Zooplankton biomass in the Oosterschelde (SW Netherlands) before,during and after the construction of a storm-surge barrier

The hydrodynamic consequences of large coastal engineering (barrier-construction) works in the Oosterschelde were: prolonged residence times of the water, increased sinking of particulate material, and higher water transparencies. This strongly influenced the phytoplankton (Bakker et al., 1990; 1994) and phytoplankton biomass increased in the shallow Eastern compartment of the Oosterschelde (Bakker & Vink, 1994) while phytoplankton concentration of the seston rose.Zooplankton biomass, especially of copepods (Temora) and meroplankton (barnacle larvae) increased during the post-barrier period in the eastern compartment. It is hypothesized that this is caused by the improved feeding conditions and the increased retention times in this area.The barrier years 1985 and 1986 were characterized by low current velocities. In the Eastern compartment, this may have favoured the development of the rotifer Synchaeta (Bakker, 1994) and of the important copepod predator Pleurobrachia (Ctenophora).In the Western compartment, zooplankton developments in the post-barrier years were rather similar to those in the pre-barrier period. This led to the disappearance of the previously existing biomass gradients West-East (maxima in West). At present a trend in the opposite direction (maxima in East) is observed.     

Turbidity,nutrients and phytoplankton primary production in the Oosterschelde (The Netherlands) before,during and after a large-scale coastal engineering project (1980–1990)

Turbidity, nutrient concentrations and phytoplankton primary production were monitored in the Oosterschelde before, during and after the construction of a storm-surge barrier and two compartment dams.Flow velocities and suspended matter concentrations decreased severely, causing an increased transparency of the watercolumn. In the eastern and northern compartments, the previously pronounced seasonal variation disappeared.Reduction of the freshwater load and decreasing nutrient concentrations in the adjacent North Sea coastal waters resulted in lower nitrite + nitrate and silicate concentrations. Autumn phosphate concentrations remained at the same level as before the nutrient reduction. Silicate was a limiting nutrient during the pre-barrier period and nitrogen and silicate were limiting during the post-barrier period.Annual patterns in chlorophyll-a concentrations in the western and central compartments showed no obvious trend; in the eastern and northern compartments higher values were measured from 1985 onwards.Primary production during the period 1980–1990 varied between 176 and 550 g C m^–2 yr^–1. The annual primary production in the western compartment had decreased, while in the central and eastern compartments annual primary production did not change: the formerly existing gradient disappeared. In the northern compartment higher chlorophyll-a concentrations and high annual production suggest that the phytoplankton could benefit from the increased transparency while nutrient concentrations were still high enough to support phytoplankton growth.Changes in photosynthetic physiological parameters were observed which suggested shade adaptation. This is in contrast to improved light conditions and reduced nutrient availability. The apparent incoherence with light-shade adaptation theory may be explained by the species shift that occurred.As a result of the opposite effects of a more favourable light climate and a reduced nutrient availability, together with the resulting species shift, the annual primary production showed a large degree of homeostasis.     

The mobile epifauna of the soft bottoms in the subtidal Oosterschelde estuary: structure,function and impact of the storm-surge barrier

Data on the mobile epifauna of the Oosterschelde estuary, collected by beam trawl, were compiled from several studies. Multivariate statistical techniques brought out the fact that the Oosterschelde, when compared with neighbouring areas, has a characteristic epibenthic fauna. Diversity as measured by Hill's diversity numbers N through N ₊ , is higher for the Oosterschelde (N₁ = 4.5) than for the Voordelta (N₁ = 3.5) and the Westerschelde (N₁ = 2.2).Four epifaunal communities can be distinguished within the Oosterschelde, the two most seaward communities being the richest. Annual production is estimated at about 6 gADW m^–2 yr^–1, annual consumption is estimated at over 25 gADW m^–2 yr^–1. These results are highly dependent on the assumptions. Over 85% of the epibenthic production and consumption in the Oosterschelde is accounted for by only six species: starfish Asterias rubens, plaice Pleuronectes platessa, bib Trisopterus luscus, brown shrimp Crangon crangon, shore crab Carcinus maenas and dab Limanda limanda. In spite of its abundance, the sand goby Pomatoschistus minutus contributes little to the production.From the available data it is difficult to assess the impact of the construction of the storm-surge barrier and the compartmentalization dams on the epibenthic fauna. The increase in flatfish in the Hammen area is probably linked to the decrease in current velocities in that area. On the other hand the increase in the gadoids bib and whiting Merlangius merlangus is predominantly due to the stronger year classes in the post-barrier time period. Lower nutrient inputs through the Northern branch, in combination with the increase of the gadoids, may have caused the decline of the brown shrimp in the Oosterschelde. A decrease has also been observed in the sandeel Ammodytes tobianus and the hooknose Agonus cataphractus.     

A new trend in the development of the phytoplankton in the Oosterschelde (SW Netherlands) during and after the construction of a storm-surge barrier

During the pre-barrier period (1982–83), the Oosterschelde phytoplankton were a diatom-dominated community, comprising a species-rich assemblage throughout the year. Assemblages of spring, early summer and summer, developed in response to a gradually evolving turbidity-light gradient during the course of the year.During the barrier-construction period (1984–87), characterized by decreasing current velocities, increasing sedimentation of suspended matter, increasing water transparencies and unchanged nutrient conditions, the growth season for the phytoplankton started earlier and lasted longer. Some flagellate species responded by much higher biomass than before. The impact of short-term climatic factors during this period, notably severe winters, could be illustrated with examples of clear responses of some species (e.g. Biddulphia aurita).In the post-barrier years (1987–90) a changed light-nutrient-salinity regime (i.e. much light, limitation of nitrate, high salinity) was demonstrated and an extended summerseason developed, without the original gradual transitions. This was reflected in an a-seasonal trend of the phytoplankton assemblage, where summer species were already observed in spring and spring species decreased in abundance. In summer small flagellates increased and some weakly silicified diatom species made their appearance. In the eastern compartment no colony formation of Phaeocystis occurred in summer and this was thought to be due to nitrate limitation. Changes in abundance of some species (Phaeocystis, Ditylum brightwellii, Skeletonema costatum), occurring during the entire period of investigation (1982–90), could be explained using field observations compared with experimental evidence from the literature.The relationship between species composition and biomass on the one hand and environmental variables on the other hand, was analysed in a Canonical Correspondence Analysis, for both compartments separately.     

Zooplankton species composition in the Oosterschelde (SW Netherlands) before,during and after the construction of a storm-surge barrier

During the period of construction of a storm-surge barrier current velocities decreased strongly and the Eastern compartment of the basin obtained a lagoon-like character. The rotifer Synchaeta spp., already abundant in the neighbouring salt and brackish lakes, profited from this condition. Higher and less fluctuating salinities caused the estuarine character of this compartment to disappear in the post-barrier years. This was reflected in the obscured succession of some Acartia species: the estuarine A. tonsa dominated in pre-barrier- and barrier years during summer, while the marine A. clausi was abundant in the post-barrier Oosterschelde during that time. Also the changed food conditions (Bakker & Vink, 1993) may have played a role in this phenomenon.Further changes in species composition were hardly observed. The main changes were of a quantitative nature (Bakker & Van Rijswijk, 1993; Tackx et al., 1993). A list of the commonly occurring species is given.     

Consumption of benthic fauna by carnivorous birds in the Wadden Sea

Consumption by carnivorous birds was estimated for the Sylt-Rømø tidal inlet in the northern part of the Wadden Sea, as well as the subarea Königshafen, a small, tidal bay. The bird community of the Sylt-Rømø Wadden Sea was dominated by Dunlin (35% of all birds counted), Eider (9%), Oystercatcher (8%), Knot (8%), and Shelduck (7%). The community in the Königshafen was dominated by Eider (20%), Knot (17%), Bar-tailed Godwit (17%), Dunlin (13%), and Oystercatcher (8%). Annual consumption was estimated at 3.4 g AFDW · m^?2 · year^?1 for the entire Sylt-Rømø Wadden Sea and 19.2 g AFDW · m^?2 · year^?1 for the Königshafen. Restricting the calculations to the intertidal area resulted in a consumption of 8.7 g AFDW · m^?2 · year^?1 for the Sylt-Rømø Wadden Sea and 17.6 g AFDW · m^?2 · year^?1 for the Königshafen. In the two areas, consumption was dominated by the Eider with 37% and 60% of the total consumption, respectively. In comparison to the western parts of the Wadden Sea the seasonal pattern of consumption as well as species composition differed, most probably as an effect of different climatic conditions, whereas annual consumption on intertidal flats seems to be in the same order of magnitude. On average, 15–25% of the mean annual macrozoobenthic biomass seems to be taken by carnivorous birds in the Wadden Sea, which is in the same order of magnitude as in other northern temperate estuarine areas.     

Selective ingestion of phytoplankton by the bivalvesMytilus edulis L. andCerastoderma edule (L.)

Blue mussles (Mytilus edulis) and cockles (Cerastoderma edule) were collected in the Oosterschelde estuary (SW Netherlands) and exposed to diets consisting of the diatomPhaeodactylum tricornutum and silt. Algal concentrations were kept constant (20.10³ cells.ml^–1), and silt concentrations were varied between 3–80 mg.l^–1 (mussel experiments) and 20–120 mg.l^–1 (cockle experiments). Chlorophyll-a concentrations in the pseudofaeces were reduced compared to the diets, indicating selective ingestion of the algae. It was estimated that the mussels increase the ingestion of chlorophyll-a 2.0 times, and the cockles increase the chlorophyll-a ingestion 2.8 times, compared to the ingestion rate without selection. The selection coefficients were not affected by the SPM concentrations used. Phaeophytin-a concentrations in the faeces showed an increase as a consequence of the digestion of the ingested algae. Digestive efficiencies of chlorophyll-a varied between 36–92% The digestive efficiencies decreased with increasing SPM concentrations.Communication no. 541 of the Delta Institute for Hydrobiological Research.     

Microphytobenthos in the Oosterschelde estuary (The Netherlands), 1981–1990; consequences of a changed tidal regime

During the period 1981–1990 the functioning of microphytobenthos in the carbon cycle was studied in the Oosterschelde, a mesotidal, euhaline estuary (SW Netherlands), both before and after completion of a storm-surge barrier in the sea ward entrance of the estuary in 1986, which reduced the tidal range to 88% and current velocities to 70% of their former values on average.The annual biomass cycle has changed from small spring and autumn peaks into a much larger summer peak. The average biomass during summer has increased from 70 to 170 mg Chlorophyll-a m². The average annual biomass has increased from 115 to 195 mg Chlorophyll-a m². As a consequence the (calculated) primary production by microphytobenthos has increased also, from 150 to 242 gC m^–2 y^–1 (14 045 to 22 265 tonnes C y^–1), and its share in the total primary production of the Oosterschelde has increased from 16 to 30%. These increases in biomass and primary production are mainly ascribed to a decrease in the dynamic forces (water current velocities) over the intertidal flats in most parts of the basin. Increased water transparency in parts of the estuary and increased import of inorganic carbon from the water column towards shoals may have contributed as well.The rate of reworking of the top layer of the soil (0–10 cm) has not changed significantly, as the decrease in Chlorophyll-a biomass with depth has hardly changed.     

Evaluation of the sample size used for the rapid bioassessment of rivers using macroinvertebrates

Experiments were designed to investigate selective predation by medium (40–55 mm carapace width: CW) and large (55–70 mm CW) Carcinus maenas when feeding on four bivalves of contrasting shell morphology. Size-selection was examined by presenting individual crabs with a wide size range of Mytilus edulis, Ostrea edulis, Crassostrea gigas and Cerastoderma edule. Medium-sized crabs preferred mussels 5–15 mm shell length (maximum shell dimension: SL) and cockles 5–10 mm SL, whereas large crabs preferred mussels 15–25 mm and cockles 10–20 mm SL. Crabs generally showed no preference for any particular size of either oyster species. Species-selection was examined by presenting individual crabs with paired combinations of the four bivalves in various proportions. When offered mussels and oysters simultaneously, both size categories of crabs consistently selected mussels, and food choice was independent of prey relative abundance. By contrast, C. maenas selected mussels and cockles as expected by the frequency in which each size category of crab encountered the preferred size ranges of prey. Crab preference clearly paralleled the rank order of prey profitability, which in turn was mainly determined by prey biomass, suggesting that active selection takes place at some point of the predation cycle. Experiments with epoxy resin models showed that initial reluctance of crabs to attack oysters was not associated with the ultimate energy reward. Moreover, they suggest that foraging decisions are partly based on evaluations of overall prey shape and volume, and that the minimum dimension of the shell constitutes an important feature which crabs recognise and associate with prey value.     

Stable isotope evidence of benthic microalgae-based growth and secondary production in the suspension feeder Cerastoderma edule (Mollusca,Bivalvia) in the Marennes-Oléron Bay

The contribution of natural food sources to the growth and secondary production of the suspension feeding bivalve Cerastoderma edule (L.) was estimated under field conditions in the Marennes-Oléron Bay (Atlantic coast, France). Monthly estimates of abundance, biomass and cockle growth were combined with seasonal analyses of ¹³C and ¹⁵N ratios of juvenile and adult cockles, together with their potential food sources [i.e. suspended particulate organic matter (POM), microphytobenthos, macroalgae and seagrass] sampled at mid-tide level in a muddy sandflat. Adult cockles grew mainly in spring, whereas juveniles grew in summer and autumn, following spat recruitment in early summer. Total annual production and elimination of cockles were estimated to be 32.5 and 34.7 g AFDW m^–2 yr^–1. Relative contributions of each year class to production were ca 40, 41, 11 and 6% for 0-group, 1-, 3- and 4-yr-old cockles in 1995, respectively. Quantitative assessment of proportions of food sources to the annual secondary production of cockles was obtained by using a simple carbon isotope-mixing model with microphytobenthos (¹³C = –16.0±0.6) and POM (¹³C = –22.2±1.1) as end-members. On average, more than 70% of the total annual cockle production originated from microphytobenthos, with a much higher contribution for the 0-group (88%) than for adult cockles (60%). The between-age difference was induced mainly by changes in the availability of food resources (benthic versus planktonic) during the non-synchronous growing seasons of juvenile and adult cockles.     

The importance of avian-contributed nitrogen (N) and phosphorus (P) to Lake Grand-Lieu,France

The largest natural lake in France, Grand-Lieu, has suffered eutrophication. The objective of the study was to estimate the annual input of nutrients (N, P) resulting from avian excrement, deposited by birds feeding out of the lake and returning to its waters for breeding or roosting, as compared to the input by the rivers that enter in the lake. Two years are compared: 1981–82 and 1990–91. About 1600–2000 breeding herons and cormorants, 20 000–33 000 wintering ducks, gulls and cormorants and 1–2.4 million starlings deposited about 5800 kg total N in 1981–82 and 7640 kg in 1990–91. Respectively, 2000 and 2530 kg total P were deposited over the same time periods. These represent 0.7% and 0.4% of the total N input of the lake and 2.4 and 6.6% of the total P input in 1981–82 and 1990–91. Starlings account for 74% of the N and mallards most of the rest. P input by starlings (36% in 1981–82, 41% in 1990–91), and by mallards and herons (35% and 27% in 1981–82 and 22% and 24% in 1990–91 respectively) plays an appreciable role among birds. During the plant growing period (April–September), the contribution by birds can increase to 37% of total P input of the lake. Piscivorous bird colonies concentrate Phosphorus 42 times more within the colony than outside the colony. Overall, the role birds play in total N and P input is relatively small due to very high inputs from human sewage and agriculture run off. The monthly mean concentration of the water of the two rivers reaches currently 10 mg l^–1 of N (to 23 mg during peak floods) and 394 mg m^–3 of P (to 468 mg during peak floods). Earlier, for example in the 1960's, water in Brittany only contained 0.1 to 1.1 mg 1^–1 of N and 1 to 5 mg m^–3 of P during the maximum flow period. At this time, birds could probably have represented annually up to 37% of the N input and up to 95% of the P input to the lake.     

The response of salt marsh vegetation to tidal reduction caused by the Oosterschelde storm-surge barrier

In 1986 a sluice gate barrier was completed in the mouth of the Oosterschelde estuary. The barrier has been partially or completely closed during 1986 and the first months of 1987. Consequently the high tides were reduced to such a level that the salt marshes were scarcely flooded. Since April 1987 the barrier has been closed on average during two out of 706 high tides a year. Although the barrier allows tidal exchange the tidal flow has been restricted as a result of the reduced width of the mouth of the estuary. This restriction of the tidal flow caused a 26 cm decrease of mean high water in the eastern part of the estuary. As a result the inundation frequencies of the salt marshes decreased.The response of salt-marsh vegetation to this tidal reduction was analyzed using annual records (1982 till 1990) of species composition in 57 permanent plots in two marshes at the southern shore of the estuary. Analysis of the response of individual species to marsh elevation in the pre and the post-barrier situation revealed that most species moved down the marsh elevation gradient. The first axis of an ordination (DCA-1) was significantly negatively related to inundation frequency. Between 1984 and 1990 all plots were displaced towards a significantly higher ordination score, indicating a trend towards a species composition from higher up the marsh. The position of most plots along DCA-1 remained stable until 1985 and started to increase in 1986 or 1987. The vegetation in plots dominated either by Halimione portulacoides or by Spartina anglica, started to change in 1985. This premature change was attributed to frost damage in January 1985. The initially high rate of change along DCA-1 decreased in 1989 and 1990. This would suggest that the vegetation re-equilibrated with the newly established tidal conditions. Further analysis revealed no significant difference in the relation between inundation frequency and sample score along DCA-1 between 1984 and 1990. This corroborates the view that species composition had re-equilibrated with the tidal conditions. Along DCA-2 the samples were displaced towards a significantly higher score as well. The change was attributed to an increase of the perennial Halimione and the annual Suaeda maritima. The annual Atriplex hastata displayed an increase from 1986 till 1988, but strongly declined in 1990. The transient response of this annual was described by DCA-3.During the pre-barrier phase several attempts had been made to predict the vegetation response to tidal reduction. We developed a multivariate model (CCA) with inundation frequency as a constraining factor to describe the relation between inundation frequency and species composition in the pre-barrier phase in 1984. Next we used this model to predict the score of the samples along CCA-1 in the post-barrier phase from the inundation frequencies in 1990. The actual ordination score in 1990 was calculated from the observed species composition. The predicted ordination scores did not differ significantly from the observed ones. In retrospect it is concluded that the response of the vegetation to the reduction of the tides, as far as the fraction of species composition which is related to inundation frequency is concerned, could have been predicted by our model.

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4. NOTES ON THE BIRDS OF THE VAN RYNEVELD'S PASS IRRIGATION LAKE,GRAAFF-REINET

McLachlan, A., Woodridge, T., Schramm, M. &; Kühn, M. 1980. Seasonal abundance, biomass and feeding of shore birds on sandy beaches in the Eastern Cape, South Africa. Ostrich 51:44-52.

The numbers of birds on three Eastern Cape sandy beaches were monitored monthly over one year during 1977/78. Dominant among 17 species were the Southern Blackbacked Gull Larus dominicanus, Black Oystercatcher Haematopus moquini, Whitefronted Sandplover Charadrius marginatus and Sanderling Calidris alba. These four species made up 95% of bird numbers and their respective biomass values were 78 652, 7 695, 578 and 3 116 kJ/km shoreline. Total food consumption for these species, calculated on the basis of estimated standard metabolic rates, was 4 159 394, 521 149, 76 768 and 363 053 kJ/km respectively. Food composition and feeding behaviour are also discussed and the total quantity of intertidal macrobenthic animals consumed by birds estimated.     

Zooplankton-trophic state relationships in some north and central Florida lakes

In a survey of eight lake systems located in north-central Florida, total zooplankton abundance showed a strong positive correlation (r²=0.87, a=0.01) with trophic state. Zooplankton abundance averaged 1.0 × 10⁵ organisms · m^–2 in oligotrophic systems and up to 8.2 × 10⁵ organisms · m^–2 in the eutrophic systems. Seasonal variations in total abundance were greatest in the eutrophic lakes where rotifers dominated and periodically produced sharp population peaks (approaching 2.0 × 10⁶· m^–2). In contrast, the more oligotrophic systems had relatively stable levels of total abundance and were dominated by copepods. Diversities of the major taxa in the lakes were variable with one to three species of copepods, zero to four species of cladocera, and two to seven species of rotifers dominant at any one time. Planktonic cladoceran communities were often composed of only one or two species. Low cladocera diversity in these subtropical systems was suggestive of increased predation pressure on this group of crustaceans. A comparison of the total crustacean abundance in the Florida systems to those of some of the Great Lakes indicated that lower standing crops of crustacean zooplankton in the Florida lakes may be a response to both predation and temperature.Contribution Number 043, Marine Science Programs Laboratory, Dauphin Island, Alabama, U.S.A.Contribution Number 043, Marine Science Programs Laboratory, Dauphin Island, Alabama, U.S.A.     

A preliminary study on the effects of oyster culturing structures on birds in a sheltered Irish estuary

This study investigated the species composition, numbers and behaviour of birds in an intertidal oyster culture area in Cork Harbour . These data were compared to a nearby area free of aquaculture within the same estuary in March 1999. Species which occurred in the aquaculture free area were also observed in the trestle-area. The most abundant species were oystercatcher Haematopus ostralegus, redshank Tringa totanus, dunlin Calidris alpina, curlew Numenius arquata, black-headed gull Larus ridibundus and common gull Larus canus. Oystercatcher, curlew, black-headed gull and common gull occurred in significantly lower numbers in the trestle area, while for redshank and dunlin the differences were not significant. The percentage of birds feeding did not differ between the two areas. Oystercatcher, redshank, dunlin and curlew mostly fed in both areas. In contrast, black-headed gull and common gull generally did not feed, but surveyed the area. Whether the trestles were covered by oyster bags or not did not have any effect on the number of birds except for the dunlin. Dunlin were significantly more frequent beneath the trestles with bags compared with those without bags. In general, the percentage of birds feeding did not differ between areas. Interspecies differences occurred with regard to the position occupied by birds at the trestles. Oystercatcher, redshank and curlew spent more time underneath the trestles. Dunlin, black-headed gulls and common gulls did not differ in numbers underneath or on top of the trestles. These preliminary observations at a single time period give some insight as to the potential interactions between shellfish aquaculture and intertidal birds.     

Taxonomic status of the extinct Canary Islands Oystercatcher Haematopus meadewaldoi

Mitochondrial genes were sequenced from four specimens of the extinct Canary Islands Oystercatcher Haematopus meadewaldoi and compared with African Oystercatcher Haematopus moquini, Eurasian Oystercatcher Haematopus ostralegus and an old unidentified extralimital ‘black’ oystercatcher specimen from The Gambia. At these loci, H. meadewaldoi was approximately 99.65% identical to multiple Eurasian Oystercatcher samples and in phylogenetic trees fell within the range of genetic variation observed in that species. The mystery Gambian bird was resolved as an extralimital H. moquini. We conclude that H. meadewaldoi was most likely a recently diverged melanistic morph or subspecies of H. ostralegus, although further genomic studies will be required to determine whether there has been a period of isolation followed by introgression.     

Assessment of Grey Heron predation on fish communities: the case of the largest European colony

The fish predation rate by Grey Heron Ardea cinerea was studied during two breeding seasons (1987–88) in the largest European colony at the Lake of Grand-Lieu (Loire-Atlantique, France). The herons' diet was compared to the available fish population of its main feeding area, the marsh of Bourgneuf (16000 ha) which is composed of former salt pans and meadows drained by a dense network of shallow ditches. This study is the first attempt to assess the predation exerted by an ardeid colony on a fish community over such an extensive natural environment. It also provides the first data about the abundance and the structure of fish communities in shallow coastal dyked marshes. For this purpose, two different sampling methods were used according to the water's salinity. In fresh waters, electrofishing was used as the removal method, and density estimates were calculated with Carle & Strub estimator (1978). Fish were caught in randomly selected stations (sections of ditches enclosed by two 5 mm mesh nets). In brackish waters, pools and ditches were drained. The distribution of the herons at the feeding areas was determined by direct observations, by counting flights from the colony, and by radio-tracking. The diet was investigated by observing adult herons on the foraging areas, and by analyzing the prey regurgitations of the young at the nests. The global food consumption was assessed from Marion (1988), according to the birds' activity determined during 5 years of radio-tracking. Altogether, at least 39 species of fish were available in the herons' feeding area (during the reference period, 87–88) and the mean fish biomass was 270 kg per ha of open water, or 30 kg per ha of marsh (open water = 11.2% of the marsh area). The fish community was dominated by eel Anguilla anguilla (145 kg ha^–1, 50,8% of the total biomass), and catfish Ictalurus melas (40 kg ha^–1, 14%). Except for small and inaccessible species (living in the deepest parts of the marsh), heron diet was very similar to fish species composition of the community occurring in the marsh. The catfish was the species captured most frequently by the heron (45% of the mass), the eel was second with 28% of the mass. The catfish was probably over represented in the diet considering that they are caught in catfish-dumps created by professional fishermen at Grand-Lieu lake, in order to reduce the density of this undesirable species. Inversely, small species such as Gasterosteus aculeatus were not found in the diet whereas they are very numerous in the marsh. On average herons of Grand-Lieu colony catch 1.92 kg of fish per ha of marsh (6% of the fish standing crops in the marsh) during the breeding season, the main predation period.     

Biomass and standing stock on sublittoral hard substrates in the Oosterschelde estuary (SW Netherlands)

From 1979 to 1991 the species composition of communities living on hard substrata (hardsub) in the Oosterschelde has been studied — in both the littoral and sublittoral zones. From 1984 onwards, biomass was also measured. This paper deals mainly with the distribution and the development of biomass on sublittoral hardsub in the Oosterschelde. Analysis has shown that the most important abiotic factors regulating the flora and fauna are: quantity and nature of the substrate; sedimentation; exposure to water movement (mainly currents); and light. The construction of the storm-surge barrier has influenced those factors. The main consequences for the flora and fauna on sublittoral hard substrata have been through the increased amount of available hard substratum by about 10% until 1984 and a further 20% from 1984 to 1987, the main barrier construction period). Within the same period (until 1987) the biomass per square metre also increased. This caused a net increase of hardsub biomass — in the sublittoral — of about 35%.After the barrier was completed sedimentation increased; in some parts of the basin hardsub organisms were covered by sediment and have not recovered; the total quantity of available hard substratum decreased by an amount yet to be established. For the purpose of this paper it is tentatively estimated at 20%, but the process is still going on.Tidal current velocities are smaller in the post-barrier situation, which caused a shift from more passive suspension feeders to more actively filtering species. The relative importance of suspension feeders on hard substrata has decreased by about 20% after the building of the storm-surge barrier. In 1990 and 1991 it increased again.Overall water transparency increased, but the lower limit of macroalgal growth has not gone deeper, as nearshore turbulence and turbidity did not change significantly.Effects on hardsub were small in the beginning. During the construction period (1985–1987) no clear effects were registered. After the completion of the barrier total species diversity increased at first, followed by a decrease from the second half of 1988 onwards. Biomass increased rather sharply, at first, but decreased very sharply in 1989. In 1990 a recovery in biomass became apparent. Developments in biomass and species composition differed per sampling location. An attempt is made to explain some of those developments, in relation to the abiotic changes brought about by the storm-surge barrier. This appeared difficult, because climatic influences obscured the effects of the barrier. The most explicit of those masking effects was brought about by a temporary, huge increase of the brittlestar (Ophiothrix fragilis). This animal covered the substratum in relatively thick layers (up to 5 cm) and more or less suffocated the other fauna. It was therefore difficult to quantify the effect of increased sedimentation on the fauna. The increase of Ophiothrix is probably not caused by the storm-surge barrier, but by a succession of several mild winters.It is clear that a new equilibrium in the basin is still to be reached. Total effects in terms of species richness and of biomass will continue to be monitored, and the results used to advise the water authorities as to management and nature friendly dike building methods.     

The effect of size and temperature on the predation of cockles Cerastoderma edule (L.) by the shore crab Carcinus maenas (L.)

Laboratory studies demonstrated that shore crabs Carcinus maenas (L.) can consume <40 cockles Cerastoderma (= Cardium) edule (L.) per individual · day⁻¹. Various predation techniques used by the crabs are reported. The time required to open and consume individual cockles increased exponentially with prey size. Small (<l5-mm shell height), easily broken cockles appeared to be the most profitable in terms of energy acquisition per unit of handling time, the optimal size of prey increasing with predator size. With unlimited prey available, however, crabs selected prey of mean size smaller than these predicted optima, and much below the maximum size they were capable of opening. Feeding rates, both in terms of cockles ingested or energy intake per day, rose steeply with increasing temperature, but the size range of prey consumed remained unchanged. These data strongly suggest that Carcinus maenas is a potentially important predator of small cockles, particularly during the wanner summer months.     

Overwintering waterfowl on Swiss lakes: how are abundance and species richness influenced by trophic status and lake morphology?

During the period 1980–1990 long-term physical, chemical and ecological studies were carried out, to study the changes induced by the building of a storm-surge barrier in the mouth of the Oosterschelde estuary and two large auxiliary compartment dams in the rear ends of the estuary. The storm-surge barrier was constructed in the mouth of Oosterschelde estuary (SW Netherlands) during the period 1979–1986. The barrier allows the tides to enter the estuary freely, and, on the other hand, the barrier guarantees safety for the human population and their properties when a stormflood threatens the area.Oosterschelde estuary is isolated from the river input, the rear ends of the ecosystem were separated from the estuary by sea-walls and the strongly decreased tidal exchange with the North Sea induced sheltered circumstances. The Oosterschelde changed from a turbid estuary into a tidal bay, and yet primary production responses appear to be robust and resilient, and the biological communities showed only quantitative shifts from the dominance of specific species assemblages to other assemblages. In many cases predicted changes in the structure of the biological communities could not be verified owing to the large natural variability mainly caused by physical factors (e.g. temperature).