A laboratory-based comparison of osmoregulatory ability at different water temperatures and salinities in the stenothermic isopod Saduria entomon (Linnaeus, 1758) and the eurythermic amphipod Corophium volutator (Pallas, 1766) from the Baltic Sea

The aim of the investigation was a comparison of osmoregulatory ability at different water temperatures and salinities by the stenothermic isopod Saduria entomon (Linnaeus, 1758) and the eurythermic amphipod Corophium volutator (Pallas, 1766) from the Baltic Sea. The experiments were performed under laboratory conditions at different water temperatures (5, 10 and 15?°C) and salinities (3, 6.6, 15 and 25 PSU for S. entomon and 3, 6.3, 15 and 25 PSU for C. volutator). The osmotic concentrations of the crustaceans’ haemolymph were determined using the melting point method. Temperature had a statistically significant effect on the osmoregulation in S. entomon, but had generally no statistical influence on the osmoregulation in C. volutator. This physiological information regarding adaptation provides a basis for predicting the distribution of these species in changing environments. The implications of the results for the comparative adaptability of the two species are considered.     

Behaviour and time allocation of the mud shrimp, Corophium volutator, during the tidal cycle: a laboratory study

Despite the numerous ecological and behavioural studies, relatively little is known about the behavioural dynamics of Corophium volutator during the tidal cycle. In the present study, the behaviour of C. volutator was observed in the laboratory in a tide-simulated aquarium, and time allocation of the different activities was quantified. Overall, the surface activity was low and showed a steep decline after submersion. A clear tide-based cyclic pattern was found, with a boost of activity immediately after first submersion, continuing during submersion, and shifting over a period with decreasing activity towards almost total inactivity at the end of emersion.     

Mass mortality in two common soft-bottom invertebrates,Hydrobia ulvae andCorophium volutator-the possible role of trematodes

Two littoral macrofaunal invertebrates,Hydrobia ulvae (Prosobranchia) andCorophium volutator (Amphipoda) suffered mass mortality on an intertidal mudflat in the Danish Wadden Sea in May–June 1990. Dissection of collectedH. ulvae individuals revealed a considerable increase from March to May in numbers of infected individuals by microphallid trematodes that useH. ulvae andC. volutator as first and second intermediate host, respectively. The numbers of infested snails were hereafter reduced by an amount equal to the observed mortality rate of snails. At the same time, theC. volutator population became extinct. Since other conceivable mortality factors could be ruled out, parasites are suspected to be the causative agent. Apart from the expected effects on potential predators by the decline in the two invertebrate populations, the benthic community changed and destabilization of the substratum occurred probably because of the die-off inC. volutator. Meteorological data suggest high temperatures as a triggering factor of the massdevelopment of the studied trematodes.     

Differential emigration causes aggregation of the amphipod Corophium volutator (Pallas) in tide pools on mudflats of the upper Bay of Fundy, Canada

We investigated mechanisms responsible for the formation of spatial patterns in a dominant macro-invertebrate, the burrowing amphipod Corophium volutator, in relation to tide pools on intertidal mudflats of the upper Bay of Fundy, Canada. A field survey of 3 different mudflats showed that density of C. volutator at low tide was consistently higher inside than outside tide pools, and a manipulative field experiment indicated that C. volutator selected tide pools over the adjacent emerged areas. Differential survival in immersed and emerged areas at low tide did not explain the observed pattern, because short-term survival of amphipods was not affected by immersion at low tide. As well, immigration rates of C. volutator into immersed and emerged areas at low tide were similar after one day, so preferential settlement did not explain aggregation of amphipods. However, marked C. volutator departed from experimental areas that were immersed at low tide at a slower rate compared to emerged experimental areas. Thus, formation of the spatial pattern observed in the field (higher densities inside tide pools) is likely the result of an effect of immersion at low tide on proportion of individuals emigrating from an area every day.     

Zur Nahrungsaufnahme vonTetrastemma melanocephalum (Nemertini)

The proboscis of hoplonemerteans is armed with a stylet apparatus, which is used for capturing prey.Tetrastemma melanocephalum (Johnston) is a common hoplonemertean in the littoral of the Baltic Sea and the North Sea. It lives in the phytal and on sand- and mudflats and occurs in large numbers in theCorophium belts in the waddenseas.T. melanocephalum feeds on arthropods, mainly on copepods and amphipods. The prey is caught with the proboscis, penetrated by the stylet and grows weak (reduction of movements) within a few seconds; it is then sucked out. During the summer, a 15–35 mm long nemertean captures about 3 specimens ofCorophium volutator of 3.5–6 mm length per day. In theCorophium belt at Sahlenburg, German Bight, North Sea, 29 nemerteans/0.25m² were found; they feed mainly onC. volutator, so that on average more than 10 000 specimens ofC. volutator/m²/month are sucked out by these nemerteans. Hence,T. melanocephalum is an important consumer ofCorophium.     

Sediment toxicity testing of lightly contaminated dredged material in The Netherlands

In the Netherlands, the decision to permit the dumping of dredged material into the sea depends on the concentrations of about 33 chemical compounds (Anonymous, 1994a). This evaluation of dredged material quality can be improved by the use of bioassays in its assessment. In this note the use of bioassays in the evaluation of lightly contaminated marine dredged material from the Netherlands is illustrated with preliminary results of bioassays with Corophium volutator and Crassostrea gigas. It is concluded that the inclusion of bioassays in the assessment of lightly contaminated sediment is useful.     

Distribution and life cycle of the North American Spionid polychaeteMarenzelleria viridis (Verrill, 1873) in the Ems estuary

In 1983 the first specimens of the North American spionid polychaeteMarenzelleria viridis were found along the European mainland shore in the Ems estuary. Since then, this polychaete has spread over several estuaries around the North Sea and the Baltic. In the inner part of the Ems estuary juveniles were predominantly present in muddy sediments high in the intertidal zone; in more sandy sediments at higher salinities juveniles and adults co-occured. Detailed information was obtained at a muddy and at a sandy station. Gametes were present in the coelomic fluid from November through March. In May new recruits were found in the sediment samples, reaching densities of over 10⁵ M⁻² at the muddy station, andc. 2000 m⁻² at the sandy station. During summer, densities decreased at the muddy station, coinciding with a density increase at the sandy station, suggesting migration of juveniles from a nursery to the adult habitat. Cage experiments showed that the decrease of juveniles at the muddy station could be attributed to migration and not to mortality due to predation. At the sandy habitat palps and anterior parts ofM. viridis made up 4–11% of the stomach content of juvenile plaice (Pleuronectes platessa). In juvenile flounder (Platichthys flesus) only in Aprilc. 10% of the stomach content consisted ofM. viridis. During 1983–1990 increasing densities ofM. viridis at the sandy habitat coincided with a reduced abundance ofNereis diversicolor, however, this inverse relationship was not found to be statistically significant. Density fluctuations ofM. viridis andCorophium volutator showed a significant positive relationship, the cause of which is not yet understood.     

Laboratory Studies of Parasite Transmission Aspects between Hydrobia spp. (Gastropoda) and Corophium volutator (Amphipoda)

Aspects of parasite transmission between Hydrobia spp. and Corophium volutator, first and second intermediate host of digenetic trematodes, were investigated under laboratory conditions. H. ventrosa is used as an intermediate host by several trematode species. Under laboratory conditions the most frequently observed emergence from H. ventrosa was of cercariae of Maritrema subdolum. The number of cercariae shed per day varied considerably. It was observed that 30 cercariae on average and up to 450 cercariae at maximum can emerge from a single H. ventrosa per day. Cercarial production continued until the death of the snails. The life-span of cercariae of the species M. subdolum decreased as the water temperature increased. It can be concluded that under natural conditions the cercariae, after emerging, have a maximum period of 1 day in which to seek out their second intermediate host C. volutator. Almost all specimens of C. volutator exposed to cercariae of the species M. subdolum died within the test period of 6 days. High average cercarial densities caused short life-spans (<50 h), while at lower densities longer survival times were possible. For cercariae of other Trematoda species, we were unable to find any equally clear evidence of a reduction in the survival rate of C. volutator within the test period. Mortality of C volutator, and other effects of infestation, as observed in our experiment, can be assumed to be a result of the penetration process of the cercariae but they also can be attributed to the mesocercariae.     

Effect of the snail Ilyanassa obsoleta (Say) on dynamics of the amphipod Corophium volutator (Pallas) on an intertidal mudflat

We investigated interactions between two dominant invertebrate species of intertidal soft-sediment environments of the northwest Atlantic, the mud snail Ilyanassa obsoleta and the burrowing amphipod Corophium volutator, on a mudflat of the upper Bay of Fundy, Canada. Distribution of I. obsoleta on the mudflat was highly patchy and negatively correlated with density of C. volutator. Manipulation of snail density in cages showed that I. obsoleta influences C. volutator; specifically, increasing density of snails reduced density, increased patchiness in distribution, decreased recapture rates and decreased immigration of C. volutator. Ilyanassa obsoleta seems to be affecting C. volutator through an influence on survival rate and emigration rate, although temporal variation in these effects was observed. Given that both I. obsoleta and C. volutator show a preference for tide pools, an important microhabitat on mudflats, snails might have a profound impact on C. volutator population dynamics.     

Experiments on epibenthic predation in the Wadden Sea

Field experiments were designed to evaluate the role of predators in the Wadden Sea, small predators like shore crabs, shrimps and gobies, and large ones like flatfish and birds. Exclosures, maintained in aCorophium volutator bed, an eelgrass bed, in a sandy and a muddy flat, protected the infauna from such epibenthic predation. The resulting changes in the macrofauna were recorded and compared with an unaffected control area. In sandy and muddy flats of the lower intertidal zone, cages (mesh size5 mm) altered abundance and composition of the infauna almost entirely. Nearly all species achieved higher population densities than in the control area, and in addition, the number of species increased as well. In the the scarcely populated mud flat a dense suspension-feeder assemblage emerged, associated with numerous tube-building polychaetes. The sand flat, normally dominated by deposit feeders, also became occupied by a dense suspension-feeder assemblage, mainly cockles. In contrast, narrowly meshed cages had only little effect in the beds of eelgrass and ofCorophium volutator. Both are positioned in the upper intertidal zone. Although a number of species still responded with significant increases in abundance, many remained indifferent or even tended to be less abundant within cages. Cages provided with a 20-mm mesh nylon net, excluding only birds, flatfish and the biggest crabs, increased significantly the survival of large-sized infaunal members. This was only apparent in the upper intertidal zone. It was concluded, that small sized epibenthic predators are the major determinants of the dynamic species abundance pattern of the lower intertidal flats. In the eelgrass bed, the meshwork of rootlets constitutes an important spatial refuge from these predators. The complex habitat structure causes a diversified faunal assemblage. TheCorophium bed in the uppermost intertidal zone is less accessable to predators like crabs, shrimp and gobies. The monotonous appearance of this faunal assemblage is assumed to be the outcome of competitive exclusion and of occasional harsh physical conditions.     

Behavioral Response of Corophium volutator to Shorebird Predation in the Upper Bay of Fundy,Canada

Predator avoidance is an important component of predator-prey relationships and can affect prey availability for foraging animals. Each summer, the burrow-dwelling amphipod Corophium volutator is heavily preyed upon by Semipalmated Sandpipers (Calidris pusilla) on mudflats in the upper Bay of Fundy, Canada. We conducted three complementary studies to determine if adult C. volutator exhibit predator avoidance behavior in the presence of sandpipers. In a field experiment, we monitored vertical distribution of C. volutator adults in bird exclosures and adjacent control plots before sandpipers arrived and during their stopover. We also made polymer resin casts of C. volutator burrows in the field throughout the summer. Finally, we simulated shorebird pecking in a lab experiment and observed C. volutator behavior in their burrows. C. volutator adults were generally distributed deeper in the sediment later in the summer (after sandpipers arrived). In August, this response was detectably stronger in areas exposed to bird predation than in bird exclosures. During peak predator abundance, many C. volutator adults were beyond the reach of feeding sandpipers (>1.5 cm deep). However, burrow depth did not change significantly throughout the summer. Detailed behavioral observations indicated that C. volutator spent more time at the bottom of their burrow when exposed to a simulated predator compared to controls. This observed redistribution suggests that C. volutator adults move deeper into their burrows as an anti-predator response to the presence of sandpipers. This work has implications for predators that feed on burrow-dwelling invertebrates in soft-sediment ecosystems, as density may not accurately estimate prey availability.     

Anthropocene invasion of an ecosystem engineer: resolving the history of Corophium volutator (Amphipoda: Corophiidae) in the North Atlantic

Resolving the natural histories of species is important for the interpretation of ecological patterns, as it provides evolutionary context for the interactions between organisms and their environment. Despite playing an integral role on the intertidal mudflats of the North Atlantic as an abundant food source for predators and as an ecosystem engineer that alters the soft sediment environment, no previous studies have provided empirical evidence to determine the biogeographical origin of the amphipod Corophium volutator. To resolve its status as introduced or indigenous in Europe and North America, we analyzed sequence data for two mitochondrial loci and two nuclear markers, aiming to determine whether the present range of C. volutator is the result of unresolved taxonomy, persistence in glacial refugia, natural trans‐Atlantic dispersal, or human‐mediated introduction. Our results demonstrate a reduced genetic diversity in North American populations that is a subsample of diversity in European populations, with coalescent analyses of mitochondrial and nuclear DNA supporting different models of multiple introductions from Europe to the Bay of Fundy and Gulf of Maine in North America. These results suggest that C. volutator was introduced to North America prior to the first surveys of local biota in the 20th Century, which has broad implications for interpretations of community and ecosystem interactions in the North Atlantic intertidal. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 115 , 288–304.     

Effects of estuarine infauna on sediment stability and particle sedimentation

Secretions produced by the two estuarine benthic invertebrates Corophium volutator and Nereis diversicolor stabilise sediments by increasing their shear strength, and reduce sedimentation of previously resuspended particles. The secretions consist of 1 to 2 m threads which bind the particles together. They are used by both species in the production of complex burrow systems in the top 15 cm of sediment. The burrow systems have been demonstrated by a new resin impregnation technique. These results have widespread implications for the stability and erosion of estuarine sediments.     

Mudflat biota since the 1930s: change beyond return?

Where, since the 1980s, patchy and variable green algal mats are prevailing, distinct belts of an amphipod (Corophium volutator) and seagrass (Zostera spp.) had dominated in the 1930s. The zonation between tide marks has been mapped in a sheltered sedimentary bay in the Wadden Sea near the island of Sylt (coastal eastern North Sea). Maps on vegetation from 1924 and on selected macrobenthos from 1932 and 1934 are compared with biannual surveys conducted from 1988 to 2006. Rising high water levels and eutrophication are suggested to be major causes of the observed long-term changes. In front of a saltmarsh, a sandy beach developed and partly displaced former cyanobacterial mats. Advancing sandiness may have inhibited C. volutator and facilitated lugworms, Arenicola marina, in the upper tidal zone. A variable occurrence of green algal mats arising in the 1980s affected infauna and seagrass by smothering the biota underneath. This dissolved a coherent belt of Zostera noltii. In the lower tidal zone, natural disturbances had lasting effects on the occurrence of mussels with attached fucoid algae. The spectrum of species became enriched by alien species (13% of macrobenthic taxa). A reversal to habitat structure and biotic zonation of the 1920–1930s does not seem possible. Aliens, in combination with climate change, are expected to further divert the ecological pattern to new configurations.     

Competition in foraging flocks of migrating semipalmated sandpipers

I examined the effect of competitor density on foraging success in staging semipalmated sandpipers (Calidris pusilla) foraging on a burrowing amphipod (Corophium volutator) in each of two study years. Little is known about the effect of competitor density when predation attempts disturb prey, causing a temporary decrease in food availability. Controlling for Corophium density and other potentially confounding factors such as temperature, pecking rate and capture rate increased linearly with sandpiper density. Success rate, the ratio of captures to pecks, was not influenced by sandpiper density. The effect of sandpiper density was similar in each of the two study years and was documented early and late in the low tide period. The increase in foraging rate is argued to be a response to increased competition for rapidly depleting prey at the temporal scale of exploitation by a flock. Potential fitness costs associated with higher foraging costs may include decreased ability to distinguish between the profitability of different prey and reduced vigilance against predators.     

Food selectivity versus prey availability: a study using the marine fish Pomatoschistus microps

Summary The food selection of the common goby, Pomatoschistus microps Krøyer, was studied in the field and in laboratory experiments on the Swedish west coast. The three most important prey organisms for P. microps in the study area were Corophium volutator, chironomid larvae and Nereis spp. Corophium was consumed more than any other prey, even when it was not the most abundant prey species in the bottom. One reason may be the higher activity of Corophium above the sediment surface, which may increase its visibility and consequently its vulnerability to visual predators. When P. microps was offered Corophium and chironomid larvae with similar exposure in laboratory experiments, it showed no preference for either of the prey items. It always took the closest mobile prey, regardless of species and size.     

Effects of Ilyanassa obsoleta (Say) on the abundance and vertical distribution of Corophium volutator (Pallas) on mudflats of the upper Bay of Fundy

Invertebrates in soft-bottom marine communities exhibit a range of responses to predators and competitors, including both emigration and avoidance. Corophium volutator (Pallas), a burrowing amphipod abundant in the upper Bay of Fundy, Canada, frequently interacts with the eastern mudsnail, Ilyanassa obsoleta (Say), as both species utilize similar habitats and food resources. Typically these interactions result in negative effects on C. volutator. There is evidence that Corophium species exhibit avoidance behaviour when interacting with other species, but whether such a response occurs in the presence of I. obsoleta is unknown. To investigate C. volutator response to I. obsoleta, total C. volutator abundance and vertical distribution within the sediment was examined in a field experiment involving I. obsoleta enclosures and exclosures at two mudflats in the upper Bay of Fundy. I. obsoleta densities were manipulated between June and August 2008 and their effects on C. volutator abundance and vertical distribution observed. Results varied between mudflats and C. volutator size classes. At Peck's Cove, total C. volutator abundance was largely unaffected by I. obsoleta. No size class exhibited a distribution shift in response to I. obsoleta, though both mid-sized individuals and adults occupied deeper layers of sediment late in the summer. Abundance of C. volutator in all size classes at Grande Anse was reduced in July in direct proportion to exposure to I. obsoleta, and adults occupied higher sediment layers in July when I. obsoleta were excluded, suggesting that I. obsoleta curtailed use of these upper layers. Juvenile abundance rebounded in August, possibly due to the mass dispersal of a newly produced juvenile cohort. Adults were reduced in all treatments in August, probably as a result of normal seasonal mortality. Differences in C. volutator responses between mudflats highlight the site-specific nature of such interactions.     

Recherches écologiques sur les faunes des plages arbitées de la région d'Arcachon

Resume 1. Le présent travail est consacré à l'étude écologique des plages abritées de la baie d'Arcachon (France). L'auteur tout d'abord, reconnaît et localise dans la région étudiée les quatres zones classiques qui partagent les estrans meubles abrités: slikke, haute slikke, schorre et zone parhalienne. Chacune d'entre elles est ensuite étudiée de manière plus détaillée, quant aux peuplements qui les occupent et aux structures édaphiques qui leur correspondent.2. Dans la slikke, l'auteur, reconnaît et décrit quatre faciès fondamentaux: les herbiers deZostera nana, les sables nus, type sables à arénicoles, les vases àNereis diversicolor et les vases à scrobiculaires. Chacun de ces faciès est analysé quant à sa teneur en eau, porosité, salinité, pH et oxygène dissous, répartition hypsométrique. En ce qui concerne la faune de ces slikkes sablo-vaseuses, l'auteur souligne l'importance de la macrofaune endogée, son caractère infralittoral, la faible mobilité de ses représentants.3. Dans la haute slikke, l'auteur reconnaît et décrit cinq faciès: les sables argileux àBledius spectabilis, les sables vaseux àCorophium volutator, les sables poreux àBledius arenarius, les aires àSpartina maritima et les microfalaises. De nouveau, sont étudiés successivement les caractéristiques abiotiques, puis les peuplements de chaque faciès; l'auteur souligne l'action réciproque des structures édaphiques sur la répartition des peuplements, mais aussi des peuplements sur les structures édaphiques. Par exempleCorophium volutator etBledius spectabilis édifient des galeries à parois stables, permanentes, qui ne peuvent être aménagées que dans certains types de sédiments; mais tandis que les galeries deB. spectabilis sont denses et étanches à la marée, celles deC. volutator sont plus dispersées et inondables; aussi la présence de l'insecte permet l'installation d'une faune d'accompagnement supralittorale, qui s'enterre au moment de la haute mer, celle de l'amphipode s'accompagne au contraire, d'une faune médiolittorale inondable.4. Enfin, dans le schorre et la zone parhalienne, l'auteur souligne l'absence de macrofaune endogée, la mobilité des peuplements vagiles épigés, leur balancement saisonnier lié au régime des marées. Il montre notamment qu'en été, le schorre abrite à la fois des espèces supralittorales marines ou halobies, et des espèces adlittorales haloxènes qui, venant de la zone parhalienne, sont attirées par la richesse trophique du Puccinellietum. En hiver, au contraire, la fréquence des immersions dans le schorre oblige la faune à refluer dans la zone parhalienne, où elle trouve son abri.5. Une classification des faciès étudiés en fonction de leur teneur en eau écologiquement utilisable par la macrofaune, un tableau de correspondance entre slikke, haute slikke, schorre et zone parhalienne d'une part, et étages intertidaux d'autre part, enfin une comparaison entre les peuplements des plages abritées et plages semi-abritées, complètent l'ensemble du travail.

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Ecological research on the faunas of the sheltered beaches of the region of Arcachon

In these ecological studies of the sheltered beaches of the bay of Arcachon (France), the author distinguishes and locates four traditional zones: slikke, upper slikke, schorre and parhalian zone. Animal communities and edaphic structures of each zone are treated in detail. In the slikke, four fundamental facies are distinguished:Zostera nana zones, bare sands (such asArenicola sands),Nereis diversicolor slime andScrobicularia slime. Each facies is analyzed with regard to its amount of water, porosity, salinity, pH and dissolved oxygen content, as well as its hypsometric distribution. With respect to the fauna of sand and slime slikkes, the importance of the endogeous macrofauna, its infralittoral characteristics and the low degree of mobility of its species are emphasized. In the upper slikke five facies are described:Bledius spectabilis clay sands,Corophium volutator slimy sands,Bledius arenarius porous sands,Spartina maritima areas and micro cliffs. Again, abiotic characteristics and animal communities of each facies are studied successively. The importance of edaphic structures for the distribution of animal communities, and vice versa, is stressed. For instance, the amphipodCorophium volutator and the coleopteranBledius spectabilis build tubes with firm and permanent inner surfaces only in certain types of sediment: yet, while theBledius spectabilis tubes are dense and watertight, those ofCorophium volutator are less tight and often flooded. Thus, the presence of the insect allows a supralittoral fauna to establish itself, whereas that of the amphipod is, in contrast, accompanied by a mediolittoral fauna resistant to flooding. In the schorre and parhalian zone, endogeous macrofauna elements are entirely absent; the vagrant epigeous species exhibit considerable degrees of mobility, migrating according to tidal rhythms. In summer, the schorre zone shelters both marine supralittoral or halobiotic species as well as adlittoral haloxene species which, coming from the parhalian zone, are attracted by the trophically rich Puccinellietum. In winter, however, frequent floodings of the schorre force the fauna to retreat to the parhalian zone for shelter. A classification is presented of the facies studied, based on the amount of water ecologically usable by their respective macrofaunas. The relationships between slikke, upper slikke, schorre and parhalian zone on the one hand, and the intertidal zones on the other are illustrated in the form of a synopsis. Finally, a comparison is made between animal communities of sheltered and of partly sheltered beaches.

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Ce travail constitue la 2ème partie d'une thèse de Doctorat ès Sciences naturelles soutenue à la Faculté des sciences de l'Université de Bordeaux (n^o d'ordre 197) et intitulée «Recherches écologiques sur les faunes des plages arbitées et des étangs saumatres de la region d'Arcachon». L'exemplaire dactylographié de la thèse in extenso a été déposé au C.N.R.S. et enregistré sous le n^o A.O. 1612.     

The influence of Corophium volutator abundance on resuspension

Two experiments were performed to test the hypothesis that Corophium volutator affects the turbidity of water in estuaries through active resuspension of sediment. One experiment was done in a flume system under different flow velocities, and one in aquaria. A diatom film developed at the sediment surface in both experiments before Corophium was added. This diatom film was supposed to have a stabilising effect on the sediment. In both experiments, the concentration of suspended solids in the water column increased with the density of Corophium individuals. No effect of flow velocity on suspended solids concentration was found. This indicates that, in our flume experiment, active resuspension by Corophium was more important than physical resuspension, at least at low flow velocity (<20 cm s^-1) and in the presence of a diatom film. The critical erosion threshold decreased with increasing Corophium density in the aquarium experiment, indicating that indirect effects of Corophium grazing may become more important at high levels of bottom shear stress. The implications of our findings for suspended solids concentration in estuarine systems are discussed.     

Feeding habits of young predatory fishes in marsh creeks situated along the salinity gradient of the Schelde estuary, Belgium and The Netherlands

Fish and macrobenthos were sampled in four different marshes along the salinity gradient of the Schelde estuary, Belgium/Netherlands, to investigate the importance of marsh creeks as foraging grounds for the dominant, larger fish species. The total density and biomass of all the main macrobenthic taxa (Corophium volutator, Nereis diversicolor, Oligochaeta, Macoma baltica and Heteromastus fliliformis) were measured. The feeding habits of the larger predatory fishes (Platichthys flesus, Dicentrarchus labrax) were investigated. Qualitative and quantitative stomach analyses included the calculation of different indices, showing the niche breadth (as diet diversity) and the niche overlap (as similarity between the predators diet) for this habitat. These analyses showed that the two most important benthic prey species for P. flesus were C. volutator and N. diversicolor. D. labrax preyed upon a wider range of species, including C. volutator, N. diversicolor, Crangon crangon, Carcinus maenas and Orchestia spp. The stomach diversity of D. labrax and P. flesus showed differences between the marshes although there was no consistent pattern in diet composition, reflecting the opportunistic nature of feeding by these large predators. The fullness indices of both flounder and sea bass did not differ significantly along the salinity gradient and the estimated minimum consumption by these predators did not indicate a top-down control of the macrobenthic community. The salt marsh creeks seem to provide excess food for the visiting fish species. The benthic prey was present in very high abundances, which may suggest that the typical nursery species such as C. crangon and C. maenas, and early juveniles of P. flesus, D. labrax and Pomatoschistus microps were not preyed upon significantly. This supports the hypothesis that salt marsh creeks provide good refuge areas for nursery species against predation by larger fish.