The Ecological State of the Waddensea. An Assessment

Since the present quality state of the Wadden Sea is judged more discordantly than that of the North Sea, it is examined whether the Wadden Sea is a separate ecosystem. The nutrient load into the Wadden Sea and the trend of the load are assessed. Biological indicators of the ecological state of the Wadden Sea are examined with the result that there are signs of a bad as well as of a good state. This contradiction is assigned to the fact that quality standards are absent. Self-protection mechanisms of the Wadden Sea are discussed with respect to their responsibility for the relatively good state. A qualitative model is proposed to explain the long-term behaviour of the ecosystem.     

Long-Term Changes in Nutrients and Mussel Stocks Are Related to Numbers of Breeding Eiders Somateria mollissima at a Large Baltic Colony

Background

The Baltic/Wadden Sea eider Somateria mollissima flyway population is decreasing, and this trend is also reflected in the large eider colony at Christiansø situated in the Baltic Sea. This colony showed a 15-fold increase from 1925 until the mid-1990''s, followed by a rapid decline in recent years, although the causes of this trend remain unknown. Most birds from the colony winter in the Wadden Sea, from which environmental data and information on the size of the main diet, the mussel Mytilus edulis stock exists. We hypothesised that changes in nutrients and water temperature in the Wadden Sea had an effect on the ecosystem affecting the size of mussel stocks, the principal food item for eiders, thereby influencing the number of breeding eider in the Christiansø colony.

Methodology/Principal Finding

A positive relationship between the amount of fertilizer used by farmers and the concentration of phosphorus in the Wadden Sea (with a time lag of one year) allowed analysis of the predictions concerning effects of nutrients for the period 1925–2010. There was (1) increasing amounts of fertilizer used in agriculture and this increased the amount of nutrients in the marine environment thereby increasing the mussel stocks in the Wadden Sea. (2) The number of eiders at Christiansø increased when the amount of fertilizer increased. Finally (3) the number of eiders in the colony at Christiansø increased with the amount of mussel stocks in the Wadden Sea.

Conclusions/Significance

The trend in the number of eiders at Christiansø is representative for the entire flyway population, and since nutrient reduction in the marine environment occurs in most parts of Northwest Europe, we hypothesize that this environmental candidate parameter is involved in the overall regulation of the Baltic/Wadden Sea eider population during recent decades.     

Long-term changes in Wadden Sea nutrient cycles: importance of organic matter import from the North Sea

The Wadden Sea is a shallow tidal area along the North Sea coast of The Netherlands, Germany and Denmark. The area is strongly influenced by rivers, the most important of which are the rivers Rhine, Meuse and Elbe. Due to the increased nutrient load into the coastal zone the primary production in the Wadden Sea almost tripled during the past few decades. A conceptual model is presented that links nitrogen input (mainly nitrate) via Rhine and Meuse with the annual nitrogen cycle within the Wadden Sea. Three essential steps in the model are: (1) nitrogen limits the primary production in the coastal zone, (2) a proportional part of the primary produced organic matter is transported into the Wadden Sea and (3) the imported organic matter is remineralized within the Wadden Sea and supports the local productivity by nitrogen turn-over. The conceptual model predicts that during years with a high nutrient load more organic matter is produced in the coastal zone and more organic matter is transported into and remineralized within the Wadden Sea than during years with low nutrient loads. As a proxy for the remineralisation intensity ammonium plus nitrite concentrations in autumn were used. Based on monitoring data from the Dutch Wadden Sea (1977–1997) the above mentioned model was statistically tested. In autumn, however, a significant correlation was found between autumn values of ammonium and nitrite and river input of nitrogen during the previous winter, spring and summer. The analysis supports that in years with a high riverine nitrogen load more organic matter is remineralized within the Wadden Sea than in years with a low nitrogen load. A comparison with older data from 1960 to 1961 suggests that the remineralisation intensity in the Wadden Sea has increased by a factor of two to three. This is not reflected by a two to three-fold increase in riverine nitrogen load from 1960 to present. It is suggested that the increased remineralisation rates in the Dutch Wadden Sea between the 1960s and the 1980s/1990s are largely caused by an increased nitrogen flux through the Channel and the Strait of Dover and by an increased atmospheric nitrogen input.     

Rapid colonization of new habitats in the Wadden Sea by the ovoviviparousLittorina saxatilis (Olivi)

The intertidal periwinkleLittorina saxatilis completely lacks larval dispersal and adult vagility is low. Although this suggests a low dispersal rate,L. saxatilis is frequently found in recently established habitats “exotic” to the Wadden Sea. Populations occur on man-made structures like dikes, breakwater and groynes, some of which are not older than several years. Furthermore,L. saxatilis is found on marsh grassSpartina anglica, introduced to the Wadden Sea in the 1920s and 1930s, as well as on mats of green macroalgae, which have become an abundant feature on the tidal flats since the late 1970s. Seagrass beds are likely to be the original habitat ofL. saxatilis in the Wadden Sea. Since seagrass populations have dramatically declined over the last decades, colonization of new habitat types enabledL. saxatilis to maintain its Wadden Sea populations despite a changing environment. Colonizers can reach new habitats by means of passive transport, especially by rafting on macrophytes and by aerial dispersal attached to birds. In thew Wadden Sea, the ovoviviparously reproducingL. saxatilis has demonstrated its ability to successfully found new populations with only a few individuals. No reduction of genetic variablility (founder effect) was observed in recently established populations.     

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.     

Eutrophication of the Dutch Wadden Sea: External nutrient loadings of the Marsdiep and Vliestroom basin

The increasing P and N content in the two main tidal basins in the western Dutch Wadden Sea, the Marsdiep and the Vliestroom basin, has been reconstructed from the 50s onwards. The area is enriched with nutrients by two sources both originating from the river Rhine, one being the discharge from Lake IJssel and the other the exchange with the coastal zone of the North Sea. Due to a buffering by Lake IJssel for about 15–20 years, the eutrophication of the western Wadden Sea showed a time lag compared with the continuously increasing nutrient concentrations in the river Rhine and the coastal zone of the North Sea. At present, the primary production in part of the area still seems to be nutrient limited in summer, while loadings have already been decreasing in recent years. So far, no severe, negative effects on the ecosystem have been reported. Some remarks are made on the eutrophication in other parts of the Dutch' Wadden Sea in relation to the hydrographic characteristics of these areas. Presented at the VI International Wadden Sea Symposium (Biologische Anstalt Helgoland, Wattenmeerstation Sylt, D-2282 List, FRG, 1–4 November 1988) Publication No. 18 of the project “Ecological Research of the North Sea and Wadden Sea” (EON)     

Neues Leben im Weltnaturerbe Wattenmeer. Globalisierung unter Wasser

Globalization under water: Alien species in the Wadden Sea World Heritage Recent investigations reveal an increasing number of non‐native species in the Wadden Sea which profit from warmer water temperatures caused by global change. These exotic species achieve highest occurrence and densities in shallow waters near the low tide water level. In this tidal zone, a highly diverse species community of algae and invertebrates became established and will continue to alter in composition. This leads to enhanced complexity of biogenic habitats and to a prevalence of filter feeding organisms. Thus, we observe a fundamental change of the whole Wadden Sea ecosystem which is without return.     

Invasive alien plants in marine protected areas: the <Emphasis Type="Italic">Spartina anglica</Emphasis> affair in the European Wadden Sea

The common cord-grass Spartina anglica, a fertile hybrid of S. maritima and S. alterniflora, was planted in the European Wadden Sea extensively during the late 1920s and 1930s to promote sediment accretion. After establishment, it colonised as a pioneer plant in the upper tidal zone, where it occurs frequently in coherent swards at the seaward front of saltmarshes and in patches on the tidal flats. Often, a conspicuous, almost monotypic, belt of S. anglica is formed. Over the last two decades, an increase in abundance and accelerated spread of S. anglica was observed, possibly promoted by warmer spring temperatures. This alien species may benefit from global warming, and there is considerable concern about its harmful impacts on the native biocoenoses and native biodiversity of the unique Wadden Sea ecosystem, encompassing effects on hydromorphodynamics and coastal protection. For a definitive assessment, however, an adequate quantification and comparison of documented and potential effects of S. anglica is important, but currently unavailable. Consequently, no management strategy exists for the prevention or restoration of the Wadden Sea ecosystem. Thus, the development of an alien species plan on the level of the Trilateral Cooperation on the Protection of the Wadden Sea is essential.     

Monitoring juvenile stocks of flatfish in the Wadden Sea and the coastal areas of the southeastern North Sea

Results are presented of an on-going monitoring program, started in 1970, of the demersal fish population in the Wadden Sea, Scheldt estuary, and along the continental coast between the Belgian-French border and Esbjerg. Particular attention is given to long-term trends in overall abundance and annual variations in spatial distribution of 0- and 1-group plaice and sole in relation to year-class strength and to variations in growth rate. Recruitment level in both plaice and sole appeared to be higher in the 1980s than in the 1970s. The most important nursery areas for plaice are the German Bight and the Wadden Sea, and the relative contributions of these areas to the total plaice stock in different years are relatively stable. In contrast, the most important nurseries for sole are along the continental coast, and the contribution of different parts of these nurseries vary considerably from year to year. Abundant year-classes of sole can originate from northern as well as from southern parts of the nurseries. Relations between survey results and year-class estimates from Virtual Population Analysis (VPA) for plaice are better compared to sole. No trends have been observed in mean length at age 0 and 1 during the years the surveys have been carried out. It is concluded that the quality of the continental coastal area and the Wadden Sea as nursery area for plaice and sole has not declined during the period of investigation. Presented at the VI International Wadden Sea Symposium (Biologische Anstalt Helgoland, Wattenmeerstation Sylt, D-2282 List, FRG, 1–4 November 1988)     

Fundamentals of an applied ecosystem research project in the Wadden Sea of Schleswig-Holstein

The aims, content, and organisatory structure of a proposed interdisciplinary ecosystem research project in the Wadden Sea of Schleswig-Holstein (W. Germany) are briefly presented. The project will include research on both fundamental as well as applied aspects of the Wadden Sea ecosystems and their interaction with local human activities. In contrast to most of the other completed or currently running ecosystem research projects on tidal coasts, a considerable part of the scientific work will also deal with aspects of ecosystem management and protection of the various marine and semiterrestrial habitats of the Wadden Sea. Considerable attention is paid to theoretical and methodological aspects of research on ecosystems and landscape units. In particular, the adoption of a hierarchical view of complex biological and environmental systems is recommended. Presented at the VI International Wadden Sea Symposium (Biologische Anstalt Helgoland, Wattenmeerstation Sylt, D-2282 List, FRG, 1–4 November 1988)     

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.     

Effects of pesticides on seagrass beds

In the German Wadden Sea there has been a remarkable decline in seagrass beds. It was the aim of this study to test whether herbicide contamination could be a reason for this. Concentrations of triazine herbicides such as atrazine, simazine and terbutylazine as well as phenylurea herbicides were measured in Wadden Sea sediments within or in the neighbourhood of seagrass meadows. Sediments were thus used as a marker for medium-term contamination of the Wadden Sea. The respective concentrations were examined in relation to the density and status of the seagrass meadows. Preliminary results show that there may be a connection between seagrass decline and herbicide contamination in the parts of the Wadden Sea sampled. A comparison with other contamination is also given. Received: 11 January 1999 / Received in revised form: 1 December 1999 / Accepted: 8 December 1999     

Earlier pupping in harbour seals,Phoca vitulina

The annual reproductive cycle of most seal species is characterized by a tight synchrony of births. Typically, timing of birth shows little inter-annual variation. Here, however we show that harbour seals Phoca vitulina from the Wadden Sea (southeast North Sea) have shortened their yearly cycle, moving parturition to earlier dates since the early 1970s. Between 1974 and 2009, the birth date of harbour seals shifted on average by −0.71 d yr⁻¹, three and a half weeks (25 days) earlier, in the Dutch part of the Wadden Sea. Pup counts available for other parts of the Wadden Sea were analysed, showing a similar shift. To elucidate potential mechanism(s) for this shift in pupping phenology, possible changes in population demography, changes in maternal life-history traits and variations in environmental conditions were examined. It was deduced that the most likely mechanism was a shortening of embryonic diapause. We hypothesize that this could have been facilitated by an improved forage base, e.g. increase of small fishes, attributable to overfishing of large predator fishes and size-selective fisheries.     

Human transformations of the Wadden Sea ecosystem through time: a synthesis

Todays Wadden Sea is a heavily human-altered ecosystem. Shaped by natural forces since its origin 7,500 years ago, humans gradually gained dominance in influencing ecosystem structure and functioning. Here, we reconstruct the timeline of human impacts and the history of ecological changes in the Wadden Sea. We then discuss the ecosystem and societal consequences of observed changes, and conclude with management implications. Human influences have intensified and multiplied over time. Large-scale habitat transformation over the last 1,000 years has eliminated diverse terrestrial, freshwater, brackish and marine habitats. Intensive exploitation of everything from oysters to whales has depleted most large predators and habitat-building species since medieval times. In the twentieth century, pollution, eutrophication, species invasions and, presumably, climate change have had marked impacts on the Wadden Sea flora and fauna. Yet habitat loss and overexploitation were the two main causes for the extinction or severe depletion of 144 species (~20% of total macrobiota). The loss of biodiversity, large predators, special habitats, filter and storage capacity, and degradation in water quality have led to a simplification and homogenisation of the food web structure and ecosystem functioning that has affected the Wadden Sea ecosystem and coastal societies alike. Recent conservation efforts have reversed some negative trends by enabling some birds and mammals to recover and by creating new economic options for society. The Wadden Sea history provides a unique long-term perspective on ecological change, new objectives for conservation, restoration and management, and an ecological baseline that allows us to envision a rich, productive and diverse Wadden Sea ecosystem and coastal society.     

Response of the Dutch Wadden Sea ecosystem to phosphorus discharges from the River Rhine

The primary production in the western part of the Dutch Wadden Sea (Marsdiep tidal basin) has been reported to have increased by a factor of two since the late 1970s. This doubling of the phytoplankton and the microphytobenthos production has been ascribed to the eutrophication of that area: mean annual phosphate concentrations have increased, whereas the nitrogen concentrations have not. Analysis of the available production data indicates a more than tenfold increase of the production of the phytoplankton in the Marsdiep tidal channel since the early 1950s. The increase in the phytoplankton production in the inner area seems to be of the same order of magnitude. The primary production of the phytobenthos and the secondary production of the macrozoobenthos seems to have increased proportionally to that of the phytoplankton. It is known that the nutrients in the Marsdiep tidal basin originate from the River Rhine. However, there are two possible routes for nutrients to reach that area. One is via the River IJssel and Lake IJssel (varying amounts of fresh water are sluiced out from the latter into the Wadden Sea). The other is along the coast of the North Sea, where the water transport is driven northeastwards by the residual current. Analysis of available data shows that the annual primary production in the Marsdiep tidal basin is mainly determined by the phosphate discharge from Lake IJssel which directly reach that part of the Wadden Sea, rather than from the River Rhine of which the water must run northward along the coast before reaching the western Dutch Wadden Sea. The close relation between fluctations in annual phosphate discharges and fluctations in annual phytoplankton primary production up to more than ten times the 1950 values indicates that primary production in the Marsdiep tidal channel and probably even in the main part of the whole basin, was and is phosphate limited.The mineralization of North Sea particulate organic matter inside the Wadden Sea, previously thought to be the main phosphate source, is questioned. The repercussions of this finding for management are discussed.     

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.     

Distribution patterns of bacterioplankton and chlorophyll-a in the German Wadden Sea

 In a first synoptic evaluation, the temporal and spatial distribution of bacterioplankton and chlorophyll-a were determined in the German Wadden Sea. Three surveys were undertaken in winter, spring, and summer of 1994 using up to eight ships simultaneously between the river Ems and Sylt island. Despite intensive hydrodynamic mixing of the Wadden Sea water, spatial gradients were obvious. The abundance of bacterioplankton ranged from 0.4 to 26×10⁵ ml^–1 and chlorophyll-a varied between <0.1 and 79 μg l^–1. In winter, relatively homogeneous distribution patterns of both parameters with small gradients were found. Highest chlorophyll-a values connected with a highly patchy structure were observed in spring, while in summer both total chlorophyll-a values and the complexity of the distribution pattern had decreased. In contrast, bacterial numbers increased steadily from January to July with the highest bacterial densities and greatest patchiness observed in summer. Moreover, in some regions of the Wadden Sea, a trophic succession of algae as carbon producers and bacteria as consumers was evident. Correlation analysis verified the relationship between bacteria and chlorophyll a, indicating bottom-up control of bacterial abundance in the northern part of the German Wadden Sea. Since the observed regression slope is remarkably low (0.12–0.46) compared to literature values (0.5–0.8), we suggest that the link between phytoplankton and bacteria found here is a special characteristic of the Wadden Sea as a transition zone between the coastal region and the outer North Sea. Received: 26 May 1998 / Accepted:12 November 1998     

Herring and Sprat Abundance Indices Predict Chick Growth and Reproductive Performance of Common Terns Breeding in the Wadden Sea

Herring (Clupea harengus) and sprat (Sprattus sprattus) are the key prey resources of common terns (Sterna hirundo) breeding in the Wadden Sea. Breeding success of the terns has been below average since 2002, coinciding with exceptionally low herring recruitment and sprat abundance. Time series of herring and sprat abundance in the North Sea and in the Wadden Sea were analyzed to explain long-term breeding success and chick development at two common tern breeding colonies. North Sea herring recruitment and sprat abundance in the Wadden Sea explained the largest part of common tern breeding success, both as single variables and in a multiple regression approach. Breeding success showed stronger correlations with herring recruitment indices derived from the North Sea region compared to the Wadden Sea. Also, herring and sprat abundance data explained more variability in breeding success than of more directly responding measures such as growth rate and maximum weight of chicks. Despite spatial and temporal incoherences between fish surveys and the common tern breeding season, breeding success of common terns reflected the abundance of their key prey fish beyond their foraging range and breeding season. We argue that the ecological connectivity between large- and small-scale herring abundance and the responsiveness of common tern breeding success is strong enough to establish a fish–seabird indicator system to be potentially valuable in monitoring and conservation.     

Maximum likelihood population size estimation of harbour seals in the Dutch Wadden Sea based on a mark–recapture experiment

1. The harbour seal population Phoca vitulina in the entire Wadden Sea was severely depleted due to a virus-epizootic during 1988. A comprehensive study on the population biology and activity patterns was subsequently initiated to design a management and conservation plan. The main objective of this study was to estimate harbour seal abundance in the different regions of the Wadden Sea.
2. We investigated the potential of a mark–recapture experiment using VHF radio-tags in combination with repeated aerial surveys to estimate the number of harbour seals in the Dutch part of the Wadden Sea. The number of harbour seals hauled-out and the presence of any radio-tagged seals was monitored during seven aerial surveys of all known haul-out sites in the Dutch Wadden Sea over the 1994 breeding season.
3. A maximum likelihood (ML) estimator was developed to infer the rate of tag-loss and the size of the local prepupping population.
4. The ML estimate of the number of harbour seals in the Dutch Wadden Sea was 1536 (95% confidence limits were 1225 and 2200). The corresponding maximum proportion of seals hauled-out was 68%.
5. The use of VHF radio-tags which can be monitored from the air provides a way of correcting aerial survey counts for the proportion of harbour seals hauled-out during the surveys. Since haul-out behaviour may be influenced by local conditions, such as exposure time of sand banks, we recommend this technique be repeated in other areas of the Wadden Sea rather than using the estimates from the current study in other areas.     

Macrozoobenthos on an intertidal mudflat in the Danish Wadden Sea: Comparisons of surveys made in the 1930s, 1940s and 1980s

To assess whether long-term faunal changes have occurred on intertidal flats in the Danish Wadden Sea, results of faunal surveys in the Skallingen area during the 1930s, 1940s and 1980s were compared. Since the earlier studies, two species have disappeared (Scrobicularia plana andPetricola pholadiformis) and two have invaded the area (Tharyx killariensis andEnsis americanus). This is, however, not a local event as species changes have occurred on a larger scale (Wadden Sea region). Comparison of abundance data did not provide evidence of changes from the 1930s until the 1980s. Spatio-temporal fluctuations in two dominant species, the mudsnailHydrobia ulvae and the cockleCerastoderma edule chosen for closer examination, could be explained by natural processes. Growth data on cockles from the 1930s and the 1980s matched perfectly. During both periods, cockles showed a much lower growth rate than generally reported from the Wadden Sea, while at the same time they occurred at high densities (>2000 ind·m⁻²). Intraspecific competition among cockles is suspected as being a major cause of the low growth rates. Estimates of secondary production and P/B-ratios of cockles were also in general agreement during the 1930s and the 1980s when corrected for differences in the age structure of the cockle populations. With the possible exception ofMytilus edulis, which according to some observations has extended its range along the low-water line, the present comparison failed to demonstrate long-term faunal changes caused by increased eutrophication. This results is expected to be representative for intertidal flats not exposed to direct impacts from terrestrial run-offs and waste-water discharges.