BULLETIN OF MARINE SCIENCE, 28(3): 501-511, 1978

NOTES ON INDO-PACIFIC CARANGID OF THE BLEEKER r. THE CARANGO/DES MALABAR/CUS GROUP

F. Williams and V. K. Venkataramani

ABSTRACT

The systematic status of two closely related of carangid fishes, genus Carallg- oides, is clarified. Detailed species accounts are given for C. malabaricus (Bloch and Schneider) and C. falamparoides Bleeker and include synonymies, diagnoses and descrip- tions, geographic variation, status of type material, material examined, and taxonomic comments.

The , one of the four sub- the two genera is the dentition. Carangoides families of the family currently is characterized by villi form teeth in bands in accepted, represent about 90% of the nom- each jaw, widest anteriorly; the outer teeth inal carangid commercial catch of 2.6 million in one or both jaws may be slightly enlarged metric tons landed throughout the world. in large specimens. In there is a Much of the Caranginae catch is probably single series of moderate to strong subequal comprised of the schooling species of horse- conical teeth in each jaw, with an inner band mackerels () and scads (Decap- of fine teeth in the upper jaw. From the tertls) . Other species, such as Caranx and dentition (and other characters) it is clear Carangoides, are major elements, because of that the two genera are morphologically their size and number, available to sub- close, as indicated by Bleeker classifying sistence and small-scale throughout them as subgenera. The use of Carangoides their wide geographic ranges. Despite their remains one of convenience until such time as economic importance, our knowledge of the we have a better understanding of the large , biology, and ecology of carangids number of species themselves, especially has remained at a relatively low level com- intraspecific variations in specific characters, pared with many other groups of fishes, with distribution, and ecology. Then a more some notable exceptions. Obviously, an im- comprehensive evaluation of the status of portant basic step in the rational utilization Carangoides and its relationships with Ca- and management of living resources is to ranx sensu stricto, as well as , , know how to identify the species involved. Gnathonodon, Selaroides and , will be Following most recent workers on ca- possible, and the phylogenetic lines within rangid fishes, Chan (1968), Smith (1965, the Caranginae may be more objectively pro- 1967a and b), Smith and Smith (1963), jected. Suzuki (1968) and Williams (1958, 1961) It should be pointed out that Glistus we recognize all, or most, of Bleeker's sub- Cuvier 1829 is a senior generic synonym of genera of Caranx (Lacepede) as genera (as Carangoides Bleeker 1851. The type species indicated in Weber and de Beaufort, 1931). is Olistus malabaricus Cuvier 1833 by sub- Two of the most speciose of those genera are sequent designation of Jordan 1917 (p. Carangoides and Caranx sensu stricto, spe- 129). To avoid additional confusion, use of cies of the former being currently restricted Carangoides is continued until such time as to the Indo-Pacific , while species of a review of carangid genera has been com- the latter are found world-wide in tropical pleted. waters. The principal character separating This is the first in a series of papers by the

501 502 BULLETIN OF MARINE SCIENCE, VOL. 28, NO.3, 1978 senior author to clarify the systematic status fork length (FL) as the critical length mea- of several species groups within the genus surement as, (1) FL is more familiar to - Carangoides. eries biologists, who have the greatcst need Smith (1967b) discussed at length the to identify carangids, and (2) FL is easier, identity of Carangoides malabaricus (Bloch and faster, and more accurate to determine and Schneider) 1801 and clearly indicated than standard length because of the difficulty that two species had been confused in past in ascertaining the posterior margin of the literature. Smith's revision of nomenclature hypural plate, especially in large fish. resulted in the more commonly cited species The following measurements were made (erroneously described in the past as C. on all specimens during thc study (though malabaricus) being allocated to C. caerule- not necessarily utilized in this paper): opinnatus (Riippell) 1830, and the other TOTAL LENGTH (TL). Length of the species redefined as C. malabaricus Bloch body from the tip of the snout to vertical and Schneider. Two of the essential differ- through tip of the longest caudal fin lobe (in ences between the two species are: (1) ex- the natural position). tension of the naked area of the breast FORK LENGTH (FL). From the tip of anteriorly above the pectoral fin base in C. snout to tip of shortest median caudal fin ray. malabaricus but not in C. caeruleopinnatus, STANDARDLENGTH (SL). From tip of and (2) significantly higher lower limb gill snout to posterior margin of hypural bones raker counts in C. malabaricus (23-26 ver- (caudal base). sus ]5-18). BODY DEPTH (Dl-P~), From origin of During the present study 389 recently ac- the first dorsal spine to dorsal point of in- quired carangid specimens from the Indo- sertion of the pelvic fin. West Pacific Ocean were provisionally iden- BODY DEPTH (D~-A~). From origin of tified as C. malabaricus Bloch and Schneider the second to the origin of the (following Smith, ] 967b). As detailed exami- anal fin. nation of the material progressed, it became PECTORALFIN LENGTH (P ). From dorsal obvious that there were probably two rather 1 point of origin of the spine to the tip of the similar species involved; the second species longest ray. was not C. caeruleopinnatus and is now PELVIC FIN LENGTH (P~). From anterior shown to be C. talamparoides Bleeker, 1852. point of origin of the spine to the tip of the Both species described here have been longest ray. found in individual lots from the same sta- HEIGHT OF DORSAL SPINE. From anterior tion in the Arabian Sea, Ceylon and Gulf point of origin to tip of the longest (usually of Thailand, thus eliminating the possibility third) dorsal-fin spine (if spine other than that the observed variations might be due to hybridization of C. malabaricus with a the third was longest, it was so noted). species such as C. caeruleopinnatus or to HEIGHT OF SECOND DORSAL FIN LOBE. environmentally induced differences through- From origin of second (soft) dorsal fin to out the range of the species. tip of dorsal fin lobe. HEIGHT OF ANAL FIN LOBE. From origin METHODS AND MATERIALS of anal fin to tip of anal fin lobe. DEPTH OF CAUDAL PEDUNCLE. Least Except for total length, all measurements depth of the caudal peduncle. were taken from point to point (straight line) on the left side of the fish (with mouth WIDTH OF CAUDALPEDUNCLE. Width of closed). Dial calipers were used for measure- the caudal peduncle at the point of least ments up to 185 mm with use of a wooden depth. meter stick above this size. We follow LENGTH OF CAUDAL PEDUNCLE. Length Smith-Vaniz and Staiger (1973) in using of the caudal peduncle from the point of WILLIAMS AND VENKATARAMAN1: SYSTEMATICS OF CARANGOIDES 503 origin of last anal-fin ray to posterior margin mediately in front of the first dorsal spine of the hypural bones. and which has often been referred to, and HEAD LENGTH (H). From tip of snout to counted by, various authors as a small pro- posterior margin of the fleshy opercular cumbent spine. membrane. ANAL FIN COUNT includes anal spines, if ORBIT. Horizontal diameter of the bony externally visible. No dissections of dorsal or orbit located (if necessary) by insertion of anal spines were made for purpose of counts dissecting needles. although radiographs were used to confirm INTERORBITAL WIDTH. Least distance meristic characters counted externally. The measured across the top of the head between last dorsal and anal fin soft rays are divided the bony orbits. to the base but have a single pterygiophore, SNOUT LENGTH. Least distance between and thus are counted as single rays. tip of snout and bony orbit. PECTORALFIN COUNT was normally made POSTORBITAL LENGTH. From posterior on the left side, though counts were made on edge of bony orbit to posterior margin of both sides in a small number of specimens to the fleshy opercular membrane. permit estimation of bilateral variation. The UPPER JAW LENGTH. From tip of snout first element at the dorsal margin of the to posterior end of maxilla. pectoral is considered a spine. SUBORBITAL DEPTH. Least distance be- PELVIC FIN COUNT was not made as it is tween bony orbit and dorsal margin of max- invariably I, 5 in carangid fishes. illa. COUNTS are for lateral gill SNOUT TO Dl' Length from tip of snout to rakers on the first arch, normally on the left origin of first dorsal fin. side. The raker at the junction of the upper SNOUT TO D~. Length from tip of snout to and lower limbs (epibranchial and cerato- origin of second dorsal fin. branchial) is included in the lower limb count SNOUTTO A~. Length from tip of snout to as the major part of the base of the raker is origin of anal fin. over the ceratobranchial. Rudimentary gill SNOUT TO Pl' Length from tip of snout to rakers, with the base width (lateral) of the dorsal point of origin of pectoral fin. raker equal to, or less than, the raker length, SNOUT TO P2. Length from tip of snout to occur at the anterior ends of the upper and anterior point of origin of pelvic fin. lower limbs and these arc included in the

BASE D2• Length of the base of the second counts, though differentiated viz. ii, 7 + 19, dorsal fin. iv = 32, only on original data sheets.

BASE A2• Length of the base of the second CURVED . Scale count is anal fin. the number of pored scales in the curved CURVEDLATERALLINE (CLL). Chord of lateral line; it is rare in Carangoides to find arc from first pored scale to junction with scutes in the posteriormost part of the curve, straight section of lateral line. but if present they are differentiated from the STRAIGHTLATERALLINE (SLL). Straight scales although counted in the total elements section of lateral line from junction with for the curve. curved section to posterior point of last lat- STRAIGHTLATERAL LINE. Scute count is eral line scute or scale on cauda] base. the number of scutes in the straight lateral MEAN LATERAL LINE RATIO (MLLR). line. A scute is a modified lateral line scale that is enlarged, usually thickened, and has SLL right SLL left / 2 its posterior margin terminated in a pointed CLL right CLL left + spine or an apex with the angle of the scute DORSAL FIN COUNT does not include the formed by this margin being about 110° or antero-dorsal projection of the third supra- less (Berry, 1959). In addition, lateral line neural, which is often externally visible im- scutes, especially in Carangoides, are pre- 504 BULLETINOF MARINESCIENCE,VOL.28, NO.3, 1978 ceded or followed by normal, or only par- (TI). Other collections from Mozambique, tially modified, lateral line scales. The ante- Malaysia, and the Philippines made under SI rior scales are counted from the junction of 3818 are also uncatalogued and are desig- the CLL and SLL to the first scute and the nated by geographic location and station posterior ones from the last scute to the last numbers (if used). Collections from the lateral line scale before the end of the hypural International Expedition plate (line of dorso-ventral flexion of tail (IIOE) cruises of the RjY ANTON BRUUN when bent sideways). Counts are thus re- are also uncatalogued and are listed by cruise corded as: scales + scutes + scales = total and station number. The material examined elements; viz. 8 + 24 + 6 = 38. sections include only specimens examined by Abbreviations used to denote institutions the authors (or associates); localities are cited are as follows: abbreviated. ANSP-Academy of Natural Sciences, Carangoides malaharicus Philadelphia (Bloch and Schneider), 1801 BMNH-British Museum (Natural His- (Fig. 1, Table 1) tory), London Scomber ma/abaricus Bloch in Bloch and BSRM-Biological Station Reference Mu- Schneider, 1801: 31 (India); Ca/"{/ngoides ta/am- seum, Porto Novo (Parrangipettai), India parah Bleeker, 1852: 64 (East Indies); ?Caranx micraspis Kner, 1868: 321, pI. 5, fig. 13 (Sunda CAS-California Academy of Sciences, Straits); Carallx impudiclIs Klunzinger, 1884: 99 San Francisco (Red Sea); Carangoides rectipillllis Williams, 1958: MNHN-Museum National d'Histoire 40 I, pI. 10, fig. 13 (Zanzibar); Carangoides 1'110111- boides Kotthaus, 1974: 40, figs. 316, 3]7 & 321 Naturelle, Paris (East Arabian Sea). NHMV-Naturhistorisches Museum, Vi- Diagnosis.-A carangid fish of the genus enna Carangoides with the naked area of the RMNH-Rijksmuseum van Natuurlijke breast extending anteriorly above the pec- Historie, Leiden toral fin base and having a total of 32-38 SU-Stanford University, Division of gill rakers on the first gill arch. Systematic Biology (collection transferred to CAS). Description (based on 310 specimens 70- TABL-Southeast Fisheries Center, Na- 240 mm FL).-Body measurements ex- tional Marine Fisheries Service, Miami pressed as percent of fork length: total (formerly Tropical Atlantic Biological Lab- length 107.4-124.3; standard length 89.2- oratory; collections of carangid fishes trans- 95.9; head 27.3-33.0; depth, Dl to P~ 37.2- ferred to CAS). 44.3, D~ to A~ 39.8-48.4; orbit 7.3-10.8; ZMUH-Zoologisches Museum, Univer- interorbital 4.9-7.0; snout 7.7-10.8; post- sity of Hamburg. orbital 8.8-12.4; suborbital 2.3-5.2; upper Uncatalogued California Academy of Sci- jaw length 10.5-13.3; maxilla width 3.4-4.7; ences specimens collected under the auspices pectoral fin 25.8-36.2; pelvic fin 4.4-12.6; of the George Vanderbilt Foundation (GVF) height Dt spine 6.3-16.0; height D~ fin are reported under their register and station lobe 11.4-21.4; height A~ fin lobe 11.3- numbers. Most collections from Ceylon 19.0; snout to Dl 35.8-42.4; snout to D~ under Smithsonian Institution Foreign Cur- 49.7-56.8; snout to A~ 49.5-57.4; snout to rency Grant 3818 (SI 3818) are uncatalogued PI 26.5-32.4; snout to P~ 33.1-38.3; base and listed by station number. Abbreviations of D~ fin 35.8-41.4; base of A~ fin 32.4- refer to the following collectors listed in 38.9; CLL 25.8-41.8; SLL 21.1-33.0. chronological of their visits to Ceylon: Dorsal fin VIII + 1,20-23; anal fin II + F. I. Schwartz (FIS); W. F. Smith-Vaniz 1,17-19; pectoral fin 1,18-19. Lateral gill (SY); P. C. Heemstra (PCH); C. C. Koening rakers on first arch 8-12 + 21-27 = 32-38; (CCK); T. R. Roberts (TRR); T. Iwamoto lanceolate easily seen on opening fish's WILLIAMS AND VENKATARAMANI: SYSTEMATICS OF CARANGOlDES 505

band on tongue, a roughly triangular patch on vomer and in narrow bands on palatines. Some scales on cheeks and postorbital, rest of head naked. Breast naked ventrally and laterally from isthmus almost to lateral line origin, but with very small scaled area ante- riorly at level of pectoral fin base (see Fig. 2a). Posterior margin of the naked area from the upper margin of operculum around axil Figure 1. Carallgoides malabariClis CAS 16498, of pectoral fin then diagonally downwards, 224 mm FL, Ceylon, 30 August 1969. and posteriorly, above pelvic fin, at a varying but usually small distance above ventral sur- face as far as origin of second anal fin. The mouth. CLL scales 54-71; SLL 19-36 weak naked area posterior to pelvics restricted scutes, and 31-55 total elements; MLLR to a varying degree (see Fig. 2a) and may 0.53-1.00. be almost isolated from rest of scaleless part Body strongly compressed (Fig. 1). Adi- of breast, or even absent in some cases. pose eyelids weak, forming rim around eye. Second dorsal and anal fins with low to mod- Mouth cleft below eye level; slightly oblique. erate falcate lobes anteriorly (increasing with Posterior edge of maxilla position ranges size); tips of rays frequently extend beyond from vertical through front edge of eye to edge of fin margins (0.20-0.25 of ray length). center of pupil. Jaws highly protrusible, Basal scaly sheath to fins especially evident lower jaw longer. Small villiform teeth in in large fish. Pectoral subfalcate in small bands in eaeh jaw; anteriorly some outer specimens to falcate in large ones; pelvics teeth conical; bands narrow posteriorly. short. Caudal deeply forked; bilateral paired Minute viJliform teeth in a central narrow caudal keels present in large specimens.

Table 1. Meristic characters of Carallgoides malabaricus and C. talamparoides

C. malabar;clls (70-240 mm FL) C. talamparaides (79-281 mm FL)

Character Range :i II Range :i II ... ---.--- D" soft rays 20-23 21.75 296 20-23 21.90 78 A" soft rays 17-19 17.88 296 17-19 17.96 76 Lateral gillrakers on first arch upper 8-12 9.94 269 6-9 7.72 65 lower 21-27 25.01 310 19-22 20.63 69 total 32-38 34.91 271 27-31 28.58 65 CLL scales left 54-70 63.14 151 60-80 70.34 35 right 55-71 64.00 67 SLL scutes left 19-36 26.76 148 20-32 26.29 28 right 18-35 27.74 139 SLL total elements left 32-52 42.63 139 32-52 39.57 28 right 31-55 42.77 134 Mean lateral line ratio (MLLR) 0.53-1.00 0.75 247 0.52-0.84 0.63 76 .'-----" 506 BULLETIN OF MARINE SCIENCE, VOL. 28, NO.3, 1978

gascar, Mozambique to Kenya, Red Sea, Arabian Sea (off of Oman, Iran, Pakistan), west and southeast coasts of India, Ceylon, northwest of Burma, Straits of Malacca, Gulf of Thailand, Java-Sumatra. Geographic Variation.- The principal me- ristic character used to separate C. mala- baricus from C. talamparoides is total gill raker count on the first gill arch. The same character has been used to compare popula- tions of C. malabariclls from five locations- Gulf of Thailand (1), Burma/Malaysia (2), South India/Ceylon (3), Arabian Sea (4), and East Africa (5). Initial examination of data suggested a possible cline from east to west (Gulf of Thailand to East Africa) and this has been examined statistically. The mean total gill raker counts for popu- lations were subjected to an analysis of vari- ance (ANOVA) to test the hypothesis Ho: III = 112 = 113 = JL4 = Ilr. (Table 2); the hypoth- esis was rejected at P < 0.0005. The data were then subjected to a Student-Newman- C.talamparoides Keuls (SNK) multiple range test with unequal sample size with the following results (Table Naked (scaleless) area of breast 2): (1) The Gulf of Thailand population is Usual outline different from that in the other areas; (2) The Burma/Malaysia population is different from ---- Variants the populations in East Africa and the Figure 2. Extent of naked (scaleless) area of Arabian Sea, but similar to that in South breast in (a) C. l11alabariclIs and (b) C. talal11- India/Ceylon; (3) The population from paroides. South India/Ceylon is different from those in East Mrica and the Arabian Sea, and; (4) The populations from East Africa and the Color (preserved).-Head and body uniform Arabian Sea are similar. brownish, darker above, more silvery below. Large conspicuous dark opercular patch Overall III oF fJ.2 = fJ.:l f JL4 = fJ.r.. often with the central area silvery or white (may be consistent artifact of preservation). Thus, it appears that there is an east to Conspicuous small black spot at caudal fork west gradient in gill raker counts, the lower (tips of central caudal rays) and more diffuse count in the Gulf of Thailand population dusky patch on upper caudal peduncle at possibly being the primitive condition in the base of caudal lobe; trailing edge of caudal species. fin dusky. Second dorsal and anal fin mem- Material Examined. branes distally dusky to black, proximally TYPE SPECIMENS: Carangoides lalamparall yellowish-pale. Dark axillary patch to pec- Bleeker, lectotype RMNH 6098 (148), Batavia, toral which is pale-hyaline; pelvics whitish. Java; Carangoides rectipinnis Williams, holotype BMNH 1958.3.26.1, Zanzibar, East Africa. Distribution.-Based on types and present OTHER SPECIMENS. Mozambique, 19°09'S, 36° collections from: the west coast of Mada- 20'E, International Indian Ocean Expedition, R/V WILLIAMS AND VENKATARAMANI: SYSTEMATICS OF CARANCOIDES 507

ANTON BRUUN,Cr. 8, Stn. 403A(IIOE-AB8-403A), 29 specimens (110-178 FL). Malagasy Republic, 16° II'S, 43°53'E, nOE-AB8-409F, 4(128-138). Mozambique, Polana, SI-3818 (no station number) uncat., 7(120-143). Iran, HOE-AB4B-248A, 7 (156-223); 253A, 5(189-207). Pakistan, 1I0E- AB4B-228A, 5(99-145); 229A, 16(131-183); 237A, 22(126-169); 240A, 5(166-198); 242A, 11 (146-172); 275A, 2(161-160 SL); 276A, 30(119- 174); 277A, 9(128-168). Northwest India, nOE- AB4B-206A, 17(142-205); 210A, 11(124-149); 226A, 19(122-171); unknown AB4B, 1(167). Southwest India, Kerala, Yizhinjam, 8°22'N, 76° 58'E, FHB 66-50, 4 ( 134-178). Southeast India, Porto Novo (Parangipettai), BSRM uncat., 75(70- Figure 3. Cal'allgoides ta/amparoides CAS 40646, 167); Tamil Nadu (Madras) TABL FHB 66-13, 0 243 mm FL, Ceylon, 22 March 1969. 5(I 38-157); Royalpuram Beach, 13°07'N, 80 20'E, FHB 66-7, 5(144-152); Tranqueber TABL FHB 66-26, 1(175). Ceylon, SI 3818, CCK 69-60, 2 (125-128); PCH 69-223, 8(133-223); TRR 70-1, Dorsal fin VIII + 1,20-23; anal fin II + 10(154-212). Northwest Burma, 20027'N, 92°20' E,1I0E-ABI-47, 14 (89-118). West coast of Thai- 1,17-19; pectoral fin 1,18-21. Lateral gill land 9°13'N, 97°51'E, UOE-ABI-Hydro 20, 7 rakers on first arch 6-9 + 19-22 = 27-31; (157-171). Malaysia, N. Penang, 5°41'N, 100°11' lanceolate, easily seen on opening mouth. E, stn. 69-79M, SI 3818uncat., 20(100-133). Gulf of Thailand, NAGA Exped., GYF 2160, 6(123- eLL scales 60-80; SLL 20-32 weak scutes, 142); GYF 2566(135); GYF 2269(135); GYF 32-52 total elements; MLLR 0.52-0.84. 1565(182). Body strongly compressed (Fig. 3). Rest of description as for C. malabaricus unless Carangoides talamparoides Bleeker, 1852 otherwise indicated. Posterior edge of (Fig. 3, Table 1) maxilla position from vertical through just Cal'allgoides ta/amparoides Bleeker, 1852: 91 in front of pupil to posterior edge of pupil. (Sumatra); ?Carallgoides gibber Fowler, 1904: Extent of naked area of breast ventrally and 512, pI. 15 upper left (Sumatra). posteriorly is variable (Fig. 2b); in about Diagnosis.-A carangid fish of the genus 70% of specimens examined posterior naked Carangoides with the naked area of the section is separate from anterior section, breast extending anteriorly above the pec- while other two states, (a) continuous with toral fin base and having a total of 26-31 anterior sections and (b) absent, are about lateral gill rakers on the first gill arch. equal in occurrence (15%). Description.-(based on 79 specimens 79- Color (preserved).-As for C. malabaricus. 281 mm FL).-Body measurements ex- pressed as percent of fork length: total Distribution.-Based on type and present length 108.3-119.4; standard length 89.2- collections from: Arabian Sea (off coasts of Oman and Pakistan), northwest and south- 95.2; head 29.1-34.3; depth, D1-P2 37.6- east coasts of India, Ceylon, Sumatra, Gulf 50.5, D2-A2 42.0-52.9; orbit 7.4-10.8; interorbital 5.9-7.2; snout 9.3-13.6; post- of Thailand, Philippines and Borneo. orbital 9.5-13.4; suborbital 3.3-6.2; upper Geographic Variation.- Total gill raker jaw length 11.7-14.1; maxil1a width 3.4-4.5; counts were used to compare populations of pectoral fin 24.0-39.8; pelvic fjn 7.5-14.4; C. talamparoides from three locations: Gulf height Dl spine 5.2-14.0; height D2 fin lobe of Thailand (1), South India/Ceylon (3), 13.2-26.3; height A2 fin lobe 11.3-24.5; and Arabian Sea (4). Initial examination of snout to Dl 39.5-45.0; snout to D2 53.8- data suggested a possible gradient from east 61.8; snout to A2 52.6-59.6; snout to P1 to west (Gulf of Thailand to Arabian Sea) 28.7-32.9; snout to P2 35.6-40.9; base of and this was examined statistically. D2 fin 34.3-40.1; base of A2 fin 30.6-35.3; The mean total gill raker counts for pop- CLL 35.6-42.7; SLL 21.2-31.0. ulations were subjected to an analysis of 508 BULLETIN OF MARINE SCIENCE, VOL. 28, NO.3, 1978

Table 2. Geographic variation in total gillraker counts of Carangoides malabaricus and C. talamparoides

Gulf of Burma/ South India/ Arabian East Thailand Malaysia Ceylon Sea Africa All Species Location (I) (2) (3) (4) (5) Locations .------.. ~._--- Range 32-34 32-37 33-37 32-38 33-37 32-38 Mean (i) 33.17 34.25 34.61 35.23 35.28 34.91 ~------.-- -- - C. malabaricus Std. deviation (S,) 0.75 1.52 0.93 1.13 1.13 1.17 Std. error (Si') 0.31 0.34 0.10 0.10 0.22 0.07 Number (n) 6 20 93 122 26 267 Range 27-30 27-30 28-31 27-31 Mean (i) 28.07 28.45 29.07 28.44 -~_. -.- --.--- C. talamparoides Std. deviation (Sx) 0.83 0.74 0.80 0.87 Std. error (5i) 0.16 0.16 0.27 0.11 Number (n) 27 22 15 64

Underlining of means (x) refers to similarities of population values derived from the Student-Newman-Kculs multiple range tests. variance (ANOY A) to test the hypothesis DISCUSSION Ho: }kl }k2 }k3 (Table 2) ; the hypothesis = = As mentioned previously, two species have was rejected at P < 0.0025. The data were continued to be confused under the identity then subjected to a SNK multiple range test of C. malabaricus. The major similarity be- with unequal sample size with the following tween the two species is the presence of a results (Table 2): (1) The population from small naked area of the breast anteriorly the Gulf of Thailand is different from that above the pectoral fin base. There is con- in the Arabian Sea, but is similar to that in siderable variation, however, in the shape South India/Ceylon; (2) The population of the posterior, ventral scaleless area (Fig. from South India/Ceylon is different from 2). that in the Arabian Sea. The key character in separating the two Overall }kl= }k2 oF }ka species is the clear, and higWysignificant, dif- ference in the mean number of total gill Material Examined. rakers on the first arch (P ~ 0.001, 2-sample TYPE SPECIMENS. Carangoides talamparoides 't' test) (Table 1). However, two other char- Bleeker, lectotype RMNH 6099(215), Sumatra; paralectotypes RMNH 26968 (191) and RMNH acters aid in the separation of the species: 25869 (204), Sumatra; Carangoides gibber Fowler, (1) Snout to Dl is normally less than 41.25% holotype ANSP 27517(67), Sumatra. FL in C. malabaricus and greater than OTHER SPECIMENS. Oman, International Indian 41.25% FL in C. talamparoides (Fig. 4); (2) Ocean Expedition, R/V ANTON BRUUN, Cr. 4B, Snout length is shorter at a given FL in C. Stn. 267A(IIOE-AB4B-267A), 1 specimen (129 FL). Pakistan 277A 1(200 SL). Northwest India, malabaricus than in C. talamparoides (Fig. 206A, 6(145-174); 210A, 10(79-141). Southeast 5). India, Porto Novo (Parangipettai), BSRM uncaL, A third character, the ratio: 15(80-171); Madras State, l3°l3'N, 80025'E, 1(145). Ceylon, SI 3818, PCH 69-223, 2(133- Base DjSnout to D~ 169); SV 69-106, 2(173-177); Valaicheherai, CAS 40646, 1(243), uncaL, 11(204-257); FJS 69-32, is usually larger at a given fork length in C. (236). Gulf of Thailand, NAGA Exped., GVF 1541 Stn. ]83,4(107-125); GVF 2335 Stn. 60-3]8, malabaricus than in C. talamparoides, but 4(98-165); GVF 2553, 3(129-164); GVF 2160 with a much lesser degree of reliance for the Stn. 60-143, 11(139-174); GVF 2483, 3(165- 176); GVF 1532 Stn. 74, 2(184-]85); GVF 1550 separation of the species than the other char- Stn. 92, (190); GVF 90, 3(152-173); 12°N, 100° acters given above. 4'E, TABL uncaL, (166). Philippines, Lingayen Certain other counts and characters, given Gulf, 51 3818, FJS 69-84, (168). Borneo, MNHN A.6285, (160). in Table 1, indicate differences, but these WILLIAMS AND VENKATARAMANI: SYSTEMATICS OF CARANGOlDES 509

120 35

30 • C.molobaricus 110 n I1UI~ • C. malaboricus 25 n C. tolomparoides • C. talamparoides SNOUT 20 100 LENGTH {mml 15

10 90

o .' I . I I I I ' I I 1 ' I L I ' I . I I I I I 80 o 40 60 80 100 120 140 160 180 200 220 240 260 280 FORK LENGTH lmml SNOUT TO 010 Figure 5. Relationship of snout length to fork 70 {mml length in C. malabaricliS and C. talamparoides.

60 oides (x = 39.57) than in C. malabaricus (x = 42.63) (P < 0.001, d.f. 166); (4) MLLR 50 is significantly higher in C. malabaricus (x = 0.75) than in C. talamparoides (x = 0.63) (P ~ 0.001, d.f. 321). 40 Kner (1868: 321) described a small speci- men from the Sunda Straits as Caranx

30 micraspis. Unfortunately, we have been un- able to locate the holotype (and second speci-

100 120 140 160 180 200 220 240 260 280 300 men mentioned by Kner) in Hamburg FORK LENGTH (mm) (ZMUH) or Vienna (NHMV), if indeed it is Figure 4. Relationship of snout to Dl to fork length extant, and have had to rely on the original in C. malabariclIs and C. talamparoides. description and text figure. The breast above the pectoral fin base appears naked anteriorly as in both species now being discussed. In appear insufficient for species separation be- the absence of gill raker counts, the sec- cause of overlapping ranges and standard ondary distinguishing characters are "snout l . deviations Means for these counts in the to DI" and "snout length as percent FL." two species were compared by 2-sample 't' Measured from the text figure these char- tests with the following results: (1) Mean acters would identify the specimen as C. number of eLL scales is significantly higher malabaricus. However, the identity cannot in Carangoides talamparoides (x = 70.34) be verified without examining the holotype, than in C. malabaricus (x = 63.91) (P ~ hence the inclusion of C. micraspis Kner as 0.001, d.f. 184); (2) Mean number of a questionable synonym of C. malabaricus SLL scutes is not significantly different be- (Bloch and Schneider). tween C. malabaricus (x = 26.76) and C. In the lectotype (RMNH 6099) of C. talamparoides (x = 26.29) (P > 0.20, d.£. talamparoides Bleeker, it is extremely diffi- 177); (3) Mean number of SLL total ele- cult, because of deterioration and damage, ments is significantly lower in C. talampar- to ascertain the breast squamation anteriorly above the pectoral fin base (W. Smith-Vaniz,

l Variances in the samples of the two species (2-tail pers. comm. and fide Smith, 1967b), though variance ratio test) differ significantly in number of scales in CLL (P 0.50). squamation clear in the paralectotypes 510 BULLETIN OF MARINE SCIENCE, VOL. 28, NO.3, 1978

RMNH 26968 and RMNH 25869. Total ACKNOWLEDGMENTS gill raker counts, though, are ::;30, and addi- We greatly acknowledge the assistance and coop- tionally in the lectotype the snout length eration of curators and staffs of institutions housing is 12.5% FL (= C. talamparaides). How- carangid fishes, particularly W. F. Smith-Vaniz ever, conclusive evidence is now available (ANSP); W. N. Eschmeyer and T. Iwamoto from the figure in Bleeker's unpublished (CAS); M. L. Bauchot (MNNH); P. Kahsbauer (NHMV); M. Boeseman (RMNH); F. H. Berry Atlas that C. talamparaides indeed has an (TABL); and H. Wilkens (ZMUH). Special thanks area of the breast naked of scales anteriorly are due to the members of the expedition to Ceylon above the pectoral fin base. (and other locations) under the Smithsonian Insti- Sreenivasan (in press), reported two speci- tution Foreign Currency Program Grant 3818 (C. R. Robins, Principal Investigator) for the excellent mens of C. talamparoides, 84 and 108 mm quality of material collected over a large size range. FL, from the southwest coast of India. Data We also thank P. V. Sreenivasan for a copy of his received from Sreenivasan (pefs. comm.) manuscript on carangids from Southwest India and indicate the specimens have gill raker counts, completed data sheets on his two specimens of C. snout length and "snout to Dl as percent lalamparoides. Work by the senior author on Indo-Pacific FL," in the same range as for C. talampar- carangid fishes was supported by the National aides. However, in the 108 mm FL speci- Science Foundation, Grants GB-31197 and GB- men, the breast does not have a naked area 3979\. Work and advice of former UMjRSMAS' anteriorly above the pectoral fin base, as in graduate students W. F. Smith-Vaniz and P. C. Heemstra during and following the project is grate- the smaller 84 mm specimen. C. talampar- fully acknowledged; thanks also go to the following aides, as now confirmed, has a small naked graduate students; 1. Burnett-Herkes, E. Chan, M. area of the breast anteriorly above the pec- Simon, and F. Vaughan at UMjRSMAS', and toral in specimens from 70-281 mm; hence, undergraduate students K. Setbacken and D. Ke]so the identity of Sreenivasan's 108 mm speci- at UCSD" who, for varying periods, assisted in measurement of fish and data processing. Radio- men is questionable. graphs were made by P. C. Heemstra and W. F. Fowler (1904: 512) described as Caran- Smith-Vaniz, and photographs by F. H. Berry and gaides gibber a 67 mm specimen from W. F. Smith-Vaniz. The figures were prepared by Padang, Sumatra. An examination of the K. Wilson. The junior author is grateful to Pro- fessor R. Natarajan, Director, Centre of Advanced holotype indicates that: (1) The breast is Study in Marine Biology, Porto Novo, for guidance, probably naked anteriorly above the pectoral help and valuable suggestions and to the University fin base, though it is difficult to be certain Grants Commission for the award of a fellowship. because of damage, and it is unclear in the The manuscript has benefited from reviews by F. text figure of the holotype; (2) The first gill H. Berry, D. P. deSylva, W. J. Richards, and W. F. Smith-Vaniz. arch is missing on the right side, and is dam- aged on the left though it appears to have LITERATURE CITED ::;31 gill rakers (= C. talamparaides); (3) Snout to Dl is difficult to measure accurately Berry, F. H. 1959. Young jack crevalles (Carallx because of damage but is probably about species) off the southeastern Atlantic coast of 41. 8% FL ( = C. talamparoides) ; (4 ) the United States. Fish. Bull. U.S. 59: 413- 535, figs. 1-98. Snout length is 8.4% FL (= C. talampa- Bleeker, P. 1852. Bijdrage tot de kennis del' roides); (5) Counts of CLL scales and SLL Makreelachtige visschen van den Soenda- scutes could place C. gibber in either C. Mulllschen Archipel. Vert. Batav. Gen. 24: malabaricus or C. talamparaides, while the 1-93. Bloch, M. E., and J. G. Schneider. 1801. Sys- MLLR (0.74) is more typical of C. mala- tema ichthyologiae inconibus cx illustratum. barkus. Berlin, ix + 584 pp., 110 col. pIs. The damage to the holotype necessitates Chan, W. L. 1968. Marine fishes of Hong Kong. Part I. Hong Kong Govt. Press, xxii + 129 pp. the inclusion of C. gibber Fowler as a questionable synonym of C. talamparoides 'University of Miami, Rosenstiel School of Marine and Atmospheric Science. Bleeker. 3 University of California at San Diego. WILLIAMS AND VENKATARAMANl: SYSTEMATICS OF CARANGOlDES 511

Fowler, H. W. 1904. A collection of fishes from Smith-Vaniz, W. F., and 1. C. Staiger. 1973. Sumatra. J. Acad. Nat. Sci. Phila. Ser. 2 12: Comparative revision of Scombroides, Oligo- 495-560, pis. VI [-XXVIII. pUtes, Parona, and Hypacant/Ills with com- Jordan, D. S. 1917. The genera of fishes, from ments on the phyllogenetic position of Campo- Linnaeus to Cuvier, 1758-1833, seventy-five gramma (Pisces: Carangidae). Proc. Calif. years with the acce;>ted type of each. A con- Acad. Sci. 39: 185-256, 26 figs., 7 tables. tribution to the stability of scientific nomen- Sreenivasan, P. V. In press. Records of six species clature. Stanford University, University Series, of carangids from the southwest coast of India. 161 pp. [ndian J. Fish. Kliinzinger, C. B. 1884. Die Fische des Rothen Suzuki, K. 1962. Anatomical and taxonomical Meeres. Eine kritische Revision mit Bestim- studies of carangid fishes of Japan. Rep. Fac. ffiungstabellen I pp. 1-133, 13 pis. Stuttgart. Fish., Prefectural Univ. Mie 4: 43-232,61 figs. Kner, R. 1868. Folge neueI' Fische aus dem Weber, M., and L. F. de Beaufort. 1931. The Museum del' Herren Joh. Cas. Godeffroy lind fishes of the Indo-Australian Archipelago. E. Sohn in Hamburg. Sitzb. Akad. Wiss. Wien 1. Brill Ltd. Leiden. Vol. 6, xii + 488 pp., 58: 26-31, 293-356. 81 figs. Williams, F. 1958. Fishes of the family Caran- Kotthaus, A. 1974. Fische des Indischen Ozeans. gidae in British East African waters. Ann. Teil, XI. Percomorphi (4). 'METEOR' Mag. Nat. Hist. Ser. 13 1: 369-430, pIs. VI- Forsch.-Ergebnisse D (17): 33-54. XIV. Smith, J. L. B. 1965. The sea fishes of southern 1961. Uraspis wakiyai sp. nov. (Pisces, Africa. 5th Ed. Central News Agency Ltd., Carangidae) from the Western Indian Ocean, Cape Town. 580 pp., III pIs., 557 figs. with a review of the species of Uraspis Bleeker, 1967a. Studies in Carangid Fishes, No. 1855 s.s. Ann. Mag. Nat. His!. Ser. ] 3 4: 65- 1. Naked thoracic areas. Occ. Pap., Dept. 87. Ichthyol., Rhodes Univ., No. 12: 139-141. 1967b. Studies in Carangid Fishes, No. DATE ACCEPTED: March 24, 1977. 2. The identity of Scomber malabariclls BLoch- Schneider, 180 l. Occ. Pap., Dept. Ichthyol., ADDRESSES: (F.W.) Division of Biology and Living Rhodes Univ., No. 13: 143-153, pis. 32-33. Resources, Rosenstie/ School of Marine and A t- ---, and M. M. Smith. 1963. The Fishes of mospheric Science, University of Miami, 4600 Seychelles. Dept. Ichthyol. Rhodes Univ., Rickenbacker Causeway, Miami, Florida 33/49; Grahamstown, South Africa. 77 pp., 99 pIs., (K. V.) Marine Biological Station, Parangipettai 608 3 figs. 502, India.