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У 17 8 5 Temporal Variation in Feeding Rhythms in a Tidal Marsh Population

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У 17 8 5 Temporal Variation in Feeding Rhythms in a Tidal Marsh Population Aquat. Sei. 66 (2004) 315-326 \t i \~7 (\jy\fj\ 1015-1621/04/030315-12 V L I t \ v ^ ¥ l n c 7 t=P C « ------------ DOi i o. 1007/s00027-004-0682-0 VLAAMS INSTITUUT VOOR U t ¿ t e ¡Aquatic S ciences e EAWAG, D M » * * 2004 FLANDERS MARINE INSTITUTE Oostende - Belgium ó 1 7 8 5 Research Article Temporal variation in feeding rhythms in atidal marsh population of the common goby (Kroyer, 1838) Henrietta Hampel1* and Andre Cattrijsse2 1 University of Gent, Department of Biology, Marine Biology Section, Krijgslaan 281, S8, B-9000 Gent, Belgium 2 Flemish Marine Institute, VLIZ (Vlaams Instituut van de Zee), Vismijn, Pakhuizen 45-52, Oostende B-8400, Belgium Received: 15 May 2003; revised manuscript accepted: 31 October 2003 Abstract. Pomatoschistus microps (Teleostei, Gobiidae) of the diel cycle is inferior in comparison with the tidal intensively uses the mesohaline marsh of Westerschelde influence on the feeding behaviour of the common goby. estuary as a nursery and foraging ground. The sampling A significant difference in foraging activity occurred be­ campaign covered the semi-lunar, diel and tidal cycles. tween the spring and neap tide. The common goby mi­ The density of P. microps and potential hyperbenthic prey grated in lower abundance into the creek during spring species in the marsh creek, fullness index, evacuation tide but foraged more intensively. At both spring and neap rates and daily ration of common goby were calculated. tide, a significant difference was found in the fullness in­ Mesopodopsis slabberi, Neomysis integer and Corophium dex between day and night. At spring tide, gobies feed volutator were the most dominant prey items in terms of more during the day, while they forage more intensively biomass. Numerically, copepods dominated the diet. Mi­ at night at neap tide. All the three cycles (tidal, diel and grating fish enter the marsh creek with a relatively empty semi-lunar) influenced the feeding rhythm of the com­ stomach and leave the marsh with a higher stomach con­ mon goby. The tidal influence is superior over the diel tent. Pomatoschistus microps seemed to feed more inten­ variation, while the explanation of the combined effect of sively during the day than the night, however the influence diel and semi-lunar cycle needs further studies. Key words. Common goby; feeding habit; tidal diel and semi-lunar cycle; marsh. Introduction The biology and ecology of Pomatoschistus microps has been intensively studied. The population structure The common goby, Pomatoschistus microps, is a small (Bouchereau et al., 1989; 1993; Pampoulie et al., 2001), gobiid fish commonly found in all European coastal wa­ breeding behaviour (Magnhagen, 1998; Pampoulie et al., ters, estuaries, fjords, salt marshes and high shore pools 2001), age-structure, growth and reproduction (Arruda (Petersen, 1919). In the intertidal area of the Wester- et al., 1993), food selection (Magnhagen, 1985) and the schelde estuary (SW Netherlands), the common goby is diet spectra of newly hatched gobies (Menher, 1992) all one of the most abundant fish species (Maes et al., 1997; received attention. Despite of these intensive investiga­ Hostens and Mees, 1999). tions, no work has been published on the influence of different cycles (tidal, diel and lunar) on the migration or on feeding rhythm of the common goby. * Corresponding author phone: +32 2 74 62 139; e-mail : [email protected]; [email protected] Three main functions have been ascribed to intertidal Published on Web: August 24, 2004 migration of fish: reproduction, avoidance of predation, 316 A. Cattrijsse and H. Hampel Short term feeding habit of the common goby and feeding (Gibson, 1993). In the Westerschelde estuary, This study aims to describe the feeding habit of the common goby uses the marsh creeks between June Pomatoschistus microps during the semi-lunar period, to and October (Cattrijsse et al., 1994). During this period, investigate the separate and combined effects of the tidal, several waves of early postlarval gobies enter the marsh diel and semi-lunar cycle on the feeding behaviour of creeks and remain there to spend their early life stage. common goby in a salt marsh creek, and to quantify the Predator avoidance may result from the huge numbers of change in tidal food consumption. juveniles that stay in small pools and seepage waters (Cattrijsse et al., 1997; Kneib, 1997). The marsh and the tidal flats must harbour favourable conditions for nesting, Materials and methods but. Cattrijsse et al. (1994) never found nests. Cattrijsse Study area et al. ( 1994) also mentioned that the common goby utilizes The studied marsh ‘Het Verdronken Land van Saeftinghe’ the marsh creeks as feeding grounds, preying mainly lies in the mesohaline part of the Westersehelde estuary upon amphipods and mysids. The common goby shifts its (Fig. 1). It is the largest estuarine brackish marsh in Eu­ feeding preference from copepods towards mysids and rope with a surface area of 2800 ha (Dijkema et al., 1984). infaunal species during its life span. All samples were collected in an intertidal creek measur­ In general, three different cycles may influence the ing 1600 m long and 4 m deep at the sampling point. Dur­ migration and feeding habits of intertidal fish species. ing tidal cycle the creek falls completely dry for at least The tidal effect is the most obvious for the movement of 5 hours. Water starts entering the marsh 3 hours before fish into intertidal areas (Gibson, 1988). The availability high water and leaves the creek 4 hours after high water. of intertidal feeding areas depends upon the tidal phase. There is no connection of any other major creek in the Several studies have investigated the tidal influence on marsh, and there are no tidal flat and pools at the mouth of the feeding behaviour of fish in the salt marshes (Weis- the creek. The channel showed a sigmoid shape close to berg et al., 1981; Rozas et al., 1988; Rozas and LaSalle, the estuary, however the creek was straight and the slopes 1990; Rountree and Able, 1992). of both banks were symmetrical at the sampling point. A second important cycle is diurnal variation. The ac­ tivity of most marine fishes is synchronised with the daily cycle of light and darkness (Gibson, 1993). The diurnal in­ Sampling fluence on the feeding activity of fish in tidal marshes has Sampling took place from a bridge spanning the creek. received some attention (Kneib and Stiven, 1978; Weis- A stow net with a 1 x 1 mm mesh size, an opening of berg et al., 1981; Cadigan and Fell, 1985; Antholz et al., 1 X 1 m and a length of 5 in was used. The net was placed 1991). Weisberg et al. (1981) argued that die! rhythms on the bottom to passively sample migrating fish and po­ would result in reduced feeding on the marsh surface and tential hyperbenthic prey species in the first metre of the an increased utilization of subtidal prey. lower water column. Hyperbenthic species were defined The semi-lunar phases form a third cyclic effect on the as animals found in the first metre of the lower water col­ migration and feeding behaviour of intertidal fish how­ umn, larger than 1 mm and capable of actively migrating ever its influence is poorly studied. Changes in abundance with the moving water. Two weights attached to the sides of fish species in marsh areas related to the semi-lunar of the frame prevented the net from being lifted by the cycle of spring and neap tides have been documented by currents, while ropes kept the net in place during sam­ Rooker and Dennis (1991) and Kneib and Wagner (1994). pling. The opening size of the net, the mesh size and the The influence of the semi-lunar cycle on the feeding ac­ thickness of the fibre were used to calculate the necessary tivity of fish has not often been investigated. Only Hamer- length of net to maintain the pressure inside the net under lynck et al. (1993) observed a semi-lunar feeding rhythm all current conditions (Tranter, 1979). This method re­ in juvenile Pomatoschistus lozanoi from a sandy beach. duced the probability that animals avoided the net. Integrated work about the influence of the tidal, diel and The net fished for one hour, was rinsed and then re­ semi-lunar cycle on the feeding habit of intertidal fish placed in the same position. After high water, the net was does not exist. turned to sample the ebb current until all water had left Marsh creeks and other intertidal habitats are believed the creek. Sampling lasted a complete tidal cycle, yield­ to play a significant role in the early life stages of many ing 3 flood (FI, F2 and F3) and 4 ebb (E4, E5, E6 and E7) fish and crustacean species as feeding grounds. A full samples on every sampling occasion. Water current and appreciation of the value of such habitats necessitates water height were measured every 15 minutes. Current more information than tidal feeding rhythms. Day-night speed was measured every 15 minutes with a current me­ changes in feeding activity and fluctuations in stomach ter placed about 20 cm above the bottom next to the net. contents over a semi-lunar cycle are needed to fully un­ Water height was measured with a ruled stake placed next derstand the functioning of such feeding grounds and to to the net in the deepest part of the creek. Temperature quantify their importance in the food web of the marsh. was recorded every hour. Aquat. Sei. Vol. 66, 2004 Research Article 317 Vttalivtgeft Hartsweert | ‘I LEGEND: Cr WTL * 5 m // / A \ \ North /T*0^ M a v .Bath t \ sea v j i .
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