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     

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.     

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.     

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.     

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.     

The exploitation of living resources in the Dutch Wadden Sea: a historical overview

An overview, based on written sources and personal observations, is presented of exploitation of living resources in and around the Dutch Wadden Sea during the past few centuries. It is concluded that before about 1900 exploitation was almost unrestricted. Exploitation of plants has been documented for saltmarshes and eelgrass beds. Fisheries have occurred for two species of hydroids, two species of polychaetes, one echinoderm species, at least seven species of molluscs, three species of crustaceans, and tens of species of fish. Hunting and egg collecting targeted almost all species of birds. Finally, two species of seals were exploited: information on exploitation of cetaceans is not available. Hence, it is likely that overexploitation may have been involved in the extirpation of several species. This supports an earlier suggestion that overexploitation played a part in the disappearance of at least 17 species from the Dutch Wadden Sea. This conclusion is confirmed by the observation that several extirpated species have returned after protective measures were introduced for the Dutch Wadden Sea area.     

Coast of change: habitat loss and transformations in the Wadden Sea

In the southern North Sea, coastal people commenced with habitat conversions 1,000 years ago. Partly interrupted in late medieval times by large-scale inundations of marshland, progressive embankments transformed the landward half of the amphibic transition zone between a limno-terrestric and a brackish-marine ecosystem into arable land and freshwater lakes. Sea walls rigidly separated the land from the sea. Dynamic transitional habitats have vanished. Areal loss has diminished the capacity of the Wadden Sea to dissipate wave and tidal energy. A coastal ecosystem once rich in marsh plants, seagrass and diatoms on mud flats became transformed into one with less autochthonous phototroph production, dominated by sandy tidal flats, and dependent primarily on allochthonous plankton supply. The large estuaries have been dredged to serve as shipping canals, and have lost most of their former retention and filter capacity. Riverine loads are now flushed right into the North Sea. Symptoms of a syndromatic coastal habitat degradation are diagnosed, leading to a decline in natural habitat diversity. The conventional on-line coastal protection may not achieve a sustainable coastal habitat configuration. At sedimentary coasts immobilised by dikes and petrified shores, a more flexible response to sea level rise is recommended.     

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.     

Common patterns in the fluctuations of macrozoobenthic species living at different places on tidal flats in the Wadden Sea

During 17 successive years (1969–1985) the macrozoobenthos has been sampled quantitatively in the latewinter/early-spring period at 15 stations scattered over Balgzand (a 50 km² tidal flat area in the western-most part of the Wadden Sea) and at 5 stations located in a small (3 km²) area 150 km away from Balgzand in the eastern part of the Dutch Wadden Sea. In 25 species, numbers per m² were, in most years, sufficiently high at 2 or more of the Balgzand stations to allow between-station comparisons of fluctuation patterns. Comparisons were made by rank correlation. Out of a total of 1003 of such comparisons that could be made with the Balgzand data, 47% yielded significantly positive correlations and less than 1% significantly negative ones. Thus, nearly half of the fluctuation patterns of the populations living at the 15 stations within the Balgzand area showed a high similarity. Synchronization of population fluctuations was augmented particularly by the incidence of severe winters (causing low spring numbers in about a quarter of the species that were sensitive to low temperatures and high reproductive success in several species during the subsequent summer) and further by the increasing trends in numbers in about half of the species, probably as a consequence of eutrophication.Similar results as on Balgzand were obtained within the restricted area Groninger Wad in the eastern part of the Dutch Wadden Sea (Essink & Beukema, this issue). Comparison of the fluctuation patterns between the 2 distant areas also yielded high numbers of significantly positive correlations, though the proportion of the patterns that were similar was lower than these proportions were within the 2 areas.It is concluded that common patterns of fluctuation in numerical densities of macrobenthic species can be assessed over vast areas. Such common patterns will represent the normal or base-line fluctuations that may be used to distinguish (as departures from such patterns) the effects of local disturbing influences.     

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.     

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.     

On the dynamics of the stocks of blue mussels (Mytilus edulis L.) in the Danish Wadden Sea

As biological basis for the monitoring programme for the commercially exploited stock(s) of mussels (Mytilus edulis L.) in the Danish Wadden Sea, samples of mussels have been collected regularly since 1986, both from sub-tidal and inter-tidal mussel beds. These samples are the basis for the estimation of total biomass. They also provide data on size frequency distributions, which have been analysed for cohort identification resulting in length at age data, which again have been used for estimating parameters (L and K) for the von Bertalanffy growth equation (VBGE) as well as mortality parameters. By applying these in the Beverton & Holt model, estimates of average biomass and annual production (P) of the mussels have been obtained together with possible fisheries yields from the beds. The growth and mortality parameters and the figures for annual production and P/B are compared with figures from other investigations. These analyses have been the basis for annual assessments of the mussel stocks, which again are used in the current management of mussel fishery in the Danish Wadden Sea.     

Bulk chemical characterization of particulate material from the Jade Bay,Lower Saxonian Wadden Sea

Data on the bulk chemical composition (C, N, P, S, total chlorophyll) of particulate matter were obtained on five occasions over one or two tidal cycles in the Jade Bay, Lower Saxonian Wadden Sea. Sampling intervals were usually one hour. The results show pronounced short-term variability for all parameters which is controlled to a large extent by physical processes such as erosion of surficial sediments at high current speeds and sedimentation at slack water. Contributions from living phytoplankton biomass were low while C∶N ratios indicate a high abundance of microheterotrophs. Organic phosphorus accounted for 36% of total P, exchangeable ammonium made up 1.9% of total nitrogen on average.     

笋壳过氧化物酶提纯及综合利用的研究

采用挥发性酸和碱调节鲜笋壳压榨液 pH值 ,分别获得无盐Fraction Ⅰ和fraction Ⅱ ;其压榨后的蒲渣作为造纸原料和栽培食用菌的基质 ;对分离Fraction Ⅰ和Fraction Ⅱ后的汁液 ,加入Fe2 +并超滤浓缩 ,经硫酸铵盐析和丙酮分级后 ,进行CM 2 2纤维素离子交换柱层析 ,以 pH5 .2乙酸钠分级洗脱 ,获得RZ为 3 .0以上的高纯竹笋过氧氧化物酶。揭示了竹笋加工成罐头食品后其尾料通过系统生化工程处理 ,获得系列产品的途径 ,减少了生产成本和环境污染 ,提高了主产品附加值 ,具有生态、经济和社会效益。     

Ecology and Bioproduction in the Microphytobenthos of the Chain of Shallow Inlets (Boddens) South of the Darss-Zingst Peninsula (Southern Baltic Sea)

The significance of epipsammic microphytobenthos bioproduction in a shallow estuary was investigated by the oxygen method. Bioproduction reached a maximum in May or June. A close correlation was found between the gross production rate and algal biomass at station K. The biomass and production rates were distinctly lower at station B than at station K as station B was more exposed. Circadian variations and depth profiles of the gross production rate correlated best with irradiation intensity. Mean gross production from April to October was 1761 O₂.m⁻² and 75 g C·m⁻².     

An evaluation of small-scale genetic diversity and the mating system in Zostera noltii on an intertidal sandflat in the Wadden Sea

Background and Aims

The dwarf eelgrass, Zostera noltii, is a predominant inhabitant of soft-bottom intertidal regions along the coasts of northern Europe. It is a monoecious, protogynous angiosperm in which the potential for self-fertilization and inbreeding are high, especially if clone sizes exceed pollen dispersal distances. The aim of the present study was to determine the relationship between mating system and clonal structure, examine the relative roles of geitonogamous selfing and biparental inbreeding, measure pollen availability (multiple paternities) and estimate pollen dispersal.

Methods

A 100-m² plot was established in a large, intertidal Z. noltii meadow on the island of Sylt in the German Wadden Sea. A total of 256 adult shoots was sampled: one from the centre of 100 fixed 1-m² quadrats (large scale resolution) and an additional 156 from within eight randomly selected 1-m² sub-quadrats (small-scale resolution). DNA was extracted from seeds and leaf tissue of all samples and genotyped with nine microsatellite loci.

Key Results

Mating system analysis revealed high multilocus and single locus outcrossing rates. Average pollen dispersal distance was nearly the same as the average genet (clone) size. Multiple paternity was common and 20–30 % of mature seeds originated from matings within the plot. Among inbred seeds, most resulted from geitonogamy rather than biparental inbreeding.

Conclusions

Moderate disturbances intrinsic to the intertidal habitat appear to facilitate seed recruitment by gap formation. Pollen dispersal distances are sufficient to maintain outcrossing and high clonal diversity.     

Evidence for metapopulation structuring in cod from the west of Scotland and North Sea

The scale of population structuring in cod Gadus morhua from the west of Scotland (ICES stock area VIa) and northern North Sea was investigated using a combination of non-genetic methods. Site fidelity of spawning aggregations was examined using historic tag-recapture experiments and individual geolocation estimates from data storage tagged fish. The extent of movements from spawning areas indicated by these two tagging approaches was broadly similar. Between 67 and 97% of adult cod remained within 100 km of spawning areas throughout the year, suggesting resident spawning groups. A small proportion of cod, however, did appear to stray between spawning areas. A comparison of the elemental signature of the otoliths of 0 year-group and the 0 year-group component from the same year-class at age 2 years indicated that most adults originate from local nursery areas. Moreover, there did not appear to be detectable exchange between spawning areas >200 km apart, such as the Clyde and Minch or Shetland and the Inner Hebrides. As such, population processes may operate at a smaller spatial scale than the stock level, with spawning aggregations functioning as local populations within a metapopulation. The relevance of this scale of dynamics is discussed in relation to recently imposed closed areas in the region.