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MARINE ECOLOGY PROGRESS SERIES Published February 28 Mar Ecol Prog Ser 1 Die1 feeding of the chaetognath Sagitta enflata in the Zanzibar Channel, western Indian Ocean Vidar Oresland* Institute of Marine Research, National Board of Fisheries, PO Box 4, 453 21 Lysekil, Sweden ABSTRACT: The d~elfeeding cycle of Sagitta enflata, the dominant chaetognath in the Indian Ocean, was investigated, using gut content analyses, between the monsoon periods on 2 occasions, October 1995 and May 1996. Zooplankton samples were collected every second hour during a 24 h period close to a dnfting buoy in the middle of the Zanzibar Channel. S. enflata made up 64 % (October) and 77 % (May) of all chaetognaths in this study. S enflata and all chaetognaths contributed as much as 30 and 33%,,respectively, of the total 90 pm net zooplankton wet weight in October (May samples could not be weighed due to occurrence of phytoplankton). S. enflata fed most actively at night. Small copepods with a mandible width <O.Of mm (prosome length <l mm) were its predominant prey. Feeding rate and mean number of small copepods found in S. enflata were 3 times higher in October than in May. In con- trast, the estimated daily predation impact on the standing stock of small copepods (by number) was around 1 % for both periods. Thls may be a negligible predation effect consi.dering the short life-span of tropical copepods. It is suggested that predation effects are related to temperature, since a sirmlar predation impact may be important in cold polar waters where copepod life-spans are much longer. Daily cannibalistic predation impact (by number) was estimated to be between 1 and 4 % (October) and between 2 and 9% (May) using assumed maximum and minimum digestion times of 8 and 2 h. Canni- balism, therefore, may be an ~rnportantsource of mortality in S. enflata populations. KEY WORDS: Zooplankton . Chaetognatha . Sagitta enflata . Flacisagitta enflata - Copepods . Feeding . Predation . Cannibalism. Indian Ocean INTRODUCTION and polar areas, as well as from deep water. In this study, the die1 feeding of a chaetognath species was The phylum Chaetognatha contains at least 115 spe- studied through gut content analysis for the first time cies (Bieri 1991).Chaetognaths are found in all oceans, in the Indian Ocean (to my knowledge). Sagitta from polar to tropical areas, and from the surface to enflata, or Flacisagitta enflata (Bieri 1991), is a cos- several thousand meters depth (see reviews by mopolitan species in temperate and warm waters Alvarino 1965, Pierrot-Bults & Nair 1991). They can (Alvarino 1965) and occurs mainly in the upper 300 m make up a large proportion of zooplankton wet weight (Pierrot-Bults & Nair 1991). It is the dominant chaetog- and are potentially one of the main sources of preda- nath species by number in the Indian Ocean (e.g. tion pressure on the copepod community. However, Furnestin & Radiguet 1964, Furnestin & Ducret 1976, our understanding of chaetognath feeding, its variabil- Nair 1977, Andreu et al. 1989, Michel 1995) and its ity, and its consequences for predator and prey popula- feeding behaviour is therefore of special interest. tion dynamics and production is limited and biased The specimens in this study were caught during two (see reviews by Feigenbaum & Maris 1984, Feigen- 24 h sampling periods between the monsoon periods, baum 1991). following a drifting buoy in the Zanzibar Channel off Most diel feeding studies carried out until now have Tanzania, East Africa. The diet, diel feeding behav- been limited to a few species in coastal areas. There is iour, feeding rates and estimates of predation impact a general lack of studies from open oceanic, tropical on standing stock of copepods-as well as the impact of cannibalism-are presented, and the results com- pared with data from other oceans. 0 Inter-Research 2000 Resale of full article not permitted 118 Mar Ecol Prog Ser 193: 117-123, 2000 bottom (35 m) at the beginning of both sampling pen- ods. Salinity was not measured. Four zooplankton sam- ples were collected every second hour (a total of 12 sampling times and 48 samples during each 24 h period), from a 7 m long open fishing boat, within 10 to 100 m from a buoy that was drifting in a northward direction. This was done in order to reduce variability in prey and predator abundance and size frequency distribution of organisms compared with sampling car- ried out at a fixed geographical position. The buoy (40 cm high) was equipped with a small blinking light so that it could be followed during the night. A nylon parachute (1.4 m in diameter) was attached to a rope 5 m below the buoy, and a 10 kg weight was attached to the end of the rope 10 m below the parachute. The positions of the buoy were deter- ZANZIBAR mined by using a GPS. In May the parachute was CHANNEL entangled during the first hours, and the drift of the buoy was affected by the wind (Fig. 1). Samples were taken using a WP2 90 pm net (UNESCO 1968).The diameter and the mouth area of the net were 0.57 m and 0.25 m', respectively. A cali- . brated General Oceanics Inc. flowmeter (model 2030) 0 10 Km was mounted halfway between the centre of the net opening and the frame. The net was hauled in by hand, Fig. 1. Sampling area in the Zanzibar Channel and the north- with the rope kept as vertical as possible, from 1 to 2 m ward drift of the buoy close to (within 10 to 100 m) which zoo- above the bottom (35 to 50 m) to the surface at a speed plankton samples were taken every second hour during two of approximately 0.5 m S-'. Sampling often took less 24 h sampling programmes (right track = 24 to 25 October 1995, left track = 27 to 28 May 1996) than 2 min and always less than 3 min. All zooplankton samples were preserved within 2 min in 5 % formalde- hyde in seawater buffered with borax. Laboratory methods. For each sampling time, be- METHODS tween 1 and 4 zooplankton samples were sorted for all chaetognaths (all species) depending on the number of The area. The Zanzibar Channel (Fig. 1) is influ- Sagitta enflata in the samples. I wanted a minimum of enced by the northeast monsoon from about December 150 S. enflata individuals from each sampling time. to March and by the southeast monsoon from about The number of S. enflata sorted and analysed for each June to October. Between monsoon periods the sea is sampling time ranged from 147-358 in October to relatively calm. The bottom depth in the middle of the 164-623 in May. All chaetognaths appeared to be in channel increases from about 30 m in the central part good condition, with most of the fragile fins intact. Spe- to about 80 m in the northern part. Throughout the cies identification of S. enflata was made according to year, from the surface to 5 m above the bottom, the descriptions given In Bieri (1991), George (1952) and temperature varies around 27°C (26 to 30°C) and salin- Lutschinger (1993).Only ind~vidualsthat were 4 mm or ity is between 34.5 and 36 psu (Wickstead 1963). longer could be identified. Shorter individuals were Detailed knowledge of the zooplankton community is regarded as belonging to a group called 'other chaeto- still lacking from this area, although Wickstead (1963) gnaths', which may have contributed to an underesti- showed that total zooplankton abundance and biomass mation of numbers of S. enflata. in the Zanzibar Channel varies considerably during Wet weight was determined for Sagitta enflata,other the year. chaetognaths, and remaining zooplankton found in the Field sampling. Two 24 h sampling programmes analysed October samples according to procedures were carried out between the monsoon periods on given in Oresland (1990).Chaetognaths were the only 24-25 October 1995 and 27-28 May 1996 in the central large (>l cm) organisms that occurred in the samples. part of the Zanzibar Channel (Fig. 1).The temperature The May zooplankton samples were not weighed due was 27°C (from mercury thermometer readings) in to the presence of dense concentrations of phytoplank- water samples taken every 5 m from the surface to the ton. Oresland: Die1 fee!ding of Sagitta enflata 119 Lengths of Sagitta enflata individuals were mea- mixture in October and 1/4000 of mixture in May) sured (after 6 to 18 mo in formaldehyde) to the nearest were taken from each mixture. Subsequent counts of mm, under a stereomicroscope (see Conway & Robins the last 4 subsamples did not change the estimated 1991 for the effects of formaldehyde preservation on abundance by more than 3%. Plots of standard error chaetognath length). Body length was measured from (SE) against the number of subsequent subsamples the anterior tip of the head to the end of the tail, showed that the last subsamples only contributed to a excluding the tail fin. Maturity stages of S. enflata minor decrease in SE (not shown, standard deviation were classified according to the development of the [SDI is given in Table 2). ovaries (Zo 1973) as follows: Stage 1, no visible ovaries; Stage 2, ovaries visible, no mature ovum; Stage 3, 1 or more mature ovum. Individuals at Stage 3 could have RESULTS been reproducing. The microdissection technique and the analyses of Abundance and size distribution chaetognath gut content-which was inferred from identification of prey mandibles, chaetognath hooks, In this study, Sagitta enflata was by far the most com- and other hard or special prey parts-followed the mon chaetognath by number.