size in Ma- Twesukdi; 23 Nov. 1959. CAS un cat. (17, 108- nila market). Herre, 1940: II (Mergui Arch., 141); Naga 60-411; SID locality 60-808; R/V [SU 37217]). Herre, 1944: 11-12 (synon- STRANGER;Arsini, Pinyo; 25 Sept. 1960. ymy; records from the Philippine Is., Borneo, China, Mergui Arch.). Marshall, 1951: 2 Java Sea: 156 specimens (56.9-177 mm SL) (Magnetic I., Qld. [FBQ 39], new Australian from 20 collections. BMN H unreg. (1, 177) ; record). Khalaf, 1961: 3: 60-61 (descrip- Malaysia, Penang; T. Cantor. SID 61-718 (4, 132- tion; Basra, N. Persian Gulf). Marshall, 170); Sunda Straits, 5-37 S, 106-02 E; Naga Ex- 1964: 100 (Qld., listed). Misra, 1962: 200 ped.; 22 March 1961. ZMK 341744-47 (4, 137- (description, in part; India and Pakistan). 169); NE of Singapore, 1-46 N, 104-25 E; R/V Hemiramphus Russeli Valenciennes in Clivier GALATHEA399; 21 June 1951. RMNH 12199 (1, and Valenciennes, 1846: 32-33 (original de- 168); Java Sea; P. Buitendijk; 1913. RMNH 12202 scription; Pondichery, India). Bleeker, 1852: (1, 128); Bay of Batavia; P. Buitendijk. RMNH 4 (Java), 10 (in key). Bleeker, 1853: 72 12205 (4, 97.7-137); East Indies; P. Buitendijk; (listed; Coromandel). Day, 1873: 292 (de- scription) . Jan. 1909. RMNH 12197 (4,81.1-107); Java Sea; Hemiramphlls lellcopterus Valenciennes in Cu- P. Bliitendijk. RMNH 12198 (3, 86.5-94.5); Java vier and Valenciennes, 1846: 48 (original de- Sea; P. Buitendijk; Aug. 1911. RMNH 12204 (10, scription; Bombay, India). Bleeker, 1853: 72 128-146); Java, Semarang; P. Buitendijk; Dec. (listed; Bombay). Day, 1873: 292 (descrip- 1912. RMNH 12201 (10,81.4-132); Pocloe-Weh; tion after Valenciennes, 1846). P. Buitendijk; Aug. 1913. RMNH 12206 (3, 75.3- H emiramphlls Eclancheri Valenciennes in Cu- 104); Java, Serna rang; P. Buitendijk; Nov. 1915. vier and Valenciennes, 1846: 51-52 (original RMNH 6953 (49, 69.1-168); East Indies; P. description; "Marquesas Is."). GUnther, 1866: Bleeker. RMNH 92138 (23, 119-176); East In- 268 (after Valenciennes). Jordan and Seale, dies; P. Bleeker. RMNH 12141a (3, 158-163); 1906: 207 (after Valenciennes). H emirhamphlls geOl·gii. Blyth, 1852: 50 (listed; Java, Batavia; P. Bleeker. RMNH 12144 (21, Ceylon). GUnther, 1866: 264 (description, 56.9-132); Java Sea; P. Buitendijk; Aug. 1905. in part; Bengal [BMNH uncat.]). GUnther, NHMV 21025-6 (2, 147-149); East Indies; P. 1909: 355-356 (description; not record of Bleeker; 1857. ZMK 341773-5 (3, 117-143); Java Pellegrin, 1898 from Guam). Weber and de Sea near Djakarta, 5-50 S, 106-59 E; R/V GALA- Beaufort, 1922: 147-149 (synonymy; descrip- THEA 459; 27-28 Aug. 1951. NMC 63-290 (1, tion, in part; East Indies). Chevey, 1934: 166); Singapore mkt.; C. C. Lindsey; Oct. 1962- 125 (listed after Tirant, 1885). Tchang, April 1963. MSNG 42572 (1,150); Java. UMMZ 1938: 343 (synonymy; Chinese records; fig. 3). Blegvad, 1944: 77-78 (synonymy, de- 171809 (8, 93.5-126); vicinity of Batavia; Hard- scription; Bushire, Persian Gulf [ZMK CN 5- enberg and C. L. Hubbs; H 29-42; 6 May 1929. 6]; fig. after Day, 1878). Munro, 1955: 74 Borneo: USNM 214086 (6, 95.5-111); Borneo, (description; in part; Ceylon; not pI. 13, fig. Sandakan Harbor; R/V TE VEGA 210; D. M. Co- 206). hen and W. P. Davis; 28 Jan. 1965. HemirlwlIlplllls pllllllatlls Blyth, 1859: 288 (original description; the Sandheads at the head of the Hooghly R. and Ceylon). Day, Rhynchorhamphus georgii (Valenciennes) 1869: 526 (description, apparently based on Figure 9b Blyth's Ceylon specimens). H elllirhamphus Cantori Bleeker, 1866: 145-146 Hernirarnphus Georgii Valenciennes in Cuvier (original description, in part; E. Indies). and Valenciennes, 1846: 37-39 (original de- Bleeker, 1871: 53 (description, in part, after scription; Bombay and Coromandel, India). Bleeker, 1866; not pI. 252, fig. 2). Bleeker, Cantor, 1849: 1230-1231 (description; Pe- 1873: 149 (listed; Amoy, China). Day, nang, Malaysia). Bleeker, 1853: 72 (listed; 1878: 514 (description; India; pI. 119, fig. 1). Bombay). Bleeker, 1858: 2 (listed; Borneo). Macleay, 1883: 593 (listed; New Guinea). Bleeker, 1859: 149 (listed; Bali). Bleeker, Tirant, 1885: 128-129 (description; Indo- COLLETTE: REVIEW OF THE GENUS RHYNCHORHAMPHUS 91 China). Day, ]889: 423-424 (description Misidentifications.-Hyporhamphl/s dl/ssllmieri (Va- after Day, ] 878; fig. 137 after Day, 1878, lenciennes) as Hemiramphl/s georgii. Pellegrin, pI. ]19, fig. 1). 1898: 228 (listed from Guam, Marianna Is. based Hemirilamplllls cantoris. Gunther, 1866: 264- on MNHN 89-210). Gunther (1909: 355-356) 265 (description, in part; Amoy, China cited Pellegrin's record. [BMNH 1860.7.20.16-17] and East Indies Hyporlwmphl/s picarti (Valenciennes) as He. [BMNH 1866.5.2.16]). georgii. TillieI', 1902: 292, 295, 299, 315, 316, Hemirampl/lls RI/sselli. Day, 1873: 292 (de- 318 (listed as a Red Sea species that had traversed scription; Coromandel Coast). the Suez Canal into the Mediterranean, one speci- Hemirampill/s pillmatl/s. Day, 1873. 292-293 men re-examined, MNHN 02-08). Fowler (1956: (description; Ceylon). 146) cited Tillier's record. Hemirhamplllls lel/copterl/s. Day, 1878: 514 Hemirampill/s depal/peratlls Lay and Bennett as (description after Valenciennes, 1846). Day, He. eclanclleri. Seale, 1906: 13 (listed based on 1889: 423 (description after Day, 1878 and three specimens from the Marquesas Is., at least Valenciennes) . two of which, BPBM 2121 and ANSP 9185 are Hemirrilampilils [sic] GeO/-gii. Meyer, 1885: 38 He. depallperatlls). (listed; Manila). Hyporlwmplls a. acutlls (Gunther) as RilyncllO- Hemiramphl/s can tori. Jordan and Seale, 1906: rhampillls geO/·gii. Fowler, 1928: 75-76 (descrip- 207 (New Guinea, after Macleay, 1883). Wu, tion based on two specimens from the Kingsmill 1929: 65-66 (description; Amoy, China; fig. (Gilbert) Is., MCZ 8777). Fowler, ]938: 87 53 ). (listed based on two specimens from the Tuamotu Hemiramphlls cantoris. Jordan and Seale, 1907: Is., ANSP 79811, 79820); 154 (listed based on 8 (description; Cavite, Philippines). Fowler, two specimens from the Society Is., ANSP 75772); 1918: 62 (listed; Philippine Is.). 179 (listed based on two specimens from Christ- LoliRorilamphl/s norman; Whitley, 1931: 105- mas Is., Line Is.; ANSP 75795). Fowler, 1949: 106 (original description of genus and spe- 56 (listed after Fowler, 1938). Fowler, 1956: 146 cies; Townsville, Qld., pI. 12, figs. 2-3). (description largely based on Hy. aClltlls). See Munro, 1957: 57 (description, N. Qld.), fig. Collette (1974b: 118). 405 (after Whitley, 1931). Whitley, 1964: 39 Hyporhampl/lls cf. dl/ssl/mieri (Valenciennes) (listed). Marshall, 1964: 98 (after Whitley, as He. georgii. Borodin, 1932: 73 (listed based on 1931), pI. 24, fig. 106 (after Whitley, 1931). a specimen from the Red Sea, VMM 863). Fowler Marshall, 1966: 174 (after Whitley, 1931), (1956: 146) cited Borodin's record. pI. 24, fig. 106 (after Whitley, 1931). Grant, Hemiramplllls robl/stlls GUnther as He. cantori. 1972: 62 (brief description). Borodin, 1932: 43 (listed based on a specimen RhyncllOrhamphus georgii. Weed, 1933: 57-58 from Brisbane, Queensland, VMM 898). See (analysis of genera; RhyncllOrlwmphlis after Collette (1974a: 43). Fowler, 1928). Fowler, 1956: 146 (in part; ZellclI'C/lOptems sp. as He. geOlgii. Herre, 1933b: reference to Blegvad, 1944). Munro, 1957: 2 (listed based on two specimens (SU 28240) from 55 (description; Qld.; fig. 391). Munro, 1958: Madang, New Guinea). Fowler (1934: 393) 135 (literature records; New Guinea region). cited Herre's record. Whitley, 1964: 39 (listed). Grant, 1965: 36 Hyporlwmphl/s sp. as He. georgii. Seale, 1935: (uncommon in Qld. waters). Munro, 1967: 346 (listed based on specimens from Tulagi, Flor- 113 (description; New Guinea; pI. 11, fig. 171 ida Is. (CAS 5773) and Sikaiana Is. (CAS 5778), after Day, 1878, pI. 120, fig. 1). Pann and b:>th in the Solomon Islands). Shcherbachev, 1972: 569-571 (comparison Hemiramphl/s far (Forsskal) as He. georgii. with R. arabiclls). Collette, 1974a: 90-92 Intengan et aI., 1956: 201, table 7 (composition of synonymy; description; Australia; fig. 21). several species of fishes, including He. georgii for Hemiramplllls georgi. Umali, 1950: 7 (listed; which they used the common name of "spotted throughout the Philippines). Herre, 1953: halfbeak" indicating that they had Hemiramphl/s 153-154 (synonymy; distribution records). far rather than R. geOl·gii). Orsi, 1974: 163 (listed based on Tirant, 1885 and Chevey, 1934; Viet Nam). Diagnosis.-R. georgii has an intermediate Hemiramphlls pll/matus. De Silva, 1956: 51 (listed; after Blyth, 1859 and Day, 1873; Cey- number of gill rakers (Tab]e 1) on the first lon and Bengal). arch (52-67, x 59.4), many fewer than R. Hyporlwmphl/s georgii. Kovalevskaya, 1965: arabicus (69-78), fewer than but overlap- 125-126 (description and figs. 1-2 of ovarian eggs; Gulf of Tonkin). ping with R. malabaricus (57-71, x 64.7), Hyporhamphlls georgi. Jones, 1969: 6 (listed; and more than but overlapping with R. naga Laccadive Archipelago). (47-59, x 52.4). The sum of dorsal plus Hyporhamplllls cantori. Talwar and Chakra- pany, 1970: 128 (re-examination of specimen ana] rays separates most R. georgii (29 or [ZSI 1300] figured by Day, 1878, pI. 119, fig. more) from most R. arabicus and R. mal- 1). aharicus (28 or less) but there is a broad RhYllcilOrhamphlis georgi. Grant, 1972: 60 (after 1965 ed.). area of overlap at 28 and 29 with R. naga 92 BULLETIN OF MARINE SCIENCE, VOL. 26, NO. I, 1976 (Table 3). R. georgii and R. naga average Hemirhamphus plumatus Blyth, 1859. higher vertebral counts (55-57, Table 4) Lectotype, herein selected, ZSI 625; Ceylon; than R. arabicus and R. malabaricus (54 or 184 mm. D 15; A 14; RGRl 14 + 43 = 57; 55). R. georgii has a more highly domed RGR2 9 + 43 = 52; upper jaw length 12.9 upper jaw than do the other species of the mm, width 9.9 mm. Paralectotype ZSI 625; genus (Fig. 9b). Both the upper jaw and the Ceylon; 143 mm. snout are significantly longer in R. georgii Blyth's original description was based on than in the other species (Table 5): upper specimens from the Sandheads at the mouth jaw 46-75 thousandths of standard length of the Hooghly R. and from Ceylon. His (x 63.0) compared to 40-69 (x's 50.9-57.0) specimens were Rhynchorhamphus because in the other three species, 216-323 (x 275.9) he clearly stated "Each nostril covered by a thousandths of head length compared to 203- remarkable plume of filaments." The dorsal 307 (x's 223.5-248.4); snout length 83-106 and anal ray counts of 15-13 also fit Rhyn- (x 94.2) thousandths of SL compared to 65- chorhamphus but do not help in deciding if 101 (x's 85.1-90.9), 365-440 (x 412.2) Blyth had R. georgii or R. malabaricus be- thousandths of head length compared to 351- cause the dorsal count is characteristic of the 424 (x's 380.8-395.8). R. georgii has former and the anal count is characteristic smaller pharyngeal teeth than R. naga (Fig. of the latter. Day (1869) re-examined much 5), but they are larger than in either R. of Blyth's material that had been transferred arabicus or R. malabaricus. from the Asiatic Society of Bengal, where Blyth had been curator, to the Government Description.-Meristic data are presented in Museum, now the Zoological Survey of In- Tables 1-4. Pectoral fin rays 10-12, usually dia. Day redescribed He. plumatus based on 11. Predorsal scales 37-45, usually 39-42. two specimens, presumably part of Blyth's Morphometric data are summarized in Table type-material, from Ceylon, giving the same 5. dorsal and anal counts ,13 and :YJl but added "upper jaw :% longer than wide" indicating Maximum known size.-Female, 231 mm that these specimens were R. georgii. I have SL (AMS B.7517, Madras). re-examined what I believe to be these two Types.-Hemiramphus georgii Valenciennes, specimens and both are R. georgii. Dr. P. 1846. Holotype MNHN B.I062; Bombay; K. Talwar has kindly checked the records at Dussumier; 1830; female, 185 mm. D 17; the Zoological Survey of India and has con- A 15; PI 11-11; RGR 16 47 63; RGR2 I + = firmed my belief that these two specimens 10 49 59. Paratype MNHN B.I061; + = are syntypes. The syntypes from the Sand- MaM Bay, Coromandel, India; Dussumier; heads are not presently in the Zoological Sur- 201 mm. vey collections. The specimen selected as Hemiramphus eclancheri Valenciennes, lectotype is in better condition and has dor- 1846. Holotype MNHN 4592; "Marquises"; sal and anal ray counts (15-14) closest to Eclancher; 107 mm. D 16; A 15; PI 11-11; those in the original description. RGR 13 41 = 54; RGR2 7 40 = 47. I + + Hemirhamphus cantori Bleeker, 1866. Hemiramphus leucopterus Valenciennes, Lectotype, herein selected BMNH 1866.5.2. 1846. Holotype MNHN B.I065; Bombay; Dussumier; 89.5 mm. D 15; A 14; PI 11- 16; East Indies; Bleeker; 166 mm. D 15; A 15; PI 10-10; RGRI 14 + 43 = 57; RGR2 11; RGR1 12 + 43 = 55; RGR2 6 + 46 = 52. 8 + 45 = 53; upper jaw length 12.8 mm, Hemiramphus russeli Valenciennes 1846. width 8.9 mm. Paralectotypes RMNH 6953 Holotype MNHN B.1067; Pondichery, In- (2,130-168); East Indies; Bleeker. dia; Sonnerat; 202 mm. D 14; A ?; PI ?-11; Bleeker's original description of Hem i- RGR1 ?; head and half the skin fastened onto ramphus cantori was based on 62 specimens a stick. from the East Indies, of which 52 arc extant: COLLETTE: REVIEW OF THE GENUS RHYNCHORHAMPHUS 93 BMNH 1866.5.2.16 (1, 166 mm) and (Table 5). R. georgii was divided into eight RMNH 6953 (51, 69.1-168). Both R. populations for meristic comparisons (Tables geOl'giiand R. naga are present in the type- 1-4). The data presented by Talwar (I 964b) series. No holotype was designated and no for a sample from the Gulf of Mannar and one has subsequently selected a lectotype. Palk Bay are included, for comparison, in The original description is apparently based Table 1 (gill rakers on the first gill arch) on both species. According to Dr. M. Boese- and Table 3 (dorsal and anal rays). There man (pers. comm.), Bleeker frequently cut is interpopulation variation. Means differ by the gill-covers on the specimen that he ex- as much as seven gill rakers on the second amined most thoroughly and he frequently arch (Table 2, Bay of Bengal x 51.50, South selected the specimen in the best condition China Sea x 58.75). The single speci- on which to base his description. None of men from Australia that was x-rayed had the extant syntypes has the gill covers cut. one more vertebra than any other specimen All the specimens are now somewhat flac- x-rayed (Table 4). The meristic differences cid and most have their beaks broken; there- seem to be random, without showing clinal fore, no specimen is in a clearly better con- variation or other coherent geographic pat- dition. Of the four largest specimens, two terns. (168 and 166 mm SL) are R. georgii and two (168 and 164 mm SL) are R. naga. All Range.-R. georgii has the widest distribu- but one of the remainder of RMNH 6953 tion (Fig. 7) of any of the species of Rhyn- arc R. naga. Most authors have considered chorhamphus. In the Indian Ocean, it is cantori to be a synonym of georgii. In the found from the Persian Gulf through the interests of nomenclatural stability, I here- Arabian Sea and Bay of Bengal to the East with select as lectotype, BMNH 1866.5.2.16, Indies and Malaysia. Records from the Red thereby fixing cantori as a junior synonym of Sea (TiIlier, 1902; Borodin, 1932; and georgii. Fowler, 1956) are based on misidentifica- Loligorhamphus normani Whitley, 1931. tions. Early reports from the Seychelle Is- Holotype AMS IA.2319; Townsville, Qld.; lands (Playfair, 1867; Bleeker, 1874) that W. E. J. Paradice; 126 mm. D 14; A 15; PI were accepted by Smith and Smith (1963) and from Mozambique (Bianconi, 1854 ) 12-12; RGR1 14 + 44 = 58; RGR~ 9 + 45 = 54; vertebrae 40 + 19 = 59; upper jaw and Mauritius (Bleeker, 1879) seem du- length 8.1 mm, width 5.9 mm. bious and have not been verified. In the western Pacific Ocean, R. georgii rvomenclatural note.-H emiramphus georgii occurs from the East Indies, Malaysia, and Jerzmanska (1968: 410-412), a fossil fish the Gulf of Thailand through the South from the Carpathian Mountains of Poland, China Sea north as far as the Swatow-Amoy is clearly not congeneric with Rhynchorham- area of China. It is found throughout the phus (49-50 vertebrae, 17-18 anal rays). Philippine Is]ands and there are records from The high number of anal rays indicates that Borneo, New Guinea, and two parts of Aus- the species is not referable to Hemiramphus tra]ia: the Northern Territory and Queens- Cuvier sensu stricto and perhaps best in- land. All records from further west into the cluded in Hyporhamphus for the present. Central Pacific appear to be based on mis- identified specimens: Guam, Marianna Is. Geographic variation: Morphometric and (Pellegrin, 1898 and GUnther, 1909); So]o- meristic characters of populations of R. man Is. (Seale, 1935); Gilbert Is. (Fowler, georgii were compared. Morphometric data 1928) ; Society Is. (Fow]er, 1938); Marque- from the Indian Ocean and western Pacific sas (Seale, 1906); and Tuamotu Archipel- populations showed no differences in ranges ago (Fowler, 1938). The holotype of He. or means; therefore, the data were combined eclancheri Valenciennes was described "aux for comparison with the other three species iles Marquises par M. ]e Dr. l'Eclancher ... 94 BULLETIN OF MARINE SCIENCE, VOL. 26, NO.1, 1976 a bord de la fregate la Reine Blanche." AMNH 32512 (25, 118-198), and USNM 214093 (10, 120-190); Prachuab Khiri Khan Prov., Ban However, examination of extensive material Khlong Wan; K. Lagler et al.; 6 Sept. 1964. from the Central Pacific for a study of Hypo- UMMZ 191500 (I, 200); Prachuab Khiri Khan rhamphus acutus (Collette, 1974b) revealed Prov., Ban Khlong Wan; K. Lagler et al. 64-1322; 12 Dec. 1964. UMMZ 191414 (4, 160-197); Pra- only four species of Hemiramphidae: Hy. chuab Khiri Khan Prov., Ban Khlong Wan mkt.; acutus (Gunther), Hy. dussumieri (Valen- K. Lagler et al. 64-1321; 12-13 Dec. 1964. KUB ciennes), Hemiramphus depauperatus Lay 1.64: 1288 (2, 142-177); Prachuab Khiri Khan Prov., Ban Khlong Wan; S. Monkolprosit; 2 Oct. and Bennett, and Euleptorhamphus viridis 1965. (van Hasselt) in the area from the Hawaiian South China Sea: 12 specimens (85.0-200 mm Islands east to Easter Island. It seems likely SL) from 6 collections. BMNH 1860.7.20.16-17 (2, 177-200); China, Amoy; Stevens. ZMK 140-1 that the type-locality of He. eclancheri is in (2, 157-177); Hong Kong. FRSHK uncat. (I, error. 153); Hong Kong, Aberdeen fish mkt.; C. Y. Lin; 26 Jan. 1961. CAS SU 25654 (I, 190); Hainan, Material Examined.-Persian Gulf: 4 specimens near Hoihow; R. C. Miller; 25 June 1930. CAS SU (148-180 mm SL) from 3 collections. FMNH 60993 (5, 85.0-99.5); Hong Kong, Tolo Channel, 51254 (I, 148); Persian Gulf; W. P. Kennedy; 22° 26' 45" N, 1440 15' 00" E; R/V ALISTER 1934. ZMK CN 5-6 (2, 165-174); Persian Gulf, HARDY;R. L. Bolin; 6 Jan. 1958. UMMZ 70353 Iran, Bushire; H. Blegvad; spring, 1937. MNHN (I, 157); China, Amoy or Swatow; Beal and 65-558 (I, 180); Mer d'Oman; EI Husseini. Steere. Arabian Sea: 8 specimens (89.5-201 mm SL) Philippine Islands-Borneo: 31 specimens (90.5- from 7 collections. MNHN B.I062 (1, 185); 218 mm SL) from 12 collections. CAS SU Bombay; Dussumier; holotype of Hemiramphus 21677 (I, 162); P. I., Samar, Borongan; A. Seale. georgii. BMNH 1889.2.1.2163-4 (2, 163-184); CAS SU 37638 (5, 155-191); P. I., Luzon, Pan- Bombay; Day. ZSI 1300 (I, 154); Bombay. gasiman Prov., Dagupan; A. W. Herre Oriental MNHN B.1065 (I, 89.5); Bombay; Dussumier; Exped.; 17 Oct. 1940. CAS SU 9619 (2, 90.5- holotype of Hemiramphus leucopterus. ZSI F.I774 96.5); P. I., Luzon, Manila; G. A. Lung; 9 May (2, 141-150); Alibag, Kolba District; German In- 1901. USNM 138678 (1,205); P. I., Samar, Cat- dian Expedition; 24 Nov. 1955. MNHN B.I061 balogan; R/V ALBATROSS;17 April 1908. USNM (1, 201); Mahe Bay, Coromandel; Dussumier. 175088 (2, 187-218); P. I., Luzon, Lingayen Gulf; ZMK un cat. (1, 184); Cochin; 1 Nov. 1972. R/V ALBATROSS;11 May 1909. USNM 138681 Bay of Bengal: 11 specimens (81.1-231 mm SL) (I, 138); P. I., Leyte, Abuyog; R/V ALBATROSS; from 9 collections. ZSI 625 (2, 143-184); Ceylon. 26 July 1909. USNM 138679 (1, 128); P. I., CMFRI uncat. (I, 180); Gulf of Manar, Pambam, Luzon, Manila mkt.; R/V ALBATROSS;16 April Mandapam Camp; 5 Jan. 1958. MNHN B.1067 1909. USNM 175077 (11,149-202); P. I., Leyte, (1, 202); India, Pondichery; Sonnerat; holotype Abuyog; R/V ALBATROSS;26 July 1909. USNM of Hemiramphus russeli. AMS B.7517 (I, 231); 138680 (1,99.0); P. I., Luzon, Manila mkt.; R/V Madras, purchased from Day, 1885; "cotype" of ALBATROSS;15 April 1909. ANSP 55963-66 (4, Hemiramphus cantori Bleeker. ZSI F.2311/2 (I, 166-206); P. I., Luzon, Manila Bay, Orien; 1. 132); Marina Beach, Madras. NHMV 5585 (2, Clemens; 11 May 1923. CAS SU 27764 (1, 145); 126-127); Madras; Novara Expedition 1857-9; North Borneo, Sandakan; A. W. Herre; 29 June Ze1ebor. NHMV uncat. (I, 217); Madras; 1886; 1929. MSNG 13979 (I, 110); Borneo, Sarawak; Day. BMNH unreg. (I, 112); Bengal; Gen. Hard- Doria-Beccari; 1866-68. USNM 138688 (I, 161); wicke. CAS SU 37217 (1,81.1); La-c-ala or Julia North Borneo, Sandakan; R/V ALBATROSS;1 I., Mergui Arch.; A. W. Herre; 18 Jan. 1937. March 1908. Java Sea: 22 specimens (130-185 mm SL) from New Guinea-Australia: 6 specimens (126-209 10 collections. MCZ 702 (I, 146); Singapore. mm SL) from 6 collections. RMNH 10321 (1, NHMV uncat. (3, 134-184); Singapore and Bunka; 209); New Guinea, Astrolabe Bay near Madang; 1874 1.2049. RMNH 12203 (I, 173); Bay of 26 Oct. 1896. CSIRO uncat. (I, 207); Australia, Batavia; P. Buitendijk; Aug. 1912. RMNH 12202 Northern Territory, WesselL, Jensen Bay; 28 July (2, 142-180); Bay of Batavia; P. Buitendijk. 1949. FBQ 39 (I, 179); Australia, Queensland, Magnetic I., Horseshoe Bay; G. Coates; Sept. 1949. RMNH 12200 (4, 130-142); Java Sea; P. Buiten- FBQ 3608 (1, 164); Australia, Queensland, off dijk; 3 Feb. 1915. RMNH 12197 (1, 162); Java Cairns. BMNH 1927.2.10.21 (I, 132); Australia, Sea; P. Buitendijk. RMNH uncat. (2, 130-168); Queensland, Townsville; W. E. J. Paradice; Aug. East Indies; P. Bleeker; 1879; out of RMNH 6953. 1924. AMS IA 2319 (1, 126); Australia, Queens- RMNH uncat. (6, 154-180) ; East Indies; P. land, Townsville; W. E. 1. Paradice; holotype of Bleeker; out of RMNH 92138. RMNH 12141a (I, Loligorhamphus normani Whitley. 185); Batavia; P. Bleeker. ZMK 139 (I, 162); Java. Gulf of Thailand: 84 specimens (114-200 mm ACKNOWLEDGMENTS SL) from 5 collections. UMMZ 191189 (11, 159- 190); Prachuab Khiri Khan Prov., S of Ban I gratefully acknowledge the assistance and co- Khlong Wan; K. Lagler, S. Punpoka et al.; L64- operation of the curators and staffs of the institu- 1268; 3 June 1964. UMMZ 191266 (31, 114-194), tions housing material of Rhynchorhamphus, par- COLLETTE: REVIEW OF THE GENUS RHYNCHORHAMPHUS 95 ticularly James W. Atz (AMNH); Reeve M. Bailey eiland Pinang. Vers!. Akad. Amsterdam 12: (UMMZ); Marie-Louise Bauchot (MNHN); E. 64-80. Bertelsen (ZMK); C. R. Boccia (VMM); Eugenia 1866. Revision des Hemirhamphes de B. and James E. Bohlke (ANSP); M. Boeseman l'Inde archipelagique. Ned. Tijdschr. Dierk. (RMNH); Dan M. Carlsson (ZMK); W. L. Chan 3: 136-170. (FRSHK); William N. Eschmeyer (CAS); Charles 1871. Scombresoces. Atlas ichthyolo- Gruchy (NMC); G. G. J. Harrison (FBQ); V. S. gique des lndes Orientales Neerlandaises. Krishnamurty Chennubhotla (CMFRI); Georg Amsterdam. 6: 40-78. Kuhlmorgan (MFLB); Tomio Iwamoto (CAS); 1873. Memoire sur la faune ichthyolo- Paul Kiihsbauer (NHMV); Karel F. Liem (MCZ); gique de Chine. Ned. Tijdschr. Dierk. 4: A. G. K. Menon (ZSI); Supap Punpoka Monkol- 113-154. prasit (KUB); Ian S. Munro (CSIRO); Jprgen G. 1874. Poissons de Madagascar et de Nielsen (ZMK); G. Palmer (BMNH); John R. l'ile de la Reunion. Pages 1-104 in Recherches Paxton (AMS); John E. Randall (BPBM); Donn sur la faune de Madagascar et de ses de- E. Rosen (AMNH); Richard Rosenblatt (SIO); pendances d'apres les decouvertes de Fran, of Australia and New Zealand. Rec. Austral. derbilt South Pacific Expedition, 1937. Acad. Mus. 29: 11-105. Nat. Sci. Phila. Monogr. 2: 349 p. 1974b. Geographic variation in the 1949. The fishes of Oceania-Supple- Central Pacific halfbeak, Hyporhamphus acu- ment 3. Mem. Bernice P. Bishop Mus. 12: fus (Giinther). Pacific Sci. 25: 111-122. 35-] 86. Cressey, R. F., and B. B. Collette. 1970. Cope- 1956. Fishes of the Red Sea and south- pods and needlefishes: A study in host-para- ern Arabia. Weizmann Science Press, Jeru- site relationships. U.S. Fish. Wild. Servo Fish. salem, 1: 240 p. Bull. 68: 347-432. Grant, E. M. ]965. Guide to fishes. Queens- Cuvier, G., and A. Valenciennes. 1846. His- land Dept. Harbours and Mar. Fish. Notes 2: toire naturelle des poissons. P. Bertrand, 280 p. Paris, 19: 544 p. ---. ] 972. Guide to fishes. Queensland Dept. Day, F. 1865. The fishes of Malabar. 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