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Diets and Metabolic Rates of Four Caribbean Tube Blennies, Genus <I>Acanthemblemaria</I> (Teleostei: Chaenopsidae)

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Diets and Metabolic Rates of Four Caribbean Tube Blennies, Genus <I>Acanthemblemaria</I> (Teleostei: Chaenopsidae) BULLETIN OF MARINE SCIENCE, 65(1): 185–199, 1999 DIETS AND METABOLIC RATES OF FOUR CARIBBEAN TUBE BLENNIES, GENUS ACANTHEMBLEMARIA (TELEOSTEI: CHAENOPSIDAE) Raymond D. Clarke ABSTRACT In order to determine the extent to which tube blennies depend on food derived from within or outside of the reef system, the diets of Acanthemblemaria spinosa, A. aspera, A. greenfieldi and A. paula were compared with food availability using plankton, benthic and gut sampling. All species fed primarily on copepods, but A. spinosa consumed calanoids and cyclopoids of planktonic origin whereas the other species consumed harpacticoids of benthic origin. This pattern correlates with the occurrence of A. spinosa in tall corals whereas the other species are found in corals close to the reef surface. Calanoids were more abundant 1 m above the reef surface than <0.5 m above the reef surface. Oxygen consumption rates were determined for the above species and Emblemariopsis pricei and E. ruetzleri. On the oxygen consumption versus body weight graph, A. spinosa shares a high slope with the Emblemariopsis species, which was three times greater than the slopes of other species of Acanthemblemaria. It is not known if the high metabolic rate of A. spinosa is a consequence of the greater food availability in its microhabitat, an adaptation for acquiring the higher quality microhabitat, or both. In any case, A. spinosa is the only Acanthemblemaria species in this study that imports nutrients to the reef by consuming plankton. Chaenopsid blennies of the genus Acanthemblemaria constitute 18 species (Almany and Baldwin, 1996) and are among the most inconspicuous diurnal fishes on coral reefs. They can be very abundant, reaching densities as high as 60 m−2 (Lindquist, 1985) and 200 m−2 (J. Buchheim, pers. comm.) and may play a significant role in the nutrient dy- namics of reef communities. They feed primarily on copepods and other small crustacea (Lindquist and Kotschral, 1987). One of the features that determines their role in the community is the extent to which their food is derived from within the reef ecosystem as compared to the water mass over the reef, but there is little information on this topic for any fishes (Sedberry and Cuellar, 1993). Studies of currents upstream and downstream of reefs have shown that in general there is a net removal of plankton and therefore an im- port of nutrients to reefs (Glynn, 1973). Many organisms, including a subset of the fish assemblages, participate in this process. Whereas the mobile planktivorous fishes have received much attention (Hamner et al., 1988; Hobson, 1991), the role of chaenopsid blennies in this process may have been overlooked. The inconspicuousness of chaenopsid blennies derives from their small size (generally 20–30 mm standard length) and their almost infaunal nature. Many species live in small cavities in hard corals created by boring organisms. Some species seem to saturate the environment, avoiding competition through precise spatial relations within and between habitats (Clarke, 1989, 1994; Buchheim and Hixon, 1992). One would expect to find a relationship between reef zone, microhabitat, and the extent to which their food is derived from the pelagic or benthic environments. This study is designed to evaluate that expecta- tion, specifically to test three hypotheses: (1) that species living furthest from the shore will consume more plankton than those living on the backreef or the shallower portions of 185 186 BULLETIN OF MARINE SCIENCE, VOL. 65, NO. 1, 1999 the forereef; (2) that the species living above the reef surface in tall corals will eat more plankton than those living on the reef surface; and (3) that metabolic rates will be higher for those species eating plankton because plankton are delivered constantly whereas benthic prey may become depleted in the vicinity of the cavities where the fish live. In addition to testing the above hypotheses, this paper clarifies the relationship between two species by showing which one has diverged from the general adaptive pattern for the genus. In St. Croix, U.S. Virgin Islands, I have previously shown that Acanthemblemaria spinosa lives higher in the corals than A. aspera, that it eats more food of planktonic origin, and that it has a higher metabolic rate (Clarke, 1989, 1992, 1996). This study builds on those observations by broadening them with more extensive sampling in a dif- ferent locality, by including sampling of food availability, and by including the diets of two additional species and the metabolic rates of four additional species. With these data, this paper shows that A. aspera is typical of other members of its genus and that A. spinosa possesses unique adaptations to occupy a different microhabitat and to feed on different food. MATERIALS AND METHODS All field work took place at Carrie Bow Cay, on the Belizean barrier reef (Rützler and Macintyre, 1982). DIET.—The design was devised to sample the potential food organisms in the same locations where the fishes feed, thus avoiding a weakness of many food habit studies (Gerking, 1994). In March 1994, I investigated Acanthemblemaria paula, A. greenfieldi, A. spinosa and A. aspera in the pavement zone, surge zone, high spur and groove zone and low spur and groove zone (Rüetzler and Macintyre, 1982; Clarke, 1994). Two species were studied in each zone, except for the low spur and groove which supported only A. aspera. Each zone was sampled twice at early (08:05–08:40), mid (13:20–13:30) and late (17:05–18:00) day for a total of 24 sample sets. As described below, a sample set consisted of: (1) a fish collection, (2) a benthic collection and (3) one or two plankton collections. (1) Five fish of each species were collected using quinaldine sulfate (see below). Immediately on surfacing, I fixed them in 10% formalin. After 2 d I placed them in 70% ethanol with a change to fresh 70% ethanol after 1 d. The complete guts were subsequently dissected out and the contents were identified to various taxonomic levels. The data for the five individuals were pooled and treated as a single sample. (2) Three pieces of coral rubble were collected by dislodging with a hammer after covering them with plastic bags. Immediately on surfacing, I fixed them in 10% formalin. Soon afterward I vigor- ously agitated the sample to dislodge the organisms and decanted the supernatant, saving the settled material in 5% formalin buffered with borax for later identification. Subsequently I randomly se- lected one of the samples, stained it with rose bengal, and identified the organisms to various taxonomic levels. The pieces of rubble were selected to resemble those locations occupied by Acanthemblemaria and although no fish were seen in the pieces before collecting, several yielded fish. The presence of fish in the samples confirms that I was collecting from locations that repre- sent the benthic food available to the fish. (3) Plankton were sampled with a diver-towed 243 μm 1:3 plankton net. The 50 cm diameter ring was formed into a 38 cm square to allow sampling from a more precise distance above the reef surface. Tows were either (a) low, ranging from the reef surface to 0.5 m above the reef surface, which is below the tops of the abundant gorgonians, or (b) high, ca 1 m (always <1.5 m) above the reef surface, which is above the tops of the gorgonians but at the level of the Acropora palmata occupied by A. spinosa. All tows were for 5 min. A current meter was not used, but as part of a different study I used the same net towed in the same way with a current meter and determined that CLARKE: DIETS AND METABOLIC RATES OF BLENNIES 187 the volume sampled in 5 min was 12.01 m3 (SE = 0.25, n = 12). Immediately on surfacing, I fixed the samples in 10% formalin. They were later transferred to 5% formalin buffered with borax. Subsequently, the samples were divided with an Alan plankton splitter (range: 1/2 to 1/32), placed in a plankton counting wheel, and identified to various taxonomic levels. The same taxonomic categories were used for gut, plankton and benthic samples. The pavement zone, which averaged ca 0.5 m in depth, was sampled differently. The plankton net was held stationary, supported with a stake, with the lower edge of the ring resting on the reef surface. The current usually kept the net filled and trailing. The net was left out for 20–30 min and sampled very different volumes from the other habitats. Current speed was not measured; it was generally weak, but it varied considerably, so the sampling effectiveness in this habitat was vari- able. METABOLIC RATES.—In March 1997, I captured the fish each morning (between 08:48 and 10:38) by squirting a 0.1% solution of quinaldine sulfate at the shelter hole and covering the opening with a test tube. The fish emerged into the tube in a few seconds. To minimize exposure to the anesthetic, I immediately transferred the fish to ventilated vials which I kept in a mesh bag, thus allowing a free exchange of seawater. Within 1 to 1.5 h of capture, I placed each fish, still in its vial, in a 300 ml BOD bottle and left it for an average of 5.2 h (range 3.5 to 8.8 h). The smallest fishes were “incubated” for the longest −1 times. O2 concentration was measured to the nearest 0.01 mg L before and after “incubation” with −1 a digital electronic O2 meter. The O2 concentrations fell to an average of 86% (5.3 mg L ) of the initial values, which is not considered to produce stress for the fish.
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