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Home , Carangidae, Carangoides, Caranx

J. mar. biol. Ass. India, 1996, 38 (1 & 2) : 118 - 123

ON THE FOOD HABITS OF THE OF THE - A REVIEW

S. SIVAKAMI Cerftral Marine fislieries Research Institute, Cochilt

An attempt has been madt in this paper to review the food habits of the fishes of the family Carangidae, Carangids are pelagic carnivores feeding mainly on and fishes. such as Megalaspis cordyln, AIepes djednbn and dnyi show a prefetence for young fishes especially Stolephorus spp. and the correspondence in the season of their landings can be utilish in enhancing the of carangids preferably using Stoleplzortis spp. as baits in gears like hooks and line. Carangids exhibit variations in food preference in juvenile and adults, the former preferring planktonic crustaceans and the latter mainly juveniles. Feeding intensity was found more in juveniles and immature fish than in breeders. The carangids are observed to feed more during night probably orientating their food using their visual organs and system. This higher rate of feeding during light may be made use of in capturing them during the late hours using artificial light.

groups such as hofie (Megalaspis cordyla), scads (Decapterus spy.), and trevallies THE CARANGIDgroup of fishes is one of the ( djedaba, A. kalla, Atule mate, car an.^ much demanded food fishes of the Indian . cnrangus, Carartgoides malabaricris, Atropus These shoaling pelagic group fonns about 3% atropus and Selaroides leptole~~is).Taxono- of the annual marine fish production of India mically, these carangids, are characterised by (Bal and Rao, 1990). Being ~aughtin a variety having the first 2 anal fin spines detached from of gears such as trawlers, drift gill net, purse the rest of the fin and by the presence of seine and hooks and line, these fishes are enlarged scutes along t5e lateral line (Fischer exploited from different levels of the water and Bianchi, 1984). Some species have detached column both during day and night. As is true fiiilets and resemble tnie inackcrel. Food and with any other shoaling group of fist~es, feeding habits of various specics of carangids carangids also migrate in pursuit of better were studied by Chidainbara~nand Venkataminan feeding and breeding grounds. Food and feeding (1946), Devanesan and Chidambaram (1948), habits of these fishes therefore become Chacko (1949), Chacko and Mathew (1954), interesting and informative from the view point Datar (1954), Kuthalingam (1955 '59) Tandon, of their distribution and abundance in time and (1959, '60), Reuben (1968), James (1967), space. Considering the importance of food Sreenivasan (1974, '79), Kagwade (1971), studies in carangids, Kuthalingam (1955) has Munasinghe (1972), Belinger and Avault (1971), Venkataramani and Nataraja~i (1983, '88), rightly remarked that 'much work has to be Sivakami (1988, '95) and Shameem (1992). A done regarding the food of horse '. review of literature shows that carangids exhibit The carangid fishery along the west and preferential feeding with varying intellsity east coast of India is supported by several during different months and also during day ON THE FOOD HABITS OF THE FISHES OF TIIE FAMILY CARANGXDAE 119

and night using their sense of sight in capturing crustaceans especially Acetes spp. followed by their prey. It is also obvious that their selective fishes (Ancltoviella)while Tham Ah Kow (1950) feeding habits can be made use of in capturing found the species off Singapore strait to feed thein in gears such as hooks and line, where selectively on zooplankton. According to their preferred food items can be used as bait. Venkataramani and Natarajan (1988), juveniles This review on the food habits of carangids is of S. leptolepis off Tuticorin prefer planktonic therefore taken up with a view to suggest crustaceans while adults fed more on juvenile probable measures to enhance their fishery using fishes. Likewise, species such as A. djedaba their preferred food as a tool. is also reported to prefer crustaceans as major The author is grateful to Dr. P. S. B. R. food by Venkataraman (1960) and Kuthalingam James, former Director, CMFRI for suggesting (1955), while according to Sivakami (1988), the preference of A. djedaba is for Stolephorus the project and for the encouragement. spp., Cynoglossus spp., and Leiognatlrus spp. M. cordyla as described by Sivakami (1995), Sreenivasan (1974), Chacko and Mathew (1954) (a) Food compositiort and Tham Ah Kow (1950) subsists mainly on A review of literature shows that carangids fish juveniles followed by crustaceans, while are generally pelagic carnivores, feeding mainly Datar (1954) concluded M. cordyla to be on juveniles of fishes (Stolephonis spp., carnivorous feeding mainly on crustaceans. D. Leiognatlius spp., Sardinella spp.), crustaceans dayi, another species of carangids is found to be a pelagic carnivore, preferring fishes and (Penaeid prawns, Acetes spp., Squilla spp., crustaceans (Sreenivasan, 1979). While medium Alima larva, megalopa, Temora spp., Ponfella sized carangids were found to prefer either spp., Labidocera spp., Lucifera spp., Zoea and fishes or crustaceans as their staple diet, smaller mysis stages for prawns, crab juveniles, species such as A. kalla was found to prefer appendages of shrimps and ostracods). Food mainly pelagic crustaceans followed by items such as polychaetes, chaetognaths, algal molluscan post larval forms. That carangids matter etc formed insignificant portions of the exhibit remarkable selectivity in feeding has food spectrum. been substantiated by the fact that in spite of (b) Food preference irt different species a variety of organisms available in the plankton, they are absent in the gut contents of carangids Though a variety of food items are noticed (Shameem, 1992). in the diet of carangids, a major preference for fish food items has been reported in species Exceptions to the normally preferred food such as M. cordvla, A. djedaba and D. dayi, items have been observed in certain species of while fonn the dominant food item carangids such as Chorinemus lyserz feeding on in species such as A. kalla (Kagwade, 1971), Hilsa spp (James, 1967) and Cara~tgoides and A. atropus (Raje, 1988). Of the different mnlabaricus feeding on Euphausids (Reuben, species of fishes preferred, Stolephorus spp. 1968). This may be explained as an adjustability has been found to be highly relished by M. to the available food in the absence of the cordyla, A. djedaba and D. dayi (Sreenivasan, preferred food. 1974, '79, Sivakami, 1988, '95). While p~ferential (c) Food preference in diflererzt size groups feeding has been established in carangids, various authors have observed intraspecific The intraspecific variations observed in variations in the food preference. Tandon (1960) the food preference among carangids are observed S. leptolepis off Palk to prefer attributable to the selective feeding in different 120 S. SIVAKAMI

age groups. Thus, while Kuthalingam (1959) especially Stolephorus spp. generally coincided found that M. cordyla upto 220 mrn size group with its landings at Vizhinjam (Sreenivasan, fed on crustaceans and those above 220 mm 1974) and at Cochin (Sivakami, 1995.). In A. from offshore region fed on fishes, according djedaba, the preference for ostracods during to Sreenivasan (1974), the species below 119 July to November had a direct correlation with mm preferred crustaceans and those beyond the abundance in the plankton off Cochin this size limit fed mainly on fishes. A similar (Sivakami, 1988). Evaluating the food index selectivity for crustaceans in the juvenile stages which signifies the level of utilisation of a and fishes by adults is also reported in S. particular group of food organism by the leptolepis (Venkataramani and Natarajan, 1988), population present in the habitat under and in D. dayi (Sreenivasan, 1979). Likewise, consideration, Venkataraman (1960) observed in A. djedaba a direct preference for fish as that the occurrence of prawns in stomach food is reported in all the stomach examined contents of djedaba coincided with the within the size groups of 150-199 mm and in seasonal abundance of prawns in the inshore about 83% of the fish within the size groups region of Calicut. Likewise, the small shrimp 240-319 mm, with crustaceans forming the Acetes dispar, Copepoda and fish juveniles and preferred food item in the size group 200-239 adults whose occurrence in the stomach contents mm (Sivakami, 1988). Though the destructive of C. djedaba was in correspondence with their effect of preference for fish juveniles by abundance in the catches (George, 1953, carangids as suggested by Chacko (1949) cannot Bhimachar and Venkataraman, 1952). be ruled out, the possibility of using the preferred fish as bait in enhancing their catch (e) Feeding intensity deserves attention. Further, the preferential Season, availability of the preferred food feeding on different food items in different size items, breeding periodicity and time of fishing groups may be explained as avoiding direct are the major factors which are found to competition and also because of the rich oil influence the feeding intensity in various content available in the planktonic crustaceans carangids. While.in M. cordyla feeding intensity which is essential for the developing gonad of was better during the post monsoon months, the sub-adult stage. Another possible with lesser percentage of empty stomachs in explanation for the change in the food preference immature and juvenile fish (Srineevasan, 1974, may be related to a change in the ability to Venkataraman, 1960), D. dayi exhibited higher search out and capture larger more mobile prey feeding intensity during both the premonsoon such as fishes and a reduced ability by larger (February-May) and post monsoon months predators to catch smaller prey such as (November and December). In this species, the crustaceans. The change in the food pattern frequent occurrence of empty stomachs with between juveniles and adult carangids may also little contents is attributed to the ratio between be dependent upon the selectivity of the gill the size of the fish and the size the prey as apparatus as demonstrated in allied Scombroid cited by Allen (1935) or to the higher caloric species (Magnuson and Heltz, 1971). value of the fish diet which necessitates only (d) Seasonal variations in feeding a lesser intake as opined by Longhurst (1957) and quoted by Sreenivasan (1974). The selectivity of different food items in carangids is often found varying during different Breeding periodicity was found to have a seasons. In M. cordyla, a preference for fish profound influence on the feeding rate in ON THE FOOD HABITS OF THE FISHES OF THE FAMILY CARANGIDAE 121 .

S. leptolepis where the fish in maturing and food can be accomplished even in low light. spent recovering condition fed more than the The preference of this species to actively moving juveniles and matured groups (Venkataramani fishes like Stolepltorus spp, Leiognatltus spp. and Natarajan, 1988). InA. djedaba also, highest Gazza spp. and spp. as food feeding intensity was observed during the post indicates that light plays an important role, spawning period (Sivakami, 1988). The while consuming coloured - in animate things influence of time of fishing on the feeding without any smell points to the less developed intensity has been discussed in S. leptolepis by olfactory sense (Sreenivasan, 1974). Confirming Tandan (1960). the habit of consuming more fish during night as a character common to carangids, Hobson O) Feeding in relation to breeding (1988) stated that the possession of large eyes Food habit is considered to be one of the enable them to hunt and prey visually after factors interlinked with in fishes. dark particularly on fish shoals. While according Various species of carangids show profound to Monteifel and Radakov (1%1)., the nocturnal interrelationships with feeding and breeding predators also use olfactory sense organs and periodicity. Major carangids as a general rule lateral line receptors to locate their prey. were found to exhibit lesser feeding intensity However, absence of food at night has been during breeding season with a higher percentage observed in the stomachs of S. leptolepis whi~h of empty stomachs in mature specimens with is attributed to the influence of time of capture the immature, developing and spent ones (Tandon, 1960). observed in an activily fed condition (Kagwade, (h) Cannibalism: 1971; Venkataramani and Natarajan 1983; Sreenivasan 1974; Raje, 1988 and Sivakami Cannibalism though not frequently 1988). Nevertheless, Tandan (1960) noticed encountered in carangids is reported in A. stomachs with food in mature specimens of S. atropos (Karuna 1959, Shameem, 1992). leptolepis as has also been observed by GENERALREMARKS Shameem 1992 that strikingly in carangids stomachs of immature and spent forms are Carangid are carnivorous fishes feeding empty. mainly on planktonic crustaceans in the early stages and on fish juveniles at a later stage of (g) Diurnal variations in feeding their lifespan. They possess moderate dentition, Diurnal rhythm of feeding depends on the well developed apparatus and a distinct method by which the fish orientates its food stomach characteristic of carnivorous fishes. and is also very closely connected with the The first gill arch with long closely knit behaviour of food organisms and also the diurnal gill rakers, characteristic of scombroid fishes dynamics of the composition of the food. Light probably acts as an efficient filtering mechanism intensity appears to play an important role in in carangids although the maximum the periodicity of abundance of the different distensibility of the mouth and oesophagus also food organisms. Carangid fishes are found to would be expected to set limits on the range have a profound diurnal variations in feeding of food sizes eaten. especially in species such as M. cordyla where the percentage contribution of heavy and Carangid fishes feed selectively on a few moderate feeding was high in night samples food items despite the availability of other food than in day samples proving that search for organisms in the environment. This pattern of 122 S. SIVAKAMI

selective feeding ensures lesser expenditure of Feeding intensity in carangids was more energy which is accomplished through their in and juvenile fish with breeders shoaling behaviour, because fish in . shoals possessing more of empty stomachs. ms may 'Onsurne food than be attributed to the very high food requirement single fish would. in the young and fast growing fish with the species like M. cordyla, djedaba and nature fish possessing a distended gonad and D. dayi which contributes to the bulk of the lethargic movement tending to feed less. carangids landed, show a preference for young fishes especially Stolephorus spp. This Carangids exhibit higher rate of feeding preference with a coincidence in their landings during night which is attributable to their may be made use of in forecasting the fishery nocturnal migrations to the sea surface. This of carangids. Further, it is possible that carangids behaviour can be utilised to capture them using fishery is improved by using Sroleplrorris spp. bag nets and trawlers along Indian coast also as bait in hook and line as is practiced elsewhere as is practiced off Manila Bay (Tiews, 1968). (Sreenivasan, 1974; Sivakami, 1995). Cannibalism is considered as an adaptation The variations in food preference during where the adult fishes, when unable to feed the young and adult stage in carangids may directly on plankton feed upon their be considered as an adaptation, since fish in planktivorous young ones (Moyle, 1982). In an advanced size group by feeding on easily carangids, since the adults do not prefer a available fish food tends to save energy which plankton diet, it is probable that the phenomenon may be spared for growth and reproduction. of cannibalism is not of common occurrence.

*ALLEN K. R.-1935. The food and migration of CHIDAMDARAM,K. AND R. S. VENKATARAMAN,1946. the perch (Perca fluviatilis) in windermere J. Anim. Ecol. Tabular statement on the natural history of certain marine 4 : 264-273. food fishes of the Madras Presidency West Coast, Madras Government Publication. BAL, D. V. AND K. VIRABHADRARAO, 1990. Marine of India. TAVA Mc GrDraw-Hill *DATA&G. G. 1954. Food and Feeding of Caranx Publishing Company Ltd., New Delhi, p. 211. rottleri (Cuv. and Val.) Proc. 41st Ind. Sci. Cong. Asroc. (3), (Abstract), 181. *BELLINGERJ. W. AND J. W. AVAULTJR. 1971. Food habits of juvenile . Trachionotus carelinirs DEVANESAN,D. W. AND K. CHIDAMBARAM,1948. in Louisiana : Trans. American Fish' Sot'* loo (3) common food fishes of the Madras Presidency. Madras 486-494. me Government Publication. BHIMACHAR,B. S. AND G. VBNKATARAMAN,1952. A preliminary study of the fish populations along the FISCHERW. AND A. G. BIANCHI.(~~S.)1984. FA0 Malabar mast. Proc. Nut. Inst. Sci. India 18 (6) : 627655. Species identification sheets for fishery purposes. Western Indian (Fishing Area 51) Prepared and printed with CHACKO,P. I. 1949. 'Ihe food and feeding habit the support of the Danish International Development of the fishes of the Gulf of Manner. Proc. Ind. Acad. Agency (DANIDA) Rome, Food and Agricultural Sci., 29B : 83-97. Organisation of the United Nations, Vol. I, Carangidae: p. 1. -AND M. J. MAW, 1954. and Fisheries of the horse-mackerel of the West Coast of GEORGF,P. C. 1953. The marine plankton of the Madras. State. Conk Mar. Biol. St. West Hill, Malabar coastal waters of Calicut with observations on the Coast. (2) : 1-12. hydrological conditions. J. Zool. Soc. India, 5 (1) : 76-107. ON THE FOOD HABITS OF THE FISHES OF THE FAMILY CARANGIDAE 123

*HOBON, S. EDMUND, 1908. Predatory behaviour Swm, A. 1992. Comparative studies on the of some shore fishes in the Gulf of California. Res. Rep. feeding habits of marine and estuarine carangids Ibid : U.S. Fish. WeIdI. Sew., 17 : 1-92. 34 (1 & 2) : 262-268.

JAMB, P. S. B. R. 1967. Some observations on SIVUI, S. 1988. Observations on some aspects the fishery of Chorinemus lysen of the Rameswaram of biology of AIepes djedaba (Forsskal) from Cochin. with notes on its biology. Indian J. Fish., 11 (1) : (1%4): I bid. 32 (1 & 2) : 107-118. 268-276. fiow~~e,V. N. 1971. 'I%e food and feeding (1995) Fishery and biology of Caravgid fish Megalaspis cordyla (Linnaeus) off Cochin Ibid, 37 habits of horse mackerel, Caran* kalla (Cw. and Val.) (1+2) : 237-248. Indian J. Fish., 14 (1-2) : 85-96. KARUNA. 1959. M.SC Thesis, Andhra University. SREBNNASAN,P. V. 1974. Observations on the food and feeding habits of the tarpedo trevally, Megalaspis KWUNGMI, M. D. K 1955. lhe food of Horse cordyla (Linnaeus) from Vizhinjam Bay. Indian J. Fish mackerel Caranx djedaba : Curr. Sci., 24. (12) 416. 21 (1) : 20-28. -1959. A contribution to the life histories - 1979. Feeding biology of the seed and boding habits of horse mackerel Megalaspis cordyla Decaptems dayi Wakiya. J mar. biol. Ass. India 21 (Linn.) and Caranr mate. J. Madras Univ. B. 29 (2) : (1 & 2) : 97-102. - 79-%. TANDON,K K 1959. The food and feeding habits *Loffim~, A. R. 1957. The food of the of Selaroides leptolepis (Cw. & Val). Curr. Sci., 29 domeml fish of a West African . J. Anim. Eccl., (2) : 62-63 26 : 369-387.

MAGNUSON,J. JOHNAND JEAN, G. HELTL 1971. 1%0. Biology and Fishery of 'Choo Gill rakers apparatus and food selectivity among mackerels, Parai' Selaroides leptolepsis (Cuvier and valenciennes) I. tunas and dolphins Fislr Bull., 68 (2): 361-370. Food and Feeding habits. Indian Journal Fish., VII (1) : 82-100. Momr, B. P. lrm D.V. RADAKOV. 1971. The adaptive significance of shooting behaviour in fishes. Res. *I'~MI AH Kow. 1950. The feed and feeding Rev. Biol. 150 : 338-345. relationships of the fishes of Singapore. Straits Coloneal ofl Fish. Pub., 1 (I), 1-35. Mona, PB78R, B. 1982. Fishes: an introduction to ichthyology, Printice, Hall. Inc Englewood, Cleffs, New *'l"mws. K. 1968. 'Report to the Government of Jersey, P. 395. Philippines on marine fishery resources. Phil. Journ. Fish. 6 (2) : 134-139. *Mwm~n,N. L R. 1972. Influence of light on diurnal behaviour of carangids in Wadge Bank trawl VBNKARARAMANG. 1960. Studies on the food and fishing. Bull. Fish. Res. Stn. Srilanka (Ceylon) 231 : feeding relationships of the Inshore fishes off Calicut on (I t 2): 9-18. the Malabar coast. Indian J. Fish: 7 (2) : 275-307. Nfrcolsm, G. V. 1963. The Ecology of fishes. Academic Press, Inc. (Lnndon) Ltd., P. 263. VENKATARAMANI,V. K. 1978. Bio systematic studies in Carangid fishes of Porto Novo coast (: Ruq S. G. 1988. Studies on the Fishery and Carangidae) Ph.D. Thesis, Annamalai University, 290 pp. biology of Atropus arropus (Block and Schneider) from Veraval, Saurashtra coast, In: Symposium on Tropical *- AND R. NATARAIAN,1983. Rec. ZOO/. Marine Living Resources, Cochin, India 12-16 January Sun. India, 81 : 369-406. 1988: Abst. 53. AND P. NATARAJAN.1988, Food and REUBEN,S. 1968. A 'note on the food of Malabar feeding habits of Selaroides leptolepis val. Off Tuticorin trewlly, Carangides malabarius (Bloch and Schn.) From coast, Mataya : 14 : 53-62. the North Western Bay of Bengal, J.mar. bid. Ass. India, 10 (1) : 182-183. *Not referred in original