Flora and fauna of the sublittoral hard substrata in the Oosterschelde (The Netherlands) — interactions with the North Sea and the influence of a storm surge barrier

Since 1979 the flora and fauna of the artificial rocky shores of the Oosterschelde estuary have been investigated by diving biologists.Spatial and temporal patterns of species diversity and biomass were studied and several groups of species (communities) were distinguished. A west-east gradient in species composition was found. Species distribution proved to be mainly determined by current velocities, silt and plankton concentrations in the water, underwater irradiance, and the nature of the substratum. Some indications of migration between the basin and the North Sea are shown. Biomass of the hard substratum fauna was compared with that of the soft bottom fauna; the ratio appeared to be about 1 : 2, which is surprisingly high considering the small area available for hard substratum fauna.The key to the interactions with the North Sea is found in the water movements and the volume of water passing through the mouth of the basin during each tide. The abiotic and subsequent biotic changes in the basin caused by the building of the storm surge barrier are discussed, and used to illustrate the measure of dependence of the Oosterschelde on the North Sea.     

Changes in the fish fauna of the Oosterschelde estuary — a ten-year time series of fyke catches

Frequency of occurrence of fish species was monitored on a fortnightly basis in four fykes and a weir in the Oosterschelde estuary from 1979 through 1988. This was done in order to record changes in the fish fauna that may have occurred as a response to the construction of a storm-surge barrier in the mouth of the Oosterschelde (1984–1986) and the concomitant building of compartmentalization dams in the landward part. These compartmentalization dams reduced the freshwater inflow into the system. Principal component analysis using the annual averages in frequency of occurrence suggests a slight shift occurred in the fish community separating a cluster of years 1979–1984 from the cluster 1985–1988. Many of the changes in individual species could be attributed to fluctuations in yearclass strength or were part of changes occurring on a wider geographical scale. The only impact of the construction works seems to be the decrease in a number of anadromous fish. Fish traps seem to be useful as a monitoring tool for a number of species. The value of the data collected could be improved if catch size and length-frequency data are recorded.     

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).     

Oosterschelde estuary (S.W. Netherlands): a self-sustaining ecosystem?

The Oosterschelde estuary has a special position among the Dutch North Sea estuaries. With relatively unpolluted water and high transparency it combines high biotic diversity and high primary and secondary production. Before 1970 a major part of the water of the rivers Rhine and Meuse flowed into the Oosterschelde. The building of a permeable storm-surge barrier (1986) decreased the exchange between Oosterschelde and North Sea. The construction of two additional dams (1987) reduced the very limited fresh water discharge on the estuary even further. The total effect of these changes was a decrease of the nutrient levels in the water column.The estuary is used intensively for culture of mussels and fishery of cockles. Both zoobenthos groups use together up to 60% (in the western part) of the organic food available in the Oosterschelde estuary. The storm-surge barrier resulted in reduced current velocities and increased sedimentation, accompanied by higher transparency, hence accelerating primary production. This phenomenon is counteracted by the lowered nutrient concentrations throughout the year.In the seventies a preliminary carbon budget study indicated that a substantial import of organic carbon was required to sustain the ecosystem. Later studies did not confirm this hypothesis.In the present paper three different methods are presented to answer the question, if internal production suffices to sustain the foodweb. At first a steady state model is applied, secondly, calculations with actually measured process rates are carried out and, finally, a dynamic simulation model is used.The conclusion is drawn that, before the construction of the storm-surge barrier, primary production of organic matter by phytoplankton is much more important than import, although the latter can not be neglected as additional food source for mussels, cockles and zooplankton. The simulation model predicts that this conclusion will not have to be changed in the future. The Oosterschelde will remain largely a self-sustaining ecosystem.     

Changes in seasonal succession of phytoplankton induced by the storm-surge barrier in the Oosterschelde (S.W. Netherlands)

Seasonal succession of the phytoplankton was followed weeklyin the eastern and western parts of the Oosterschelde estuary,before (1982–1983) and during (1984–1986) the constructionof the storm-surge barrier. Construction of the barrier seriouslyinfluenced the hydrodynamics of the estuary. In the easternpart, decreased current velocities led to the emergence of avertical salinity gradient, to an increasing sedimentation ofsuspended matter and to a rise in transparency during a prolongedperiod of the year. The relationship between species compositionand biomass on the one hand and environmental variables on theother hand, was analysed in a Canonical Correspondence Analysis,for both compartments separately. The analysis revealed thatthe phytoplankton assemblage more and more obtained a summercharacter, extending its growth season both earlier and laterin the year. The transition of spring to summer phytoplanktonassemblages proceeded parallel to the main light-turbidity gradient.The factor light did not only explain the seasonal pattern,but also the long-term trend from pre-barrier to barrier period.Partial Detrended Correspondence Analysis after correction forthe mean seasonal pattern confirmed this trend significantly.     

Spatial and temporal patterns of intertidal macrobenthic populations in the Oosterschelde: are they influenced by the construction of the storm-surge barrier?

The construction of a storm-surge barrier in the mouth of the Oosterschelde caused important hydrodynamical and morphological changes that could influence the macrobenthic populations. This paper is one in a series of five all dealing with the effects of the storm-surge barrier on macrozoobenthos and analyses the spatial and temporal distribution of macrozoobenthos in the Oosterschelde and its relationship with some environmental parameters, based on two large scale sampling campaigns, one before and one after the completion of the barrier.The sediment of the sampling stations was fine, well sorted sand, with an average mud content of about 2.5 %. Only in the Krabbenkreek the sediment was coarser in 1989. The tidal elevation of the sampling sites decreased significantly in 1989.The density of macrozoobenthos was significantly lower, the biomass higher in 1989. The density was dominated by deposit feeders, the biomass by filter feeders. The difference in biomass between both years was mainly due to a substantial increase of the biomass of filter feeders in 1989. The number of species per station was significantly smaller in 1989 than in 1985. Between 1985 and 1989, frequency of occurrence decreased in 34 versus 13 which increased, density increased in 13 species and decreased in 34 species, biomass increased in 18 species and decreased in 29 species.Based on TWINSPAN several clusters of stations, each with a different faunal composition, were identified. These clusters did not form distinct zones on the tidal flats but were dispersed widely.The relationship between density and biomass of different trophic groups and the mud content of the sediment and the depth was analysed. This relationship sometimes differed clearly between years. The correlation coefficient of a multiple regression between density and biomass of individual species and environmental factors, although significant in most cases, was very low, indicating that only a small proportion of the species variability was explained. The relationship between benthos and environmental factors was further analysed by canonical correlation analysis and multivariate discriminant analysis that gave different results for the 1985 and 1989 data. This is probably due to the broad tolerance of the species to the range of the environmental variables found in our study area. From a TWINSPAN of the density data of 1985 and 1989 together we could conclude that, although the environmental parameters in a group of stations, showing a large faunal similarity in one year, did not change, the faunal composition did. This indicates that faunal changes are not necessarily linked to changes in the measured environmental parameters.In the discussion the different factors affecting macrobenthic populations are situated and it is suggested that the macrobenthic populations are probably more towards the nonequilibrium end of the continuum between nonequilibrium and equilibrium communities as defined by Wiens (1984).Although the impact of the construction of the barrier on the macrobenthic community seems at present to be rather small this does not mean that on the long-term there will be no effect.     

Tidal reduction and its effects on intertidal hard-substrate communities in the Oosterschelde estuary

Developments of intertidal hard-substrate communities in the Oosterschelde estuary were examined in perpendicular transects between high-water line and low-water line in the period 1982–1992. Prior to the beginning of the Oosterschelde estuary works a typology of communities was established and an overall survey of the estuary was carried out. The communities contain flora (algae) as well as fauna. Due to asphalting of dikes in 1986, much of the surface of several communities has been destroyed. The originally well developed communities with large species-richness have not returned. The small reduction in tidal amplitude due to the construction of the storm-surge barrier had a minor effect on the zonation of communities. In the upper part of the intertidal zone the boundaries of the communities moved 0.5–1.0 m downward in the transects along the dike-slopes. At an average inclination of 18° this means a vertical shift of about 15–30 cm. This reflects the reduction of the tidal amplitude: the high-water line shifted ca 22 cm downward. In a number of places sedimentation has caused a reduction in the number of smaller seaweed species in the lower eulittoral zone. At monitoring locations presence of the original communities is rather unchanged. Rare species like Pelvetia canaliculata, Actinia equina and Gelidium pusillum have been able to maintain quite successfully.     

Plankton size distributions and trophic relations before and after the construction of the storm-surge barrier in the Oosterschelde estuary

Phytoplankton and zooplankton biomass distributions were calculated on a carbon basis for the inland part of the Oosterschelde, in the period before (1983), during (1984, 1986) and after (1987, 1988) the construction of the storm- surge barrier. In all years studied, both phytoplankton and zooplankton distributions are very irregular, and little consistent patterns emerge. The data were used to test the model of Sheldon et al. (1977). The observed standing stock ratios of zooplankton to phytoplankton agree with the model predictions in 1983, and are slightly higher during the period 1984–1987. In 1988, the model predictions are very different from the observed values, because of important changes in the zooplankton species abundance occurring in this year.     

Artificial substrata in a shallow sublittoral habitat: do they adequately represent natural habitats or the local species pool?

Artificial substrata have been advocated as tools which have considerable potential for monitoring both natural and anthropogenic effects on invertebrate communities of shallow coastal environments. In this experiment, community structure was compared between two dominant natural algal habitats (kelp holdfasts and algal turf) and artificial substratum units (ASUs; nests of pan scourers) deployed in close contact with, and 20 cm above the substratum. Univariate and multivariate statistical analyses were applied to the data to determine the similarity of community structure between the four different habitats. In addition, recently developed measures of taxonomic distinctness were applied to the data from both sets of artificial substrata to determine if they provided a representative sample of the local epifaunal species pool and thus have the potential to be used as surrogate samples for this important faunal group. There were marked differences between community structure in each of the habitats. Both sets of artificial substrata were dominated by tubicolous polychaetes with abundances that were more than an order of magnitude greater than in the holdfast and turf samples. The fauna recruiting to the artificial substrata deployed above the substratum showed the lowest values in the univariate summaries of diversity and evenness and were unrepresentative of the local species pool. Artificial samples deployed in contact with the substratum showed greater diversity and evenness but were still mostly unrepresentative of the local species pool. The tendency for both sets of artificial substrata to under-sample amphipods and to be dominated by suspension-feeding polychaetes suggests that methods using these units may be relatively insensitive to the effects of anthropogenic impacts (e.g. sewage outfalls) where shifts in community structure including increased dominance of suspension-feeders and polychaetes and a reduced dominance of amphipods have been observed. Further studies, including the evaluation of temporal variation in community structure related to the time at which the ASUs are deployed and duration of deployment, are needed to test the wider utility of artificial substrata as tools for monitoring shallow, sublittoral, epifaunal communities.     

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.     

Changes in basin geomorphology after implementation of the Oosterschelde estuary project

The completion in 1986/87 of an open storm-surge barrier in the inlet and of secondary dams in the landward parts of the Oosterschelde tidal basin (SW Netherlands) has had and will continue to have a significant impact on geomorphological developments. An analysis of historic data, and of recent detailed bathymetric and morphodynamic process data, indicates that former trends have reversed. At present the Oosterschelde is a sedimentation basin with a degrading intertidal area and silting up of channels. The continuing reduction in intertidal area, the decreasing geomorphological gradients, the increasing fine sediment content of channel deposits, combined with a general reduction in hydrodynamics, imply significant ecological effects.     

Hydrodynamic characteristics of the Oosterschelde in recent decades

Hydrodynamic conditions in the Oosterschelde have changed as a result of the Delta Project. Engineering works were carried out between 1960 and 1970 in the northern branch of the Oosterschelde. The Grevelingen dam was built in 1964 and the Volkerak dam was completed in the spring of 1969. With the completion of these dams, the northern branch of the Oosterschelde was cut off from other estuaries, and river flow was then regulated by a sluice complex in the Volkerak dam. Between 1980 and 1987 a storm-surge barrier was constructed across the mouth of the Oosterschelde, and two compartmentalisation dams were built in the eastern part of the basin. The storm surge barrier decreased the effective cross-sectional area at the mouth from 80 000 m² to 17 900 m². The compartmentalisation dams reduce the surface area of the Oosterschelde basin from 452 km² to 351 km². The hydraulic impact of the works is demonstrated with data from field measurements and data from model calculations.     

Changes in the waterbird populations of the Oosterschelde (SW Netherlands) as a result of large-scale coastal engineering works

Between 1982 and 1987, the construction of a storm-surge barrier and two secondary dams in the eastern and northern parts of the Oosterschelde/Krammer-Volkerak area resulted in the loss of 33% of the 170 km² of intertidal area in the estuary. Consequences for non-breeding waterbirds were evaluated on the basis of monthly high-tide counts during five seasons before and three seasons after the construction period.In the entire Oosterschelde/Krammer-Volkerak area, numbers of wintering waders decreased but those of ducks increased. Peak numbers and total number of bird-days changed little, but the seasonal pattern shifted from a midwinter maximum to a peak in autumn.In the Oosterschelde (excluding the Krammer-Volkerak), where 17% of the tidal flats disappeared, species feeding mainly on open water remained stable or increased. Species dependent on intertidal areas for foraging (mainly waders and dabbling ducks) generally decreased. Total density of intertidal foragers decreased slightly. In most intertidal species, the Oosterschelde wintering population showed a stronger decrease, or smaller increase, than was shown during the same period by numbers in Britain and Ireland which were taken as an index of the total W-European winter populations. Changes varied considerably between species, and were correlated with their distribution within the estuary. Species concentrated in the eastern sector, where most habitat loss occurred, declined more than species with a more westerly distribution.Results indicate that intertidal foragers forced to move from the enclosed parts of the estuary were not generally able to settle into the remaining intertidal areas. Both dispersal to adjacent areas (mainly by dabbling ducks) and mortality during severe winter weather (in some wader species) may have contributed to the declines. Populations of intertidal foragers apparently were (and consequently still are) close to carrying capacity, and further changes in capacity, as foreseen from geomorphological changes still under way in the estuary, are likely to be reflected in bird populations.Numbers of waders moulting in the Oosterschelde in late summer declined strongly compared to numbers in other seasons. Increased disturbance due to recreational activities may have played a role during this time of the year.     

Policy planning in the Oosterschelde estuary

During the execution of the Delta project the awareness of the environmental values of estuaries was growing strongly. This awareness has led to a reconsideration of the original proposal to close off the Oosterschelde, and to an alternative view of the management of the estuarine areas and of the civil engineering structures. Now, the completed storm-surge barrier and two compartmentation dams, together with the reinforced dikes, guarantee a sufficient safety against flooding.After the decision to build a storm-surge barrier it was soon recognized that a coherent plan would be needed for the management and development of the Oosterschelde. A policy plan was accordingly produced in which the broad outlines of the policy were set out regarding the various potential uses of the estuary and the possible means of effecting them. Because of the periodic improvement and adjustment of the policy plan an environmental research programme was set up. Also, environmental research played a significant role in the decision-making processes for the completion of the barrier and the compartmentation dams as well as for the management of the storm-surge barrier.The coping-stone of the Delta Project marks the beginning of a new period in the water management of the Netherlands: the period of integrated water management.     

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.     

Changes in the sublittoral hard substrate communities in the Oosterschelde estuary (SW Netherlands), caused by changes in the environmental parameters

In order to assess the effects of the execution of the Delta Project, the sessile sublittoral communities on hard substrates in the Oosterschelde estuary and the environmental parameters were quantitatively investigated from 1985 till 1990. During the construction period of the barrier, three communities were sampled in the photic zone and four in the aphotic zone. The distribution of the communities in the photic zone seemed to be determined by the exposition to water movement and depth, while the communities in the aphotic zone were restricted to geographic areas, with differences in tidal current velocities: the mouth of the estuary, the Hammen, the central part and the Zijpe. Two years after the completion of the enclosure works, the community structure changed rapidly, caused by decreases of tidal current velocities, increases of the amounts of sedimentation, especially of fine sediments, and an increase of the transparency of the water. Changes within the associated vagile animals showed the same tendency as the sessile communities: under less exposed conditions the number of organisms remained the same or increased, while at some locations this increase was nullified by increasing amounts of sedimentation.     

Mussel culture in a changing environment: the effects of a coastal engineering project on mussel culture (Mytilus edulis L.) in the Oosterschelde estuary (SW Netherlands)

To evaluate the effects of a large scale coastal engineering project on the mussel (Mytilus edulis) bottom culture in the Oosterschelde estuary (S.W. Netherlands), mussel growth and production in the period 1980–1990 are studied in relation to food supply and the hydrodynamic conditions. Due to the construction of a storm-surge barrier and two additional dams, the risk that mussels are swept away by high current velocities decreased, resulting in an increase of the area in the Oosterschelde potentially suitable for mussel culture and in food availability now being more important as a limiting factor. For the Oosterschelde, a clear relation between mussel growth, stock sizes, and phytoplankton dynamics has been demonstrated. The meat yield of mussels landed in autumn — which is an index for growth rate — seems to be determined by the phytoplankton production in the preceding summer. In years with dense bivalve stocks, phytoplankton production and meat yields are relatively low. It is concluded that an increase of the mussel biomass cultured can result in a reduction of the primary production and, consequently, in a deterioration of the growing conditions for suspension-feeders in the estuary. This conclusion is supported by model calculations. An expansion of mussel culture in the new Oosterschelde is therefore dissuaded. Apart from primary production and stock sizes, food supply for mussels on culture lots appeared to be controlled by the horizontal advection of phytoplankton between and within the tidal channels. An observed decline in mussel landings from certain areas is attributed to the reduced mixing energy of the estuary in relation to the present distribution of the lots over the estuary. Production figures from the experimental lots, established in 1988 in the newly available areas, demonstrate that the yield of mussels can be enhanced by relaying culture lots towards the areas where the phytoplankton is produced. It is expected that by redistributing the culture lots, without expanding the biomass cultured, the carrying capacity of the Oosterschelde for mussel culture can be maintained.     

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.     

Effect of sedimentological and hydrodynamical changes in the intertidal areas of the Oosterschelde estuary (SW Netherlands) on distribution,density and biomass of five common macrobenthic species: Spio martinensis (Mesnil), Hydrobia ulvae (Pennant), Arenicola marina (L.), Scoloplos armiger (Muller) and Bathyporeia sp.

In order to evaluate the impact of the construction of the storm-surge barrier and secondary dams on macrobenthos of the tidal flats in the Oosterschelde (SW Netherlands), changes in distribution, density and biomass of five common species (Spio martinensis, Hydrobia ulvae, Arenicola marina, Scoloplos armiger and Bathyporeia sp) were analysed.Data on macrobenthos were collected from 1979 to 1989 on five different tidal flats. Changes in sediment texture and hydrodynamic factors during the construction and after the completion of the coastal engineering project were taken into account.Three severe winters in a row caused more disturbance in the population of the main predator of S. armiger than did the hydrodynamical changes. A temporary prolongation of the emersion time (in 1986 and 1987) caused a temporary decrease in juvenile A. marina. But afterwards they still occupy the same nursery grounds. Increased wave action on the edges of the flats probably created new niches for Bathyporeia sp. and Spio martinensis, replacing other benthic species.It is not yet clear what has caused the decline of H. ulvae in many places in the Oosterschelde estuary. Parasitic infestation is one of the possibilities.     

Long-term changes (1979–89) in the intertidal macrozoobenthos of the Oosterschelde estuary: are patterns in total density,biomass and diversity induced by the construction of the storm-surge barrier?

To evaluate the effects of the construction of a storm surge barrier in the Oosterschelde, long-term patterns (1979–89) in abundance and biomass of the intertidal macrozoobenthos were studied at 14 permanent stations. Additionally, data of a large-scale survey in late summer 1985 and 1989 were analysed. In this paper, patterns in general parameters are discussed.Late summer values of total biomass, total density, species richness, diversity and abundance- and biomass ratio show no overall significant trend during the study period. The changes in the hydrodynamics and the morphology of the Oosterschelde after the completion of the storm surge barrier do not seem to have influenced the normal patterns in benthic populations. The observed patterns are determined by the occurrence of severe versus mild winters, rather than by hydrodynamic changes caused by the construction of the barrier. Low biomasses, high densities (particularly of opportunistic species) and higher stress-values (abundance- and biomass ratio) in 1985(–87) indicate a temporal disturbance by severe winter weather. At the elevated COST-station 27, total biomass decreased sharply in 1985, due to a short-term increase in exposure time, caused by the manipulation of the storm surge barrier.