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.     

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.     

An analysis of mussel bed habitats in the Dutch Wadden Sea

A habitat suitability analysis for littoral mussel beds in the Dutch Wadden Sea was carried out. The analysis was based on the presence of mussel beds in the years 1960–1970, and a number of environmental characteristics: wave action, flow velocity, median grain size, emersion times and distance to a gully border. The habitat model describes mussel bed appearance quantitatively. It predicts the distribution of mussel beds quite well, as well as the distribution of spatfall in the years 1994 and 1996. From the analysis we found that wave action (maximum orbital velocity) was the main structuring factor. A low orbital velocity was preferred. Neither very low, nor maximum flow velocities were favourable for mussel beds. Very coarse sands or silty environments were not preferred. Sites close to the low water line showed lower mussel bed appearance; when emersion time was above 50% , hardly any mussel beds could be found. The habitat suitability analysis and the construction of a habitat suitability map was performed in the framework of the discussions on a further or reduced exploitation of the tidal flats in the Dutch Wadden Sea by cockle and mussel fishery activities. Electronic Publication     

General Patterns in Invasion Ecology Tested in the Dutch Wadden Sea: The Case of a Brackish-Marine Polychaetous Worm

The success of invasive aquatic species is determined by a variety of attributes such as wide environmental tolerance, high genetic variability, short generation time, early sexual maturity, high reproductive capacity, and a broad diet. Usually, introduced species, after some time lag since inoculation, show an exponential population increase and expansion. Maintenance of the immigrant species at a high population level will be dependent on interspecific competition with native species and availability of habitat and food. Eventually, the immigrant population may decline, for instance due to increased predation pressure, parasite infestation or loss of genetic vigour. These characteristic patterns in invasive species are reviewed for the case of the North American spionid polychaete Marenzelleria cf. wireni in the Dutch Wadden Sea. This species was first recorded in estuaries and coastal waters of the European continent in the Ems estuary (eastern Dutch Wadden Sea) in 1983. In the western part of the Dutch Wadden Sea the first specimens were found in 1989. The Ems estuary population showed the typical lag-phase, explosive increase, stabilisation, and eventual decline. In the western part of the Dutch Wadden Sea the latter two phases have not yet developed. The strong development and stabilisation of the population in the Ems estuary may have been caused by the availability of a yet not utilised food source. The species' final decline remains largely unexplained.     

The Balgzand (Dutch Wadden Sea) tidal flats as a source of soluble silicate

Summary The huge numbers of molluscs, worms and bacteria found in the sheltered tidal flats of the Dutch Wadden Sea, decompose large amounts of organic matter. In this process nutrients incorporated in the particulate organic matter are transferred to the dissolved state. The importance of this process for nutrient budgets was first demonstrated by POSTMA (1954) and later by various authors with emphasis on tidal watersheds and sheltered bays (see Neth. J. Sea Res., 8 (2/3). 1974).The nutrient budgets of the Balgzand tidal flats, as a typical example of a sheltered bay, have been studied during one tidal period by TIJSSEN and VAN BENNEKOM (1976) and recently for a longer period in a cooperative project by the Studiedienst Hoorn of Rijkswaterstaat (DE BOER, 1978) and the Netherlands Institute for Sea Research (MANUELS, 1978). A qualitative measure for the high intensity of mineralization are the high concentrations of nutrients in the water on the Balgzand, compared to those in the Marsdiep (Table I).Simple box models can be used to derive the intensity of nutrient production from the concentration differences in which also the water depth and the exchange velocity of the water with adjacent areas contribute. As a first approximation all mineralization is thought to be situated in the surface sediment (VAN BENNEKOM et al., 1974) and the characteristics parameter becomes the nutrient production per m² per day. For reactive silicate this parameter shows a pronounced seasonal cycle. In January, February and Marchin situ production is hardly detectable while in July and August 500 mg SiO₂ m^–2d^–1 is found. For comparison, weathering of soil by rain water gives 4–30 mg SiO₂.m^–2d^–1 with only slight seasonal variations. The difference is at least partially explained by assuming a different source: diatom frustules in the case of the Balgzand and silicate minerals in the case of normal weathering.The scarce data available indicate that the seasonal cycles of in situ production of phosphate, nitrate and ammonia on the Balgzand are less pronounced than for reactive silicate. The picture is further complicated because ammonia is always produced, but sometimes nitrate is consumed, apparently by denitrification.The nutrient ratios change during the year (Table II). Especially in summer the contribution of silicate is relatively large. This is in marked contrast with the nutrient ratios in the coastal sea water where during the summer the amount of silicate is relatively low. The nutrient ratios are probably an important factor in the seasonal and regional species composition of the phytoplankton.     

A sketchy outline of the fate of organic matter in the Dutch Wadden Sea

Conclusion In ecosystems like the Dutch Wadden Sea a carbon budget can be made roughly However, there are in space as well as in time a lot of gaps in the observations. Especially from subtidal sediments of inlets and gullies are no observations available. Also the quantification of the different functional groups of the anaerobically respiring organisms (manganese reducers, denitrifiers, iron reducers, methane formers) in the decomposition is absent. And what is the role and quantitative importance of fermenting organisms? Finally, the quantification of the use of the reduced end products of anaerobic respiring organisms is unknown.     

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)     

Chemical monitoring in the Dutch Wadden Sea by means of benthic invertebrates and fish

In monitoring, it is of utmost importance to carefully define the purpose, the sampling strategy, as well as the analytical chemical and statistical requirements. Surveys are appropriate for describing the geographical variation in environmental contaminant levels. Repeated surveys and recurrentdata collection at permanent locations provide means of detecting temporal trends. Results are presented here of surveys on pollution by trace metals, polychlorinated biphenyls and organochlorine pesticides in the Ems Estuary and Dutch Wadden Sea usingMytilus edulis, Mya arenaria, Arenicoia marina, Nereis diversicolor andCrangon crangon as test organisms. Trends towards decreasing pollution by mercury are illustrated by monitoring data onMytilus edulis andZoarces viviparus. It is stressed that the results of chemical monitoring in organisms may be interpreted only in termser the biological effects on the basis of relevant toxicological knowledge and/or additional bio-assays. Presented at the VI International Wadden Sea Symposium (Biologische Anstalt Helgoland Wattenmeerstation Sylt, D-2282 List, FRG, 1–4 November 1988)     

Now an empty mudflat: past and present benthic abundances in the western Dutch Wadden Sea

The benthic fauna of two areas in the western Dutch Wadden Sea, Posthuiswad and Staart van Schieringhals, was described in 1930–1960 and again between 1996 and 2005. Here, we document the changes. Whereas both areas formerly had high densities of species that biogenically structured the intertidal mudflats such as mussels Mytilus edulis and cockles Cerastoderma edule, by 1996 they had shown a tenfold decrease in the densities of molluscs, with no recovery till 2005. Although the number of species of polychaetes and crustaceans may not have changed much, their relative abundance did. Nowadays, more polychaete species are common than before. We briefly discuss whether the changes in benthic community composition could be due to industrial fishery practices or eutrophication effects.     

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)     

An estimate of the sustainable rate of shell extraction from the Dutch Wadden Sea

1. Shell production by cockles Cerastoderma edule was studied to examine whether or not the present licensed rate of shell extraction in the Dutch Wadden Sea exceeds the current rate of shell addition to the exploitable stocks.
2. Long-term data on numbers of cockles and weights of their shells were used to estimate their annual production on Balgzand, a 50-km² tidal flat area in the western-most part of the Wadden Sea. During the 1969–97 period, it amounted to an average of 125 g m^–2, including 107 g m^–2 of shells large enough to be exploitable for shell-lime fishery.
3. The very irregular annual recruitment of cockles was the main cause of the wide 95% confidence limits (74 and 140 g m^–2 year^–1) of this 28-year estimate. Moreover, high mortality rates in severe winters substantially reduced production per recruit in some year classes.
4. About one-third of the estimated production does not reach exploitable stocks, because it is fragmented by birds (particularly eider ducks), permanently buried in the sediment, or removed by the fishery for live cockles.
5. During the last few decades, the estimated mean amount added annually to the exploitable stocks was 88 million kg or 132 000 m³ of large cockle shells. This amount compares favourably with the current annual level of removal of 134 000 m³ of shells, three-quarters of which are cockles.
6. Even at temporarily lower production rates, the exploitation of shell stocks at its present rate is not expected to lead to a rapid exhaustion of the existing stocks in the tidal inlets of the Dutch Wadden Sea, as these stocks will be in the order of a few million m³.     

Recent changes in the contributions of river Rhine and North Sea to the eutrophication of the western Dutch Wadden Sea

From 1955 to the mid 1980s the loads of both nitrogen and phosphorus from the river Rhine to the Dutch coastal area, the Wadden Sea included, increased. Since 1985 the phosphorus loads has decreased significantly, while the nitrogen load remained about the same.Annual primary production in the western Dutch Wadden Sea has increased fromc. 40 g C m^–2 (1950) to 150 (mid 1960s) and over 500 g C m^–2 (1986). The biomass of macrozoobenthos has more than doubled since 1970. Simultaneously, the meat yield of cultured blue mussels (Mytilus edulis), has increased since the 1960s. Previously, it was indicated that the increase in primary production of the phytoplankton over the period 1950 to 1986 was stimulated by the load of dissolved inorganic phosphate from Lake IJssel, a reservoir supplied by Rhine water. Since 1990, however, primary production has been higher than was expected from decreased phosphate loads from Lake IJssel. It is argued that this lack of response may have been caused by increased concentrations of dissolved inorganic phosphate at sea originating from increased inflow from a.o. the Strait of Dover, which compensate for the decrease in phosphate from the rivers, possibly in combination with a significant improvement of the light conditions of the water in the Wadden Sea.     

The political culture of poaching: a case study from northern Greece

Poaching has deep social and cultural roots and its meanings are multi-layered. This article explores the meanings attached to the practice of illegal hunting and fishing around Lake Kerkini in northern Greece. Here poaching must be considered in the context of a disordered ecosystem, where the dominance of locally maligned fish and bird species results from economic and environmental policy designed to benefit distant farmers. We conclude that poaching cannot be understood only as an individual action, but as one where collective and personal identities are defended in the face of seemingly irrevocable economic and social decline. The discussion shows that poachers identify different kinds of poaching. Some of the most apparent forms of poaching, done by local inhabitants, may be less damaging than other commercially oriented forms, including by outsiders. Poaching is motivated through a complex mix of factors. Our data lead us to discuss two manifestations of poaching (a) poaching as a form of collective resistance; and (b) poaching as a violation of culturally valued types of human-nature interaction. Some people who admit undertaking what they perceive as least detrimental forms of poaching are antagonistic towards what they construe to be truly harmful forms. Such people appear willing to act and to support actions against types of poaching they agree to be threatening. This is a message with potential importance for environmental management strategy.     

Accumulation rates of soil organic matter in wet dune slacks on the Dutch Wadden Sea islands

Background and aims

A long-term monitoring program (ranging from 16 to 77 years) on the Dutch Wadden Sea Islands provided well-documented examples of vegetation succession in wet dune slacks. We used this opportunity to study soil organic matter (SOM) accumulation in relation to vegetation succession. The aim of this paper is to identify the factors which regulate accumulation rates of SOM in wet dune slacks.

Methods

We used several soil chronosequences using data from the monitoring program together with data from a long-term research activity and more recent measurements. We used several soil chronosequences using data from the monitoring program together with data from a long-term research (up to 150 years) and more recent measurements. Field measurements included pH, soil organic matter, above ground standing crop and water levels. Water level regimes (inundation duration and mean minimum water level), were simulated using a hydrological model. Capable of simulating inundation duration and water-level fluctuations, this model used field measurements collected over more than 5 years, as well as precipitation and evapotranspiration data collected over a period of 25 years.

Results

Sampling two synchronic chronosequences showed that SOM accumulations increased linearly during the first 50–60 years and then levelled off. Sampling various diachronic chronosequences over time showed a wide variation in accumulation rates. Slacks with low productive species, such as Littorella uniflora, showed low accumulation rates (0.02–0.08 kg/m²/year), and persisted even over a period of more than 90 years. In contrast, slacks dominated by high productive species, such as Phragmites australis, showed ten times higher accumulation rates (0.17–0.26 kg/m²/year) over a similar time period and comparable annual inundation periods (176–240 days). A multiple linear regression showed that variation in SOM accumulation rates was best explained by above-ground biomass of the vegetation.

Conclusions

We conclude that the rate of SOM accumulation in wet dune slacks is primarily controlled by plant above-ground biomass. Both above-ground biomass and SOM accumulation can remain very low over a long period of time when dune slacks are flooded during most of the year and plants with adaptive traits are able to maintain vegetation succession at a pioneer stage.     

Variations in the mussel population of the Dutch Wadden Sea in relation to monitoring of other ecological parameters

The pattern of distribution of intertidal mussel beds is relatively constant over a number of years although their surface area can vary greatly. The abundance of mussels shows much greater fluctuations. In the western part of the Dutch Wadden Sea, west of the Terschelling tidal divide, the amount of mussels on natural beds fluctuated between 1 and 24 million kg fresh weight during the years 1949 to 1988. In the eastern part of the Dutch Wadden Sea the biomass varied between 5.5 and 180 million kg. The influence of the mussels on the ecosystem therefore can be very different between years. When many mussels are present the whole watermass can be filtered every few days. In years with few mussels present the filtering may take one month. It is argued that monitor programmes for a.o. nutrients, chlorophyll and growth rates of benthic organisms are of limited value if there is no indication about the total amount of mussels in the area. Presented at the VI International Wadden Sea Symposium (Biologische Anstalt Helgoland, Wattenmeerstation Sylt, D-2282 List, FRG, 1–4 November 1988)     

Long-term changes in macrozoobenthic abundance on the tidal flats of the western part of the Dutch Wadden Sea

For 20 years (1969–1988), larger bottom animals have been sampled quantitatively once or twice annually at 15 stations scattered over Balgzand (a 50 km² tidal flat area in the westernmost part of the Dutch Wadden Sea). In 29 species, numbers were sufficiently high to allow a statistical evaluation of the fluctuation patterns of their annual densities. The results revealed two main trends: (1) a sensitivity to low winter temperatures in 12 species, causing low densities in these species immediately after a severe winter (1979, 1985, 1986 and 1987) and relatively high densities during a period with some mild winters in succession (1973–1975); (2) an upward long-term trend in 11 (other) species, causing upward trends (viz. roughly a doubling) of total macrozoobenthic biomass and production over the 20-year period of observation, probably as a consequence of increasing eutrophication. By far the major part of the species thus exhibited either of these two patterns, causing total biomass and species number to be governed largely by the above two trends. Results of less frequent sampling (once per 5 or 10 years) of 26 transects scattered over the ≈500 km² of tidal flats of the whole western half of the Dutch Wadden Sea showed that the two trends also represent the changes occurring in a much larger area. Some local departures from the general patterns are discussed and related to specific causes. Presented at the VI International Wadden Sea Symposium (Biologische Anstalt Helgoland, Wattenmeerstation Sylt, D-2282 List, FRG, 1–4 November 1988)     

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.     

The Birth,Growth and Death of Intertidal Soft-Sediment Bivalve Beds: No Need for Large-Scale Restoration Programs in the Dutch Wadden Sea

Ecosystems - Recruitment and fate of all 1436 mussel and oyster beds of the Dutch Wadden Sea were studied for the period 1999–2013. Cox’s proportional hazard rate model with covariates...     

The importance of intraspecific competition in a Littorina littorea population in the Wadden Sea

Two field experiments were carried out totest whether effects of intraspecific competition ina Littorina littorea population can be detectedin a short-term investigation. Different size classesof L. littorea showed no significant differencein preferences when offered four kinds of eitherpossible food or substrata (Fucus vesiculosus,Ulva lactuca, Carcinus maenas, brick).Large and medium winkles preferred Fucusvesiculosus, followed by Ulva lactuca. Deadshore crabs (Carcinus maenas) were the leastpreferred objects for all size classes. On the firstday of the experiment bricks were more attractive tosmall littorines than to larger ones. Considering allfour days, the same ranking occurred for all sizeclasses: Fucus vesiculosus > Ulvalactuca > brick > Carcinus maenas. The reactionofjuveniles to increased densities was examined using anin situ caging experiment on a mussel bed. Meshsize of the cages allowed adult densities to beincreased while juveniles could escape by passingthrough the meshes. However, there was no significantemigration of small winkles even from cages with 10 to20 times natural density of large individuals. Ofgreater importance was the original number of winklesat the site. The available resources on the musselbeds appear to be sufficient to maintain a highpopulation density. Intraspecific competition does notseem to play a major role in this L. littorea-population.