Macrofauna on flood delta shoals in the Wadden Sea with an underground association between the lugwormArenicola marina and the amphipodUrothoe poseidonis

Living conditions for macrofauna on flood delta shoals are determined by surf, strong currents and sediment mobility. Thus, a unique assemblage of invertebrate species colonize these far off-shore, low intertidal flats. We here describe the macrobenthic fauna of emerging shoals in the Wadden Sea between the islands of Römö and Sylt. Besides ubiquitous macroinvertebrates of the intertidal zone and species which attain their main distribution in the subtidal zone, the flood delta shoals are characterized by organisms adapted to live in these highly unstable sediments, like the polychaetesSpio martinensis, Streptosyllis websteri, Magelona mirabilis, Psammodrilus balanoglossoides, the pericarid crustaceansCumopsis goodsiri, Tanaissus lilljeborgi, Bathyporeia sarsi and a few others. Average abundance (1440 m^?2 of ind >1 mm) and biomass (12.9 g AFDW m^?2) were low compared to other intertidal habitats in the Wadden Sea. Biomass was dominated by largesized individuals of the lugwormArenicola marina. The U-shaped burrows of these polychaetes were inhabited by high numbers ofUrothoe poseidonis. Maximum densities of these amphipods occurred in the deepest parts of the burrows. Sampling at approximately montly intervals revealed no apparent seasonality ofU. poseidonis abundance. Together with smallCapitella capitata, these amphipods constitute a deep-dwelling component of the macrofauna associated with lugworms, which is separated from all other macrofauna living at the sediment surface. As a response to rising sea level and increasing tidal ranges, we expect the unstable sandy shoals, inhabited by numerousSpio martinensis andUrothoe poseidonis, to expand within the Wadden Sea at the cost of stable sandy flats with abundant macrofauna.     

Macrofauna distribution and bioturbation on tidal confluences of the dutch Wadden Sea

Macrofauna density and bioturbation intensity (measured with X-ray radiography), were studied in the Dutch Wadden Sea near Ameland and in Mok Bay, Texel, in September 1988. The bivalveMacoma balthica and the polychaeteHeteromastus filiformis were widely distributed in the areas studied. The cockleCerastoderma edule and the polychaeteScoloplos armiger did not occur in areas with high clay content, while the molluscsHydrobia ulvae andMya arenaria preferred fine sediments. The polychaeteNereis diversicolor preferred the higher parts of the intertidal. In the Ameland area disturbance of primary sediment structures was highest near the shore and near the middle of the tidal confluence where physical reworking was low. The sheltered Mok Bay sediments were completely reworked by deposit-feeders. Bioturbation intensity and deposit-feeder (bioturbator) density were, however, not highly correlated. A number of stations showed higher bioturbation towards the surface and this may be related to the high abundance of near surface dwelling macrofauna, due to eutrophication and organic enrichment of the sediments in recent years.     

Effects of the severe winter 1995/96 on the benthic macrofauna of the Wadden Sea and the coastal North Sea near the island of Sylt

The development of benthic macrofauna in the Wadden Sea and in the coastal North Sea after the severe winter of 1995/96 is compared with the preceding years with mild to moderate winters. In the intertidal of the Wadden Sea, ice-drift and low temperature caused the expected changes in species composition by increasing winter mortality in sensitive species, and by exceptionally high recruitment of some species during the succeeding summer. In the shallow subtidal (10–20 m depth), similar winter effects were observed. However, recovery of many subtidal populations was still incomplete until the summer of 1997. It is suggested that this was due to hydrographic conditions that carried many larvae or drifting juveniles into more distant offshore areas. This may have limited larval supply and may have delayed recovery at the onshore sites. Since in the eastern North Sea severe winters are accompanied by frequent easterly winds, it is not clear whether decreasing winter abundances in some species were due to increased mortality, or to a seaward dislocation of organisms. Received in revised form: 7 May 2001 Electronic Publication     

Major Changes in the Ecology of the Wadden Sea: Human Impacts, Ecosystem Engineering and Sediment Dynamics

Shallow soft-sediment systems are mostly dominated by species that, by strongly affecting sediment dynamics, modify their local environment. Such ecosystem engineering species can have either sediment-stabilizing or sediment-destabilizing effects on tidal flats. They interplay with abiotic forcing conditions (wind, tide, nutrient inputs) in driving the community structure and generating spatial heterogeneity, determining the composition of different communities of associated species, and thereby affecting the channelling of energy through different compartments in the food web. This suggests that, depending on local species composition, tidal flats may have conspicuously different geomorphology and biological functions under similar external conditions. Here we use a historical reconstruction of benthic production in the Wadden Sea to construct a framework for the relationships between human impacts, ecosystem engineering and sediment dynamics. We propose that increased sediment disturbances by human exploitation interfere with biological controls of sediment dynamics, and thereby have shifted the dominant compartments of both primary and secondary production in the Wadden Sea, transforming the intertidal from an internally regulated and spatially heterogeneous, to an externally regulated and spatially homogenous system. This framework contributes to the general understanding of the interaction between biological and environmental control of ecosystem functioning, and suggests a general framework for predicting effects of human impacts on soft-bottom ecosystems.     

Eutrophication and mussel culture in the western Dutch Wadden Sea: Impact on the benthic ecosystem; a hypothesis

Since 1950, two large-scale changes have taken place in the western Dutch Wadden Sea, namely the eutrophication of the area and the introduction of an extensive mussel culture. Although eutrophication in the fresh waters started already around 1950, nutrient concentrations in the western Wadden Sea remained fairly constant until about 1970, due to the retention of nutrients in Lake IJssel, the main source. From 1970–1980 concentrations increased strongly, and during the last years the situation has stabilized. Mussel culture was introduced in 1950 and expanded during the next decade to an area of 70 km², all situated in the sublittoral area. From 1960 the area of mussel culture remained about constant with fluctuating yields of between 35 and 120 millions of kg fresh weight. Due to a lack of data for the period until 1970 the impact of eutrophication and mussel culture on the ecosystem cannot be assessed. From 1970 onwards an increased biomass and production of the macrofauna in the intertidal zone has been observed, which is attributed to eutrophication. The hypothesis is postulated that the introduction of mussel culture between 1950 and 1960 has resulted in an increased food competition in the area, leading to a decreased stock of the macrofauna in the intertidal. Eutrophication from about 1970 onwards has improved the food conditions and as a result both the macrofauna in the intertidal and the mussel in the sublittoral area would have increased in biomass, allowing higher maximum yields of the mussel culture. The importance of monitoring programs is stressed to follow these trends in the near future and to check the above hypothesis in areas where it is decided to introduce or intensify mussel culture. Presented at the VI International Wadden Sea Symposium (Biologische Anstalt Helgoland, Wattenmeerstation Sylt, D-2282 List, FRG, 1–4 November 1988) Publication No. 30 of the project “Ecological Research of the North Sea and Wadden Sea” (EON)     

Historical changes and inventory of macroalgae from Königshafen Bay in the northern Wadden Sea

Knowledge on the distribution, abundance and species richness of intertidal macroalgae occurring on sandy and muddy flats of the German Wadden Sea is still incomplete. We summarize published and unpublished information available on the presence of macroalgae on the tidal flats of Königshafen Bay (island of Sylt, North Sea), one of the more extensively studied areas of the Wadden Sea. A total of 46 green algal species, 36 brown algal species and 26 red algal species has been recorded within the last 120 years on soft and hard substrata of Königshafen Bay (disregarding species found unattached or drifting). Several of these species were only temporarily resident on the tidal flats. Today, at least 35 green, 15 brown and 12 red algal species occur within or close to Königshafen Bay. Significant long-term changes in species abundances have occurred in all three major groups of algae: Since the late 1970s, dense green algal mats dominated byEnteromorpha flexuosa, E. radiata andE. prolifera have occurred regularly on the intertidal flats, whereas a general decrease of brown and red algae has been documented. Two red algal species,Gracilaria verrucosa and its epiphyteCallithamnion corymbosum, were conspicuous members of the macroflora until the middle of this century. Although still present in the 1980s, they have now disappeared completely. On the other hand, the brown algaSargassum muticum has begun to colonize mussel beds. The causal background of long-term changes in the macroalgal flora of Königshafen Bay is discussed. Owing to substantial nomenclatural changes during the last 120 years, a revised species list with authors’ names and synonyms is included.     

Bio-availability of phosphorus in sediments of the western Dutch Wadden Sea

The purpose of this study was to make a prognosis of the effects of extended purification of terrestrial waste water, reaching the Wadden Sea by the River Rhine and Lake IJssel, on the phosphate concentration in the western Wadden Sea.The quantities of different phosphorus fractions in intertidal and subtidal sediments of the Marsdiep tidal basin (western Dutch Wadden Sea) were measured. Different methods are applied to determine the amount of phosphorus that can be released from these sediments. The direct bioavailability is determined by inoculating sediment suspensions with a natural mixture of precultured micro-organisms from the sampling area. A second approach is the measurement of the phosphate release under different redox conditions. Sequential extraction of sediment samples with different solvents is also applied. Under the present conditions and compared to the nutrient loads from fresh water (Lake IJssel) and from the North Sea, the phosphorus stored in the sediments of the western Dutch Wadden Sea plays a minor role in the total supply to micro-algae and bacteria. The bulk of the biologically available phosphorus in the sediments originates from the metal-associated fraction. Releasable phosphate may contribute to the local annual primary production to an extent of ca 45 to ca 150 g C m^–2 a^–1. The total amount of phosphorus in the sediment (mainly calcite associated) is twice to 6 times the biologically available amount.     

Thermal tolerance ranges and climate variability: A comparison between bivalves from differing climates

The climate variability hypothesis proposes that in variable temperate climates poikilothermic animals have wide thermal tolerance windows, whereas in constant tropical climates they have small thermal tolerance windows. In this study we quantified and compared the upper and lower lethal thermal tolerance limits of numerous bivalve species from a tropical (Roebuck Bay, north western Australia) and a temperate (Wadden Sea, north western Europe) tidal flat. Species from tropical Roebuck Bay had higher upper and lower lethal thermal limits than species from the temperate Wadden Sea, and Wadden Sea species showed an ability to survive freezing temperatures. The increased freezing resistance of the Wadden Sea species resulted in thermal tolerance windows that were on average 7 °C greater than the Roebuck Bay species. Furthermore, at a local-scale, the upper lethal thermal limits of the Wadden Sea species were positively related to submersion time and thus to encountered temperature variation, but this was not the case for the Roebuck Bay species. A review of previous studies, at a global scale, showed that upper lethal thermal limits of tropical species are closer to maximum habitat temperatures than the upper lethal thermal limits of temperate species, suggesting that temperate species are better adapted to temperature variation. In this study, we show for the first time, at both local and global scales, that the lethal thermal limits of bivalves support the climate variability effect in the marine environment.     

Associations between macrobenthos and invasive cordgrass, Spartina anglica, in the Danish Wadden Sea

We conducted a survey to provide knowledge on mechanisms controlling spatial and temporal variability of macrobenthos in an intertidal Wadden Sea area partly covered by invasive Spartina anglica. Benthic macrofauna was collected seasonally at seven stations along a transect covering non-vegetated mudflat and vegetated marsh areas. Shannon diversity index was consistently higher for macrobenthos in the open mudflat compared to the marsh area. Infaunal species, like Arenicola marina, Tubificoides benedeni and Macoma balthica, were more abundant in the mudflat than the marsh, while the opposite was evident for epifaunal species, like Hydrobia ulvae and Littorina littorea. The infaunal crustacean, Corophium volutator, on the other hand, appeared particularly attracted to the mudflat–marsh boundary. The biomass of below-ground plant materials and macrodetritus was positively correlated with the total macrofaunal abundance, while the biomass of dead below-ground plant materials alone showed a significant negative correlation with infaunal abundance. Total macrofaunal abundance was highest during summer in the mudflat, while no seasonal pattern was evident in the marsh. The negative influence of S. anglica invasions on infaunal abundance may have serious implications for higher trophic levels, such as waterbirds, that forage in Wadden Sea areas and thus for the overall biodiversity and ecosystem functioning.     

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.     

Prediction of benthic community structure from environmental variables in a soft-sediment tidal basin (North Sea)

The relationship between benthos data and environmental data in 308 samples collected from the intertidal zone of the H?rnum tidal basin (German Wadden Sea) was analyzed. The environmental variables were current velocity, wave action, emersion time (all of which were obtained from a 2-year simulation with a numerical model) and four sediment grain-size parameters. A grouping of sample stations into five benthos clusters showed a large-scale (>1?km) zoning of benthic assemblages on the tidal flats. The zoning varied with the distance from the shore. Three sample applications were examined to test the predictability of the benthic community structure based on environmental variables. In each application, the dataset was spatially partitioned into a training set and a test set. Predictions of benthic community structure in the test sets were attempted using a multinomial logistic regression model. Applying hydrodynamic predictors, the model performed significantly better than it did when sediment predictors were applied. The accuracy of model predictions, given by Cohen’s kappa, varied between 0.14 and 0.49. The model results were consistent with independently attained evidence of the important role of physical factors in Wadden Sea tidal flat ecology.     

Effects ofFucus vesiculosus covering intertidal mussel beds in the Wadden Sea

The brown algaFucus vesiculosus formamytili (Nienburg) Nienhuis covered about 70% of mussel bed (Mytilus edulis) surface area in the lower intertidal zone of Königshafen, a sheltered sandy bay near the island of Sylt in the North Sea. Mean biomass in dense patches was 584 g ash-free dry weight m^?2 in summer. On experimental mussel beds, fucoid cover enhanced mud accumulation and decreased mussel density. The position of mussels underneath algal canopy was mainly endobenthic (87% of mussels with >1/3 of shell sunk into mud). In the absence of fucoids, mussels generated epibenthic garlands (81% of mussels with <1/3 of shell buried in mud). Mussel density underneath fucoid cover was 40 to 73% of mussel density without algae. On natural beds, barnacles (Balanidae), periwinkles (Littorina littorea) and crabs (particularly juveniles ofCarcinus maenas) were significantly less abundant in the presence of fucoids, presumably because most of the mussels were covered with sediment, whereas in the absence of fucoids, epibenthic mussel clumps provided substratum as well as interstitial hiding places. The endobenthic macrofauna showed little difference between covered and uncovered mussel beds. On the other hand, grazing herbivores — the flat periwinkleLittorina mariae, the isopodJaera albifrons and the amphipodsGammarus spp. — were more abundant at equivalent sites with fucoid cover. The patchy growth ofFucus vesiculosus on mussel beds in the intertidal Wadden Sea affects mussels and their epibionts negatively, but supports various herbivores and increases overall benthic diversity.     

Temporal variations in microbenthic metabolism and inorganic nitrogen fluxes in sandy and muddy sediments of a tidally dominated bay in the northern Wadden Sea

Factors controlling seasonal variations in benthic metabolism (O₂ flux) and dissolved inorganic nitrogen (DIN) fluxes were examined during a 12–14 month period at three intertidal Wadden Sea stations. Since the flux measurements were made as small-scale laboratory core incubations, the results are primarily related to the microbenthic community (microalgae, bacteria, micro-, meio- and small macrofauna) and cannot be considered representative of the total benthic community in the Wadden Sea. Furthermore, it has to be emphasized that light intensity during day-time simulations were constant and saturating at all times. Benthic primary production and oxygen uptake appeared to be temperature dependent with a ‘seasonal Q₁₀’ of 1.7–1.8 and 2.7–4.3, respectively. Inundation had no effect on oxygen fluxes as evidenced by similar sediment respiration with and without water cover. A stronger temperature dependence of primary production in muddy than in sandy sediment indicated that the overall control in the latter may be complex due to factors like macrofaunal grazing and nutrient availability. Benthic respiration may not be controlled by temperature alone, as sedimentary organic matter content correlated significantly with both temperature and benthic respiration. Annual gross primary production in high intertidal sandy sediment was 10 and 50% higher than in low intertidal sandy and muddy sediments, respectively. Since annual benthic community respiration was 2 times higher in muddy than sandy sediments, the annual net primary production was about 0 in the former and 17–19 mol C m^?2 yr^?1 in the latter. However, heterotrophic contribution by larger faunal components as well as removal of organic carbon by waves and tidal currents, which are not included here, may balance the budget at the sandy stations. There was no or only weak relationships between (light and dark) DIN exchange and factors like temperature, sedimentary organic content, and oxygen fluxes. Factors related to nutrient fluxes, such as denitrification and nutrient concentration in the overlying water, may have hampered any such relationships. In fact, DIN fluxes at all three stations appeared to be strongly controlled by DIN concentrations in the overlying water. On an annual basis, the sediment appeared to be a net sink for DIN.     

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.     

The colonization of two new brackish water habitats on the isle of Texel (the Netherlands)

Summary As a result of the reconstruction of a dike on the Dutch island of Texel two small areas of the Wadden Sea were embanked in 1977 and 1978. In 1981 both by now blocked brackish waters were investigated for macrofauna and macroflora. The vegetation on these former parts of the Wadden Sea was still in a colonizing state. Many small patches ofRuppia spp. were found, contrary to older brackish waters with more extendedRuppia beds. The striking dominance ofR.maritima agrees with its large seed production enabling to colonize quickly new areas. In one of the ponds, where the maximum salinity of 23 Cl^– exceeded far the tolerance limit of this species given in the literature (15 Cl^–),R.maritima was still more numerous thanR.cirrhosa, a species which is more tolerant to salinity fluctuations. Several years after isolation these areas are still inhabited by a few Wadden Sea macrofauna species (e.g. Carcinus maenas, Hydrobia ulvae andCapitella capitata) who managed to survive the drastic environmental changes. In addition, a large number of species characteristic for blocked brackish waters has already colonized the new ponds. One of the two, that has always completely been isolated from other inland water, was inhabited in 1981 by 13 brackish water species. In the future the two new ponds on Texel are expected to loose their pioneer characteristics. There are reasons to expect that most of the Wadden Sea macrofauna relics will soon become extinct. A more extendedRuppia vegetation will probably develop together with a relative increase ofRuppia cirrhosa.     

Diversity and vertical distribution of cultured and uncultured Deltaproteobacteria in an intertidal mud flat of the Wadden Sea

The diversity and distribution of Deltaproteobacteria in an intertidal mud flat of the German Wadden Sea was characterized by molecular biological techniques and cultivation. A 16S rRNA gene library generated with general primers (303 clones) suggested that sulfate-reducing bacteria (SRB) related to Desulfobulbaceae and Desulfosarcina were abundant. Fluorescence in situ hybridization (FISH) with probes targeting these groups was used to characterize their vertical distribution. The combination of FISH with catalysed reporter deposition (CARD-FISH) significantly enhanced the detection of selected subgroups of Deltaproteobacteria, particularly in deeper sediment layers. Up to 11% of all cells were assigned to SRB. Organisms related to Desulfosarcina and Desulfobulbaceae were the dominant SRB in the surface sediments. Two abundant subpopulations of Desulfosarcina-related bacteria were identified by FISH. The SRB community differed between the sampling site and a sandy intertidal flat chosen as a reference. Enrichments and MPN cultures inoculated with surface sediment were monitored by FISH. Nine strains of Deltaproteobacteria were isolated. Four strains were related to Desulfobulbaceae, such as Desulfobacterium catecholicum and Desulfocapsa spp. A subgroup including clone sequences and strains related to D. catecholicum could be detected in situ by a specific FISH probe. The first physiological experiments suggested specific functional roles for the isolates. Two strains utilized environmentally relevant compounds in coastal areas such as catechol and nitrate. One strain related to Desulfocapsa spp. disproportionated thiosulfate and might thus contribute to the sulfur isotope fractionation at the study site. A Fe(III)-reducing strain was obtained that affiliated with the Pelobacter-Desulphuromonas group. This group accounted for up to 6% of total cell numbers and even exceeded SRB numbers in upper sediment layers. These bacteria might substantially contribute to carbon mineralization via dissimilatory reduction of, e.g. Fe(III).     

High mortality of Pacific oysters in a cold winter in the North-Frisian Wadden Sea

Mortality of introduced Pacific oysters (Crassostrea gigas) was studied in the northern Wadden Sea in response to an ice winter. After a decade of mild winters, in January and February 2010, the first severe winter occurred since the Pacific oysters became dominant on former intertidal blue mussel (Mytilus edulis) beds in the North-Frisian Wadden Sea. After the ice winter, mortality of Pacific oysters on densely populated beds in the List tidal basin reached about 90%, indicating much higher losses in comparison to former mild winters. At lower densities between the islands of Amrum and Föhr, oysters were less or even not affected. Although Pacific oysters are assumed to be very tolerant to frost, the duration of cold water- and air temperatures accompanied by mechanical stress of the ice burden might have caused the high mortality in the winter 2009/2010 in formerly dense beds.     

The Dutch Wadden Sea: a changed ecosystem

Since 1600 the surface area of the Dutch Wadden Sea decreased considerably by successive reclamations of saltmarshes. In 1932 the Zuiderzee (3200 km²) was closed off from the Wadden Sea causing in the remaining part an increase in tidal range and current velocities. In 1969 the Lauwerszee (91 km²) was closed off and turned into a freshwater lake as well. Man's use of the Wadden Sea changed simultaneously. Dredging in harbours and shipping routes, and extraction of sand and shells became common practice. Extraction of sand increased manifold between 1960 and 1985. These activities did contribute to the turbidity of the Wadden Sea water. Discharge of nitrogen and phosphorus compounds into the western Wadden Sea increased also manifold since 1950, causing an increase in phytoplankton production, duration of phytoplankton blooms, and intertidal macrozoobenthic biomass. Loads of metals and organochlorine contaminants entering the Wadden Sea were hard to estimate. Fisheries changed drastically since the 1930's. Fishing for ‘Zuiderzee’ herring came to an end shortly after closing off the Zuiderzee. The anchovy fishery ceased in the 1960’, that for flounder in 1983. Undersized brown shrimps were fished until 1971. Selective shrimptrawls and sorting devices with flushing seawater were introduced to reduce mortality among young flatfish and shrimps. Oysters became extinct in the 1960's due to over-exploitation of the natural beds. Production of mussels increased more than ten times between 1950 and 1961 due to ‘culturing’, catches of cockles increased slowly between 1955 and 1984. Whelks were fished until 1970. The most important changes in the biotic system of the Wadden Sea, increased production of microalgae and intertidal macrozoobenthos, can be attributed to increased nutrient loads. Eutrophication provided ample food supply for mussels which are harvested mainly by man and eider duck, and may have caused also increased growth in juvenile plaice. Increased turbidity may have impaired life conditions for adult dab, and have presented also recovery of sublittoral eelgrass beds after their disappearance in the 1930's due to the ‘wasting disease’. Increased turbidity in the Wadden Sea is probably caused by closing off the Zuiderzee (1932), a significant increase of dredge spoil disposal near Hoek van Holland between 1970 and 1983, and a more than 10-fold increase of mussel culturing in the Wadden Sea since 1950. Stocks of several bird species breeding in the Wadden Sea area suffered great losses in the early 1960's due to a pesticide accident. Most of the breeding populations have recovered. PCB's caused a reproduction failure among harbour seals in the 1970's. Since 1980 official Dutch policy aims at multiple use of the Wadden Sea, with emphasis on protection and restoration of the natural environment. The 3rd Water Management Plan (1989) aims at a development of the Wadden Sea ecosystem towards the situation of ca. 1930, i.e. without undoing present sea dikes and reclaimed polders. Management of the Dutch Wadden Sea will therefore have to focus mainly on reduction of eutrophication, pollution and turbidity. Some management options are discussed.     

Variations in the sulfide regime and the distribution of macrofauna in an intertidal flat in the North Sea

Sulfide concentrations were measured in the inner and outer Königshafen (Sylt, Wadden Sea) from November 1990 to December 1991 to assess the fluctuations of sulfide levels in natural tidal habitats. Three different areas were compared: (1) muddy sediment (2) fine-medium sand, and (3) a mussel bed. Other abiotic factors such as Eh, pH, temperature, grain size and organic content were measured. After assessment of the macrofaunal distribution, an attempt was made to relate the distribution to the sulfide concentrations in the benthic habitat. Sulfide concentrations varied between sites throughout the year with considerable differences ranging from below 5 μM (limit of detection) to about 1 mM (Oct. 1991). The faunal composition (Table 2) at the different sites hardly varied; it was always dominated by annelids: The cirratulidTharyx marioni was the most abundant species in the upper layers of all sites, where it occurred at low sulfide concentrations (<50 μM).Heteromastus filiformis was commonly found in the deeper sediment layers of the muddy site where it was regularly exposed to sulfide levels around 75 μM and peak concentrations in autumn up to 1 mM.Capitella capitata, Tubificoides pseudogaster andTubificoides benedii were very common in the upper sediment layers where sulfide levels were about 150 μM in autumn. These species also occurred, however, in the deeper layers with higher sulfide concentrations. These results document not only the wide annual range of sulfide concentrations at different depths in a tidal flat, they also emphasize that under natural conditions tidal flat annelids are exposed to considerable concentrations of hydrogen sulfide.     

Long-term studies of macrozoobenthos in intertidal and shallow subtidal habitats near the island of Norderney (East Frisian coast,Germany)

Near the East Frisian island of Norderney two sites are investigated permanently in order to study long-term fluctuations of macrozoobenthos: one transect (since 1977) at the northern side in shallow subtidal waters and another one (since 1976) at the sheltered southern side in the intertidal area of the Wadden Sea. Since 1980 the investigations have been continued in the frame of COST-47, sedimentary intertidal programme (including the shallow subtidal habitats colonized by the Macoma balthica community).The results up to 1984, respectively to 1985, are presented and the changes of abundance of the dominant species are discussed. Discussed are also the influences of water temperatures and sediment disturbances caused by wave action. There is evidence that the intertidal variety of the Macoma balthica community shows a greater stability than the subtidal variety.