BULLETIN OF MARINE SCIENCE. 30(1): 13-20, 1980

NOTES ON INDO-PACIFIC CARANGID OF THE CARANGOIDES BLEEKER II. THE CARANGOIDES ARMATUS GROUP

F. Williams, P. C. Heemstra, and A. Shameem ABSTRACT The systematic status of two closely related of carangid fishes. genus Carangoides, is reviewed and clarified. Detailed species accounts are given for C. arma/us (Riippell) and C. hedlandensis (Whitley), and include synonymies, diagnoses, descriptions, geographic variation, status of type material, material examined, and taxonomic comments.

This is the second in a series of papers by the senior author to clarify the systematic status of several species groups within the genus Carangoides. A general introduction to the series was given in the first paper by Williams and Venkataramani (1978). M. M. Smith (1973) discussed in detail the identity of Sciaena armata Forsskal, 1775, which name has been commonly applied to a deep-bodied carangid with the breast naked of scales to the pectoral fin base and having elongated soft dorsal and anal fin lobes. Evidence was presented which clearly indicated that Sciaena armata Forsskal and Citula plumbea Quoy and Gaimard, 1824, should be con- sidered nomina dubia, and that the first secure name for this fish, large mature males of which are noted for having elongated central dorsal and anal fin rays, was C. ciliarius (Ruppell, 1830). During the present study of Indo-Pacific species of Carangoides, early data analyses indicated that there were probably two closely related species with a deep body, naked (scaleless) breast, and with the middle dorsal and anal fin rays elongated in large males. These species are now identified as C. armatus (Ruppell, 1830), and C. hedlandensis (Whitley, 1934), and are clearly separated by the number of lateral gill rakers on the first gill arches. Citula ciliaria Ruppell, a name used by some authors for these species, is here considered a nomen dubium (see Discussion below). It should be pointed out that Citula Cuvier, 1816, and Olistus Cuvier, 1829, are senior generic synonyms of Carangoides Bleeker, 1851; the type species are respectively citula Cuvier, 1833 = Caranx armata Ruppell, 1830, by monotypy and Olistus malabaricus Cuvier, in Cuvier and Valenciennes, 1833 (preoccupied by Scomber malabaricus Bloch and Schneider, 1801, next available name = OUstus hedlandensis Whitley, 1934), by subsequent designation of Jor- dan, 1917, Genera of fishes, pt. 1, page 129.

METHODS AND MATERIALS

Methods used in this study are as given in Williams and Venkataramani (1978). Abbreviations used for institutions cited are as follows: AMS-Australian Museum, Sydney; CAS-California Academy of Sciences, San Francisco; FMNH-Field Museum of Natural History, Chicago; MNHN-Museum national d'Histoire naturelle, Paris; SMF-Natur-Museum und Forschungs-Institut, Senckenberg, Frankfurt am Main; TABL-Southeast Center, National Marine Fisheries Service, Miami (formerly Tropical Atlantic Biological Laboratory; collections of carangid fishes transferred to CAS). Uncatalogued California Academy of Sciences specimens collected under the auspices of the George Vanderbilt Foundation (GVF) are reported under their register and station numbers. Most collections from Ceylon made under Smithsonian Institution Foreign Currency Grant 3818 (SI 3813) are uncataIogued and are listed by station number and abbreviations refer to the following collectors listed in chronological of their visits to Ceylon: F. J. Schwartz (FJS); W. F. Smith- Vaniz (SV); P. C. Heemst.ra (PCH); C. C. Koenig (CCK); T. R. Roberts (TRR); T. Iwamoto (TI). Other collections 13 ]4 BULLETINOFMARINESCIENCE.VOL.30. NO.I. 1980

a b Figure I. Carangoides armatus: (a) Male, CAS 16330, 269 mm FL, Ceylon, 12 August 1969; (b) Female, CAS 16331,236 mm FL, Ceylon, 27 July 1969.

from Mozambique, Ma]aysia and Philippines made under SI 3818 are also uncatalogued and are designated by geographic location and station numbers (if used). Collections from the International Indian Expedition (lIOE) cruises of the R/V ANTON BRUUN are also uncatalogued and are listed by cruise and station number. The material examined sections include only specimens examined by the authors (or associates); localities are abbreviated.

Carangoides armatus (Ruppell, 1830) (Figure la and b, Table 1)

Citu/a armata, Ruppell, 1830, 103 (Red Sea); Caranx citu/a Cuvier, in Cuvier and Valenciennes, 1833, 94 (type locality unknown, probably Red Sea); Caranx cirrhosus (Ehrenberg) Cuvier, in Cuvier and Valenciennes, 1833, 94 (Red Sea); Caranx rastrosus Jordan and Snyder, ]908, 37 (Taiwan); Caranx (Citu/a) sch/egeli Wakiya, ]924, ]70, pI. 20, fig. 2 (Nagasaki, Japan); Caranx (Citu/a) ciliaris Wakiya (non Ruppell), 1924, 172, pI. 21 (Taiwan); Citu/a pescadorensis Oshima, 1925,395, pI. I, fig. 2 (Pescadores Is., Taiwan); Carangoides armatus Williams (non Forsski\.]), 1958,398 (Zanzibar). Diagnosis.-A carangid fish of the genus Carangoides with the naked (scaleless) area of the breast extending up to the pectoral fin base but not above it, very deep body with a steep dorsal head profile, and a total of 30-37 lateral gill rakers on the first arch. Description (based on 84 specimens, 57-503 mm FL).-Body measurements ex- pressed as percent offork length: total length 102.9-121.1; standard length 90.0-

95.8; head 24.1-31.4; depth, Dl to P2 35.9-54.9, D2 to A2 38.6-55.6; orbit 6.4- 12.0; interorbital 6.5-10.3; snout 6.9-9.8; postorbital 9.8-12.7; suborbital 2.4-4.0; upper jaw length 10.2-13.5; maxilla width 2.8-4.6; pectoral fin 23.3-41.1; pelvic

Table I. Meristic characters of C. armatus and C. hed/andensis

C. a'marlls (57-503 mm FL) C. "ed/andensi.' (71-283 mm FL)

Character Range Range n

O2 soft rays 19-22 20.19 84 20-22 20.97 66

A2 soft rays 16-18 16.95 84 16-18 17.05 66 Lateral gill rakers on first arch upper 10-15 12.32 78 6-1 ] 7.85 78 lower 20-24 21.94 78 14-]7 15.73 78 total 31-37 34.26 78 20-27 23.58 78 CLL scales (left) 57-77 66.39 57 59-77 68.93 60 SLL scutes (left) 11-24 17.07 82 17-29 22.00 74 SLL total elements (left) 25-43 32.69 59 29-41 34.6] 70 WILLIAMS ET AL.: SYSTEMATICS OF CARANGOIDES 15

Figure 2. Extent of naked (scaleless) area of breast in (a) Carangoides armatus and (b) C. hedlan- dens;s.

fin 10.6-26.7; height Dl spine 8.6-14.1; height D2 fin lobe 18.1-62.9; height A2 fin lobe 18.0--49.2; snout to Dl 35.7--41.3; snout to D2 50.3-55.9; snout to A2 53.2-

62.6; snout to PI 25.4-31.9; snout to P2 31.7--42.5; base D2 fin 34.2--47.1; base of A2 fin 31.4--42.4; CLL 34.5--42.6; SLL 24.6-35.0 VIII + I, 19-22; anal fin II + I, 16-18; pectoral fin 1, 18-20. Lateral gill rakers on first arch 10-15 + 20-24 = 31-37; lanceolate with small bony setae on inner edge. CLL scales 57-77; SSL 11-24 weak scutes and 25--43 total ele- ments; MLLR 0.68-0.89. Body strongly compressed and deep; head profile very steep but relatively straight from nape to snout (Fig. I). Adipose eyelid weak, forming rim around eye. Mouth cleft at or below level of eye. Posterior edge of maxilla ranges from vertical through front edge to about posterior edge of pupil. Jaws protusible, lower jaw longer. Juveniles (less than 60 mm FL) with 1-3 irregular rows of small conical teeth in jaws; larger fish with small conical teeth in bands, widest ante- riorly. Villiform or very small conical teeth in curved band on palattnes, in wedge shape on vomer, in central band on tongue. Some scales on cheeks and postor- bital, rest of head naked. Breast naked (scaleless) ventrally and laterally from isthmus up to pectoral fin base; posterior margin of naked area running diagonally from posterior point of pectoral fin base to anus or just in front of anus, that is to about opposite tip of innermost ray of pelvic fins (Fig. 2a). Second dorsal and anal fins with strong falcate lobes, which are produced into long filaments (may exceed 50% fork length) with increasing body size. In mature males, greater than 210 mm FL, 3 to 12 of the central dorsal fin rays (ranging from #6-18) and 0-10 central anal fin rays (ranging from #6-15) are produced into filaments of greatly varying lengths. Eleven of 15 mature females had no elongated central dorsal or anal fin rays, but four specimens had from 5 to 7 central dorsal fin rays (ranging from #7-11) produced into filaments. Number and degree of extension of rays did not appear to be related to stage of maturity of gonads or size of fish. Basal scaly sheaths to both second dorsal and anal fins. Pectorals subfalcate in small specimens to falcate in larger ones; pelvics short, fitting in ventral groove which is rather more prominent than in most carangids but not as long or pronounced 16 BULLETIN OF MARINESCIENCE. VOL. 30. NO. I. 1980

Table 2. Geographic variation in total counts in C. armatus

(I) (2) (3) East Africa! Gulfof Thailand! South India! All Location Red Sea Hong Kong{I'aiwan Ceylon locations

Range 31-33 33-36 31-37 31-37 Mean (x) 32.99 34.20 34.48 34.25 Standard deviation (Sx) 0.76 1.14 1.38 1.44 Standard error (SJ 0.29 0.36 0.18 0.16 Number (n) 7 10 60 77

Underlining of means (x) refers to similarities of population values derived from the Student~Newman~Keuls multiple range tests.

as in Atropus atropus. Caudal deeply forked; bilateral paired caudal keels pres- ent. Examination of radiographs shows no hyperostosis in this species. Color (freshly preserved).-Up to 440 mm FL body generally silvery white, dusky or bluish above, and pale whitish below. At 58 mm FL, six dark crossbars still visible on body posterior to pectoral fin base. Head silvery white; nape, snout, postorbital and opercula tend to be dusky bluish, more so with increasing size of fish. Fairly large black opercular patch. First dorsal fin pale grey in small speci- mens, blackish in large specimens. Second dorsal and anal fins pale hyaline. Caudal fin pale hyaline with trailing edge dusky; black patch on peduncle at base of upper caudal fin. With increasing size, greater than 200 mm FL, second dorsal, anal, and caudal fins becoming dusky to black; leading edge of anal spine whitish as may be anal fin ray tips and scaly sheath. Pectorals pale hyaline, upper edge dusky, in specimens less than 200 mm FL, blackish hyaline at greater than 200 mm FL; axil dusky black. Pelvics generally black, though leading edge may be whitish in specimens less than 100 mm FL but becoming paler with increasing size until at 200 mm pale or only slightly dusky. Distribution.-Based on types, present collections, and confirmed identifications from: Madagascar, East Africa, Red Sea, Ceylon, south and southeast of India, Gulf of Thailand, Hong Kong, Taiwan, Okinawa, and Japan. Geographic variation.-The principal meristic character used to separate C. ar- matus from C. hedlandensis is total gill raker count on the first gill arch. The same character was used to compare populations of C. armatus from three lo- cations-East Africa/Red Sea (I), Gulf of Thailand/Hong Kong/Taiwan (2), and Ceylon/South India (3). The mean total gill raker counts for populations were subjected to a one-way analysis of variance (ANOY A) to test the hypothesis Ho: ILl = 1L2 = 1L3 (Table 2); the hypothesis was rejected at P < 0.0005. The data were then subjected to a Student-Newman-Keuls (SNK) multiple range test, with unequal sample size. The test shows ILl ~ 1L2 = 1L3 (Table 2), indicating that the East Africa/Red Sea population is different from the Gulf of Thailand/Hong Kong/ Taiwan population and the South India/Ceylon population, which are similar to each other.

Material Examined.

TYPE SPECIMENS: Citula armata Riippell, lectotype SMF 1601 (152 mm FL), Massawa, Red Sea; paralectotype SMF 15074 (154), Massawa, Red Sea. Caranx citula Cuvier, holotype MNHN 6182 (178), locality unknown, probably Red Sea; paratype MNHN 6181 (182), Red Sea. Caranx rastrosus Jordan and Snyder, holotype FMNH 55363 (288), Takao, Taiwan.

OTHER SPECIMENS: Madagascar, Baie d'Ambor, JR27 2(57 and 104 mm FL). Zanzibar, EAMFRO K38(61), KI8 (162 and 177), Kl1 (169 and 170), Kl(350); TABL 105612 (166). Red Sea, MNNH 6179 WILLIAMS ET AL.: SYSTEMATICS OFCARANGOlDES 17

a b

Figure 3. Carangoides hedlandensis: (a) Male, CAS 16332, 198 mm FL, Gulf of Thailand, 23 May 1961; (b) Female, CAS 16333, 190 mm FL, Gulf of Thailand, 10 May 1961.

(147). Southeast India, Tamil Nadu, Mandapam, FHB 66-36 (153 and 162), 66-37 (104); \3°\3'N, 80025'E, FHB 66-13 3(175-237); Porto Novo, FHB 69-]7 TABL 106472 (184). Eastern India, Andra Pradesh, Visakhapatnam, uncal. (153). Ceylon SI 38]8, CCK 69-8 (2\3), 69-9 (204),69-33 (137), 69- 43 (60), 69-45 (348), 69-54 (71),69-58 (223),69-60 (83), 69-132A (58); FJS 69-26 (179), 69-55 3(3]5- 395), 69-64 (249), 69-65 6(210-280), 69-70 (311); PCH 69- J 85 8( 110-171), 69-189 (352), 69- J 93 (258 and 298),69-]93 CAS ]6330 (269),69-2]7 (434), 69-218 (85 and 121),69-263 (239), 69-3094(86-104); SV 69-83 (337 and 504), 69-150 (491), 69-159 CAS ]6331 (236 and 253), 69-160 CAS 1646] (127 and 197), 69-164 3(250-328); TI 70-326 (108). Gulf of Thailand, GVF 92 (288), GVF 15]24(131-144),1548 (93), 1550 (191). Hong Kong, GVF HK1l2 (305). Japan, Nagasaki, FMNH 59402 (77).

Carangoides hedlandensis (Whitley, 1934) (Figure 3 a and b, Table 1)

Olistus malabarieus Cuvier, in Cuvier and Valenciennes, 1833, 103 (Malabar); Caranx plumbeus Jordan and Seale (non Quoy and Gaimard), 1906, 223, fig. 28 (Samoa); Caranx (Cirula) armalus Wakiya (non Forssk~l), 1924, 169, pI. 20, fig. I (Taiwan); Caranx (Cilula) plumbeus Wakiya (non Quoy and Gaimard), 1924,36], pI. 20, fig. 3 (Kii, Japan); Cilula armala Oshima (non Forssk~I), 1925, 396, Taiwan; OIislus hedlandensis, Whitley, 1934, 156, fig. 2 (Port Hedland, Western Australia); Cirula ciliaria Nichols (non Ruppe])), ]940, 201 (Bali, Indonesia); Cilula plumbeus Suzuki (non Quoy and Gaimard), 1962, 173 (Japan). Diagnosis.-A carangid fish of the genus Carangoides with the naked (scaleless) area of the breast extending up to the pectoral fin base but not above it, a very deep body and a steep dorsal head profile with distinct break in the profile in the interorbital region in adults, and a total of 20-27 lateral gill rakers on the first arch. Description (based on 79 specimens, 71-283 mm FL).-Body measurements ex- pressed as percent of fork length: total length 108.5-118.5; standard length 90.5-

94.3; head 25.5-29.9; depth, D1-P2 42.3-49.4, D2-A2 43.8-51.7; orbit 8.3-10.7; interorbital 8.1-10.1; snout 7.0-9.0; postorbital 10-12.8; suborbital 2.8-5.1; upper

Table 3. Geographic variation in total gill raker counts in C. hedlalldellsis

(I) (2) (3) South India! Northern Gulf of (4) All Location Ceylon of Bengal Thailand Philippines locations

Range 20-25 22-24 22-25 23-27 20-27 Mean (x) 23.25 23.38 23.21 24.25 23.42 Standard deviation (S~) 1.\3 0.74 0.89 1.22 1.1/ Standard error (S~) 0.18 0.26 0.24 0.35 0.13 Number (n) 40 8 14 12 74 18 BULLETINOFMARINESCIENCE,VOL.30. NO. I. 1980 jaw length 11.0-13.1; maxilla width 3.0-4.3; pectoral fin 22.2-40.9; pelvic fin

10.4-25.4; height DI spine 9.6-15.2; height D2 fin lobe 21.0-62.1; height A2 fin lobe 16.8-52.0; snout to Dl 34.6-42.8; snout to D2 50.4-56.5; snout to A2 54.2-

60.0; snout to PI 25.9-30.5; snout to P2 31.6-37.4; base D2 fin 38.0-42.9; base A2 fin 32.1-37.8; CLL 34.9-44.9; SLL 23.4-32.6. Dorsal fin VIII + 1,20-22; anal fin II + I, 16-18; pectoral fin I, 18-20. Lateral gill rakers on first arch 6-11 + 14-17 = 20-27; lanceolate with small bony setae on inner edge. CLL scales 59-77; SSL 17-29 weak scutes and 29-41 total ele- ments; MLLR 0.60-0.88. Body strongly compressed and deep; head profile extremely steep with a dis- tinct break in contour in interorbital region which becomes more pronounced with increasing size (Fig. 3). Rest of description as for C. armatus unless oth- erwise indicated. Extent of naked (scaleless) area of breast similar to that in C. armatus but slightly more constricted below pectoral fin base than in that species (see Fig. 2b). In mature males greater than 165 mm FL 3 to 8 (usually 5-7) of the central dorsal fin rays (ranging from #5-13) and 3 to 8 (usually 5-6) of the central anal fin rays (ranging from #5-10) are produced into filaments. In 9 mature fe- males of 185 to 271 mm no central dorsal or anal fin rays were produced into filaments. Examination of radiographs shows no hyperostosis in this species. Color (preserved).-As for C. armatus except as follows. At greater than 150 mm FL pelvic fins appear to .be .variable in color from pale to dusky-blackish. First dorsal fin black in specimens less than 80 mm FL, becoming greyish above this length. Distribution.-Based on holotype, present collections, and confirmed identifica- tions from: Seychelles, south and east coasts of India, Ceylon, northern Bay of Bengal, Gulf of Thailand, Philippines, Taiwan, Okinawa, Japan, New Guinea, northwest of Australia, and Samoa. Geographic variation.- Total gill raker counts were used to compare populations of C. hedlandensis from four locations-South India/Ceylon (1), northern Bay of Bengal (2), Gulf of Thailand (3), and Philippines (4). The mean total gill raker counts for populations were subjected to a one-way analysis of variance (ANO- VA) to test the hypothesis Ho: f.Ll= f.L2 = f.L3 = f.L4 (Table 3); the hypothesis was accepted at P > 0.05, indicating no significant differences among the popula- tions.

Material Examined. TYPE SPECIMENS: GUstus malabaricus Cuvier, holotype MNHN 3061 (283 mm FL), Malabar, S. India; paratype MNHN A.588 (243), Seychelles. GUstus hedlandensis Whitley, holotype AMS 1-12957 (171), Port Hedland, western Australia. OTHER SPECIMENS: South India, Tamil Nadu, Cape Comorin, FHB 66-45 TABL (162); Eastern India, Andhra Pradesh, Kakinada, 11(154-227). Ceylon, SI 3818, CCK 69-12 (105),69-42 (91),69-51 14(71-92),69-54 (90),69-1]5 (77); FJS 69-64 (260), 69-65 (249),69-683(262-271),69-84 (90); PCH 69- 223 (192 and 246); SV 69-81 (213 and 278). Bay of Benga], 20027'N, 92°20'E, AB47 CAS 164629(138- ]6]). Gulf ofThailand. GVF 1548(135),22]4 (165), 2494 3( 125-145),2546 3(179-222),2566 (207),2567 CAS 16333 (182 and 190),2574 (]89), 2600 CAS 163323(164-]98),2872 (135). Philippines, Bataan, CAS 16467 (115); Lingayen Gulf, FJS 69-84 7(185-216). New Guinea, MNHN 6180 (110), MNHN A6I02 (172). Taiwan, FMNH 58932. Japan, Kii, FMNH 59426 (144). DisClission.-In our opinion Smith (1973), as mentioned earlier (p. 18) presented conclusive evidence why Sciaena armata Forsskill and Citliia pilimbea Quoy and Gaimard should be considered nomina dllbia. She also indicated that the first secure name for armatus of authors was Citula ciliaria Riippell, 1830. An WILLIAMS ET AL.: SYSTEMATICS OF CARANGOIDES 19

excellent description and text figure of that species was presented by Ruppel! but without a count of gill rakers. This turns out to be a critical omission in view of our discovery of two closely related species differing only in the number of gill rakers. C. ciliaria is represented in the col!ections of the Senckenberg Mu- seum by two syntypes: SMF 9475, a dried mount (examined by W. F. Smith- Vaniz), and SMF 14129, a dried whole specimen (examined by F. W.). Radio- graphs provided by Dr. W. Klausewitz (SMF) indicate that the gill arches were removed, as was the custom, at the time of preparation of the specimens. Ruppel! iAhis handwritten manuscript of the "Verzeichniss" (1852) mentioned two dried specimens and one in alcohol but this latter specimen has not been found (Klau- sewitz, pers. comm.). The two dried specimens (SMF 9457 and 14129) both still have attached the original Ruppel! labels with the older label with the annotation "Citula ciliaria (Rupp.) Mare rubr." and the other with the annotation "Otistus Riippelli, Cuv. leone: Ruppel! Atlas T.25 F.8 Mare Rubrum. Geschenk von Dr. Riippell 1828" confirming that O. riippelli Cuvier should be considered an objec- tive synonym of C. ciliaria Riippel!. No consistent (objective) measurable char- acter other than gill raker count permits separation of the two species at all life history stages. Thus C. citiaria Ruppel!' is indeterminate and must be considered a nomen dubium. In 1830 Ruppell also described Citula armata based on material from the Red Sea which differed from his C. ciliaria in not having the elongated dorsal and anal rays of the latter. Smith (1973) also discussed why Ruppell believed himself to be the author of Citula armata and not Forsskal (1775). C. armata Ruppel! is represented in the SMF by three syntypes: a dried specimen, SMF 2853, a ra- diograph of which indicates a lack of gill arches, and two alcohol specimens, SMF 1601. We have examined the latter specimens and they have gill raker counts of 12 + 20 = 32 (FL 192) and 11 + 21 = 32 (FL 194). Thus the name Citula armata Ruppel!, 1830, is available for the species shown to have 30-37 lateral gill rakers on the first gill arch; this means that by fortunate coincidence the well-used name of Carangoides armatus is conserved by an alteration of authorship from Forsskal to Ruppel! (as hoped for by Smith, 1973, p. 354). Spec- imen SMF 1601 of FL 152 is hereby designated as the lectotype of Citula armata Riippell and SMF 15074 of 154 mm FL the paralectotype. The first secure name for the other member of the species group, the species with the low range of gill raker counts (20-27), is Carangoides hedlandensis (Whitley, 1934). We have now also determined that Otistus malabaricus Cuvier, 1882, indeed represents this species but the specific name malabaricus is un- available in Carangoides, being preoccupied by Scomber malabaricus Bloch and Schneider, 1801 (=Carangoides malabaricus, see Williams and Venkataramani, 1978). Although Whitley described C. hedlandensis in 1934, there have been few, if any, references to the species since that time. This is not surprising as the two species, C. armatus and C. hedlandensis, are very similar except for gill raker counts, and shape of the head profile in adults. We believe that many col!ections previously identified as C. armatus Forsskal will contain specimens of both C. armatus Riippell and C. hedlandensis. No attempt has been made in this study to check all descriptions of C. armatus derived from existing collections, and to separate these into the component species. Neither species appears rare in the Indo-West Pacific Ocean. The key character in separating the two species is number of gill rakers, and the difference between the mean total counts in C. armatus (i = 34.26) and C. hedlandensis (x = 23.58) is highly significant (P ~ 0.001, df 154, 2-sample '1' test) (see Table 1). Certain other counts (Table 1) show differences, but these appear 20 BULLETINOFMARINESCIENCE,VOL.30, NO. I, 1980

insufficient for species separation because of overlapping ranges and standard 1 deviations • Mean counts for the two species were compared by 2-sample 't' test with the following results: (1) Mean number of eLL scales is significantly higher in C. hedlandensis (x = 68.93) than in C. armatus (x = 66.39) (P < 0.002, df 115); (2) Mean number of SLL scutes is significantly higher in C. hedlandensis (X = 22.0) than in C. armatus (x = 17.07) (P ~ 0.001, df 154); (3) Mean number of SLL total elements is significantly higher in C. hedlandensis (X = 34.61) than in C. armatus (x = 32.69) (P < 0.001, df 127).

ACKNOWLEDGMENTS

We greatly acknowledge the assistance and cooperation of curators and staffs of institutions housing carangid fishes, particularly J. Paxton (AMS); W. F. Smith-Vaniz (ANSP); W. N. Eschmeyer and T. Iwamoto (CAS); M. L. Bauchot (MNHN); W. Klausewitz (SMF); and F. H. Berry (TABL). Special thanks are due to the members of the expedition to Ceylon (and other locations) under Smithsonian Institution Foreign Currency Program Grant 3813 (C. R. Robins, Principa] Investigator) for the ex- cellent quality of material collected over a large size range. Work by the senior author on Indo-Pacific carangid fishes was supported by the National Science Foundation, Grants GB-31 ]97 and GB-3979t. Work and advice of former UMJRSMAS2 graduate student W. F. Smith- Vaniz during and following the project is gratefully acknowledged; thanks also go to former graduate students J. Burnett-Herkes, E. Chan, M. Simon, and F. Vaughan at UM! RSMAS, former undergraduate students D. Kelso and K. Setbacken at UCSD3, and to D. W. Berry of Miami, who for varying periods, assisted in measurements of fish and data processing. Radiographs and photographs were made by W. F. Smith-Vaniz and F. H. Berry. The manuscript has benefited from reviews by F. H. Berry, D. P. de Sylva, W. J. Richards and W. Smith-Vaniz.

LITERATURE CITED

Cuvier, G. L. C. F. D., and A. Valenciennes. 1833. Histoire naturelle des Poissons. 9:i-xxxix and 1-512, pI. 246-279. F. G. Levrault, Paris. Jordan, D. S., and A. Seale. 1906. The fishes of Samoa. Description of the species found in the archipelago, with a provisional check-list of the fishes of Oceania. Bull. U.S. Bur. Fish. 25: 173- 455, fig. I-I], pI. 33-53. Jordan, D. S. and J. O. Snyder. 1908. Description of three new species of carangoid fishes from Formosa. Mem. Carnegie Mus. Pittsburgh 4: 37-40, pI. Nichols, J. T. 1940. On CitLlla ciliaria Riippell. Copeia 1940: 20]-202. Oshima, M. 1925. A review of the carangoid fishes found in the waters of Formosa. Philippine J. Sci. 26: 345-413, I pI., 2 fig. Riippell, W. P, E. S. ]830, Atlas zu der Reise in nordlichen Afrika. Zoologie, Fische des Rothen Meeres. 16: 95-118. Riippel!, W. P. E. S. 1852. Verzeichniss in dem Museum der Senckenbergischen-Fische und deren Skelette. Frankfurt am Main, iii-iv + 1-40. Smith, M. M. 1973. Identity of Caranx armatLIs (Pisces: ). Copeia 1973: 352-355. Suzuki, K. 1962. Anatomical and taxonomical studies on the carangid fishes of Japan. Rep. Fac. Fish. Prefec. Univ. Mie 4: 43-232. Wakiya, Y. 1924. The carangoid fishes of Japan, Ann. Carnegie Mus. ]5: 139-293, Whitley, G. P. 1934. Studies in Ichthyology No.8. Rec. AustrI. Mus. 24: 153-163,3 fig. Williams, F. 1958. Fishes of the family Carangidae in British East African waters. Ann. Mag. Nat. Hist. 13: 369-430. Williams, F., and V. K. Venkataramani. 1978. Notes on Indo-Pacific Carangid fishes of the genus Carangoides Bleeker. I. The Carangoides malabaricLls group. Bull. Mar. Sci. 28: 501-511.

DATE ACCEPTED: December 8, ]978.

ADDRESS: (FW) Division of Biology and Living ResoLlrces, Rosenstiel School of Marine and At- mospheric Science, University of Miami, 4600 Rickenbacker CaLlseway, Miami, Florida 33/49; (PCH) J. L. B. Smith Institute of Ichthyology, Rhodes University, Grahamstown, SOLlth Africa; (AS) De- partment of Marine Science, Andhra University, Waltair, Visakhapatnam 530003, India.

I Variances in the samples of rhe two species appear to differ significantly (2-taiJed variance ratio test) . • University of Miami, Rosenstiel School of Marine and Atmospheric Science.

3 University of California at San Diego,