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Diet Composition and Feeding Habits of Atlantic Bumper, <I BULLETIN OF MARINE SCIENCE, 72(3): 675–683, 2003 DIET COMPOSITION AND FEEDING HABITS OF ATLANTIC BUMPER, CHLOROSCOMBRUS CHRYSURUS (PISCES: CARANGIDAE), LARVAE IN THE SOUTHERN GULF OF MEXICO Marina Sánchez-Ramírez ABSTRACT Chloroscombrus chrysurus (Atlantic bumper) is an ecologically important species in the southern Gulf of Mexico, as it is abundant throughout the area and contributes to the diet of a great number of species of recreational and commercial value. For this reason, changes in the size of its populations can have an effect on these fisheries. The larval stage of fishes generally has the highest mortality rate, and because mortality rate is often influenced by prey availability, this study focused on the feeding habits of Atlantic bumper larvae. Zooplankton samples were collected during five oceanographic cruises (1987– 1993) and the digestive tracts of 297 larvae, 1.03–8.07 mm SL, were analyzed. Larvae of this species fed mainly during the day, and feeding incidence was greatest for postflexion larvae (values of 100%) when the larvae capture their prey more efficiently. Larvae fed primarily on Penilia avirostris (Cladocera), copepods and nauplii, and the number and size of prey increased with larval development. Fish play a very important part in the dynamics of marine ecosystems. Year class strengths are largely determined during the larval stage because variability in the high mortality rates during the early life history stages may result in large differences in the magnitude of recruitment and abundance of the adult population (O’Connell and Raymond, 1970; Cushing, 1975; Laurence, 1977; Ivlev, 1961 vide in Last, 1978; Houde, 1978; Hunter, 1981; Lasker, 1981; Hjort, 1914, 1926 vide in Towsend, 1983; Hjort, 1914 vide in Hempel, 1984). The survival of larvae depends upon such factors as transportation by currents, ad- equate food and the evasion of predators (Ahlstrom and Moser, 1976). The abundance of food directly affects growth and mortality rates (Ivlev 1961 vide in Last, 1978; Waytt, 1972). Well-fed organisms are more robust, active, and less subject to predation and dis- eases. Their ability to search for food also increases (May, 1974; Laurence, 1977; Last, 1978; Alvariño, 1985). McGurk (1986, 1987) proposed that temperature controls the rate of development of larvae, and may control the duration of the period within the interaction between preda- tion mortality and patchiness operates, but Pepin (1991) also pointed out that high growth rates require high ingestion rates, which in turn require that greater numbers of prey items be encountered and results in increased encounters with predators. Because feeding is important to mortality of the early life stages of fishes, information on the diet of fish larvae is necessary in population studies. The Atlantic bumper (Chloroscombrus chrysurus) is a typical and ecologically domi- nant species in the fish community of the southern Gulf of Mexico (Yáñez-Arancibia and Sánchez-Gil, 1986). It spawns inside the 40 m isobath throughout the year, with a maxi- mum during the warm spring–summer season (Flores-Coto and Sánchez-Ramírez, 1989). The Atlantic bumper constitutes an important food source for many fish of recreational and commercial value, thus the abundance of Atlantic bumper may ultimately affect the biomass of other fishes (Shaw and Drulliger, 1990). 675 676 BULLETIN OF MARINE SCIENCE, VOL. 72, NO. 3, 2003 The purpose of this study was to determine the diet and feeding habits of Atlantic bumper larvae during development and among seasons, in the southern Gulf of Mexico. METHODS FIELD METHODS.—The study area was located in the southern Gulf of Mexico, at 18°06'– 21°00' N and 90°26'–97°20' W (Fig. 1). Zooplankton samples were collected on board the R/V JUSTO SIERRA during five cruises between 1987–1993 (Table 1). Zooplankton sampling consisted of a double oblique plankton tow following a circular trajectory using a Bongo net with 333 and 505 µm mesh sizes. During the Mopeed V cruise, sampling was carried out at several depths (2, 10, 20 and 45 m) throughout the water column, with an opening-closing net with a 75 cm diameter mouth and a 505 µm mesh size netting. Samples were fixed with 5% formalin neutralized with sodium borate. LABORATORY ANALYSES.—The Atlantic bumper larvae were separated, identified, and preserved in 70% ethanol. Larvae that were not twisted and curled were selected for the diet analysis. They were measured for standard length (SL) or notochordal length when the flexion of the notochord was not complete, and for the length of the upper jaw, under a dissecting microscope with a micrometric ocular and a 0.01 mm precision. The C. chrysurus larvae were grouped by stage of development into preflexion (3.0 mm SL), flexion (3.1–4.0 mm SL) and postflexion (4.1–9.0 mm SL), as established by Sánchez- Ramírez and Flores-Coto (1993). The digestive tracts of 297, 1.30–8.07 mm SL, Atlantic bumper larvae collected during the four climatic seasons were dissected in a damp cham- ber using very fine needles. The contents were identified to the lowest possible taxon and counted. The prey were measured for width (short axis) under a microscope with a micrometric ocular. DATA ANALYSES.—The feeding incidence was calculated as the percentage of larvae with food in the digestive tract with respect to the number of larvae analyzed for the day (6:40–17:40 hrs) and night (20:36–5:58 hrs) periods, as well as for each stage and cli- matic season. The index of relative importance (IRI) proposed by George and Hadley Figure 1.- Study area. SANCHEZ-RAMIREZ: FEEDING OF ATLANTIC BUMPER LARVAE 677 Table 1. Summary of research cruises to the southern Gulf of Mexico. Ceruise Dnat Sseaso Number of Station O7GMEX II 2r7 July– 5 August, 198 s8umme 6 O7GMEX III 2l8 November– 5 December, 198 f4al 4 O8GMEX V Arugust 1– 9, 198 s6umme 7 M2OPEED II Jgune 20– 24, 199 s1prin 2 M3OPEED V Frebruary 12– 18, 199 w0inte 2 (1979) and modified by Towsend (1983) was calculated to analyze the importance of each food category in the diet of the Atlantic bumper larvae at each developmental stage and for each climatic season: 100()Xa IRIa = n ∑ Xa a=1 where IRIa= Index of relative importance for food item a, Xa = % frequency of occurrence + % number of food item a, and n = number of different food items in the larvae of each sample. The size of the mouth was calculated using the index proposed by Shirota (1970) to analyze the relationship between the size of the mouth and the size of the injested prey: MUJ= 2() where M = relative mouth size (mm) and UJ = length of the upper jaw (mm). RESULTS AND DISCUSSION FEEDING INCIDENCE.—Chloroscombrus chrysurus demonstrated a clear tendency to feed during the day. Of the 181 larvae collected during the day (6:40–17:40 hrs) feeding inci- dence was 79.6%, whereas of the 116 collected during the night (20:36–5:58 hrs) feeding incidence was only 12.9% (Table 2). As Hunter (1981) mentioned, marine fish larvae are visual feeders. The feeding is confined to daylight hours as indicated by analysis of other species (Arthur, 1976). In nearly all species of fish the visual cells comprise only cones in the larval stage and there is no movement of retinal masking pigment when the larvae are subjected to changing conditions of illumination (Blaxter and Staines, 1970). A pure- cone retina is thus adequate for first feeding, and experiments have demonstrated that light is required for feeding by many species, at least in the early larval stages (Blaxter, Table 2. Diel feeding incidence of Chloroscombrus chrysurus larvae collected in the southern Gulf of Mexico. N = number of digestive tracts analyzed. HNdR Wdith foo Wdithout foo % with foo 6:40−117:40 148 174 3679.5 20:36−065:58 151 11130 12.9 678 BULLETIN OF MARINE SCIENCE, VOL. 72, NO. 3, 2003 1986) when they need light to locate their prey, which they do mainly by sight. This has been observed in the majority of fish larvae (Hunter, 1981) including clupeids such as Brevoortia tyrannus (June and Carlson, 1971), B. patronus (Govoni et al., 1983) and Sardinops sagax (Arthur, 1976), engraulids such as Engraulis mordax (Arthur, 1976), scombrids such as Katsuwonus pelamis, Thunnus maccoyi and T. alalunga (Young and Davis, 1990), carangids such as Trachurus declivis (Young and Davis, 1992) and sciaenids such as Leiostomus xanthurus and Micropogonias undulatus (Govoni et al., 1983), among others. The digestive tracts of Atlantic bumper larvae smaller than 3.0 mm SL contained food, in contrast with another carangid, Trachurus symmetricus, whose larvae must be > 3.0 mm SL to start feeding (Arthur, 1976, 1977). Although these are not closely related spe- cies, it is possible that this difference in size at the start of feeding is characteristic of organisms of lower latitudes being able to develop faster (Sánchez-Ramírez and Flores- Coto, 1993), and capture prey more efficiently at smaller sizes. Size at the time of hatch- ing differs between the two species as well: 0.76–1.02 mm SL for Atlantic bumper (Sánchez-Ramírez and Flores-Coto, 1998); 1.9–2.4 mm SL for the Jack mackerel, T. symmetricus, (Alhstrom and Ball, 1954 vide in Watson et al., 1996), which in this case results in the larvae smaller development at 3.0 mm. On the other hand, Arthur (1976) suggested that the high feeding incidence may be related to a looped gut in the larvae of these species, which reduces regurgitation at the moment of capture and preservation. Thus, as the smaller larvae of the Atlantic bumper have a more markedly looped gut than those of the jack mackerel, regurgitation may be reduced and the presence of food more obvious in Atlantic bumper larvae.
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