Variability of the water temperature in the western Wadden Sea on tidal to centennial time scales

Daily observations of the sea surface temperature in the Marsdiep tidal inlet, which connects the shallow Dutch western Wadden Sea with the deeper North Sea, already started in the summer of 1860, over 140 years ago. Since the year 2000 the sampling frequency has strongly increased because of the use of electronic sensors and data logging by computer. Analysis of these temperature data has revealed variations with time scales from tidal, daily, seasonal, inter-annual, to centennial. The tidal temperature variations are generated by advection of the seasonally varying temperature gradient between Wadden Sea and North Sea, while the daily variations are mainly caused by the daily variation of solar radiation. The seasonal variation in sea surface temperature only lags a few days behind the coastal surface air temperature, contrary to the sea surface temperature in the deeper nearby North Sea, which is delayed with about 1 month. The North Atlantic Oscillation index has been used as large-scale proxy for the atmospheric forcing of the Wadden Sea temperature. Only for the winter and spring a significant correlation is found between temperature and the winter index. However, this correlation is so strong that also the annual mean temperature is correlated significantly with the North Atlantic Oscillation. At longer time scales, from decadal to centennial, also large temperature variations are observed, of the order of 1.5 °C. However, these are not related to long-term changes of the North Atlantic oscillation. These long-term temperature changes involve a cooling of about 1.5 °C in the first 30 years of the record and a similar warming in the last 25 years. In between, these long-term changes were smaller and more irregular. Similar conclusions can also be applied to individual seasons as well as to the date of the onset of spring.     

Variability of the salinity in the western Wadden Sea on tidal to centennial time scales

Daily observations of the salinity of the Marsdiep tidal inlet, which connects the Dutch western Wadden Sea with the North Sea, already started over 140 years ago, in 1860. Since the year 2000 the sampling frequency has increased because of the use of electronic sensors. Analysis of these salinity data have revealed variations on time scales from tidal (~ 12 hour), seasonal, inter-annual, and multi-decadal, to centennial. The contributions of the salinity variations in the Marsdiep for these different spectral bands or time scales are all of the order of a standard deviation of 0.5 to 1. The centennial variation, which can be expressed as a 140 year long salinity trend, is related to engineering works on the rivers Rhine and IJssel, which already started in the early 18-th century, and more than doubled the magnitude of the freshwater content of the western Wadden Sea since then. In contrast with this anthropogenic salinity trend, the climatic variability of the precipitation over western Europe, and the connected changes in the Rhine discharge, are mainly responsible for the inter-annual variations in the salinity and/or freshwater content of the western Wadden Sea. Since variations in salinity and freshwater content also reflect variations in the terrigeneous and river influence on the Wadden ecosystem, e.g. via the nutrient content, it can be expected that the ecology of the Wadden Sea also experienced changes on centennial time scales.     

Benthic photosynthesis in submerged Wadden Sea intertidal flats

In this study we compare benthic photosynthesis during inundation in coarse sand, fine sand, and mixed sediment (sand/mud) intertidal flats in the German Wadden Sea. In situ determinations of oxygen-, DIC- and nutrient fluxes in stirred benthic chamber incubations were combined with measurements of sedimentary chlorophyll, incident light intensity at the sediment surface and scalar irradiance within the sediment. During submergence, microphytobenthos was light limited at all study sites as indicated by rapid response of gross photosynthesis to increasing incident light at the sea floor. However, depth integrated scalar irradiance was 2 to 3 times higher in the sands than in the mud. Consequently, gross photosynthesis in the net autotrophic fine sand and coarse sand flats during inundation was on average 4 and 11 times higher than in the net heterotrophic mud flat, despite higher total chlorophyll concentration in mud. Benthic photosynthesis may be enhanced in intertidal sands during inundation due to: (1) higher light availability to the microphytobenthos in the sands compared to muds, (2) more efficient transport of photosynthesis-limiting solutes to the microalgae with pore water flows in the permeable sands, and (3) more active metabolic state and different life strategies of microphytobenthos inhabiting sands.     

Spatial and temporal variability in bio-optical properties of the Wadden Sea

The Wadden Sea, a shallow coastal area bordering the North Sea, is optically a complex area due to its shallowness, high turbidity and fast changes in concentrations of optically active substances. This study gathers information from the area on concentrations of suspended particulate matter (SPM), Chlorophyll-a (Chl-a), and Coloured Dissolved Organic Matter (CDOM), on total absorption and beam attenuation, and on reflectances from the whole area. It examines the processes responsible for variations in these. Sampling took place at 156 stations in 2006 and 2007. At 37 locations also the specific inherent optical properties (SIOPs) were determined. Results showed large concentration ranges of 2–450 (g m^-3) for SPM, 2–67 (mg m^-3) for Chl-a, and 0–2.5 m⁻¹ for CDOM(440) absorption. Tides had a large influence on the SPM concentration, while Chl-a had a mainly seasonal pattern. Resuspension lead to a correlation between SPM and Chl-a. The absorption of CDOM had a spatial variability with extremely high values in the Dollard, although the slope of CDOM absorption spectra was comparable with that of the North Sea. The Chl-a specific pigment absorption proved to be influenced by phytoplankton species and specific absorption of non-algal particles at 440 nm was correlated with the mud content of the soil at the sample locations. SPM specific absorption was not found to correlate with any measured factor. As the concentrations of optically active substances changed, we also found spatial and temporal variability in the absorption, beam attenuation and reflectances. Reflectance spectra categorized in groups with decreasing station water depths and with extreme CDOM and SPM concentrations showed distinguishable shapes.     

Endoparasites in common eiders Somateria mollissima from birds killed by an oil spill in the northern Wadden Sea

Mass mortalities of common eiders Somateria mollissima have been ascribed to high parasite loads. However, the actual role of parasites in mortalities is disputed as in the case of a mass mortality of eiders in the Wadden Sea in the winter of 1999/2000. A critical evaluation of the role of parasites in eider mass mortalities is hampered by (1) a lack of data on actual parasite loads of the birds involved, (2) missing regional data for comparison, and (3) a lack of unbiased samples: investigations are often based on dead beached individuals, which are presumably the more heavily infected birds of a population and thus more likely to die and be washed ashore. Although published data on parasite loads in birds of the winter 1999/2000 mortality are available, no data on background parasitism in eiders from the Wadden Sea exist, making an evaluation of the potential role of parasites in this mortality event difficult. By investigating endoparasites of 102 eiders affected by an oil spill in the northern Wadden Sea in winter 1998/1999, we provide a data set of background parasitism in wintering eiders from the Wadden Sea. We found 13 different parasite taxa with high prevalence values (% infected birds) in the acanthocephalan Profilicollis botulus, the nematode Amidostomum acutum, cestodes and trematodes. In some taxa we observed pronounced differences in prevalence values between juvenile eiders and adults, as well as between adult sexes. The parasite composition shows that bivalves, crabs (Carcinus maenas) and other crustaceans are important sources of infections by being intermediate hosts. This is partly mirrored in the food content of eider stomachs where bivalves and crabs were predominantly found. Intensities of the acanthocephalan P. botulus, suspected of causing eider mortalities, were especially high in juveniles (1112 ± 416 ind per infected host), but lower in adult males (40 ± 7) and adult females (81 ± 18). However, no extraordinary mortality event was observed in the winter of 1998/1999, indicating no or a very weak effect of the parasites on host condition. A comparison with the parasite loads of eiders from the mass mortality in the winter of 1999/2000 shows that parasite numbers were by no means exceptional for birds from the area. Hence, parasites alone are unlikely to have caused this mortality. Regional background parasite loads are important to differentiate between primary and secondary roles of parasites in anomalous mortality events.     

Seasonality of microzooplankton grazing in the northern Wadden Sea

A sequence of nine dilution experiments was conducted according to Landry and Hassett [Landry, M.R., Hassett, R.P., 1982. Estimating the grazing impact of marine microzooplankton. Mar. Biol. 67, 283–288] in the northern Wadden Sea from March until October 2004 to investigate the seasonality of microzooplankton grazing. From March until April, no grazing was observed. Microzooplankton grazing started in May (0.66 d^− 1) and increased until August (1.22 d^− 1). In October microzooplankton grazing was low again (0.17 d^− 1). Phytoplankton growth rates varied between 0 and 1.1 d^− 1. Since the reliability of dilution experiments is still frequently discussed in literature, we tested if our data obtained by dilution experiments reflected short-term in situ phytoplankton dynamics of the study site. We scaled experimental growth rates to water column irradiance, calculated short-term chlorophyll-a dynamics and compared the results to in situ measured chlorophyll-a concentrations. Calculated chlorophyll-a concentrations correlated significantly with in situ measured chlorophyll-a concentrations but slightly overestimated the in situ measured chlorophyll-a. This overestimation was in the range of phytoplankton assimilation reported for the Wadden Sea benthos. We will show that microzooplankton grazing had a large impact during the Phaeocystis bloom and during summer suggesting that a large proportion of phytoplankton biomass remained the pelagic food web. Microzooplankton grazing did not impact the diatom spring bloom and its demise.     

The Influence of the Cockle, Cerastoderma edule, on the Macrozoobenthic Community of Tidal Flats in the Wadden Sea

Abstract. The cockle Cerastoderma edule is one of the large and important species of the tidal flats of the Wadden Sea. C. edule disturbs the upper sediment layer due to its crawling and “shaking” behaviour. About 7% of a cockle population move within one week. The mean distance of movement is -4 cm per week, which, in the case of a 2-year-old cockle with a length of ?3 cm, means that ?20 cm² of sediment is disturbed to a depth of ?3 cm. Besides these measurable movements, cockles also regularly “shake” themselves, thereby disturbing the sediment around them to at least ?0.5 cm. At a density of 500 cockles per m² of a mean length of 3 cm, ?16% of the sediment surface is occupied by cockles and another ?29% is disturbed by cockles within one week. To study the effects of cockles on recruitment of other benthic species, densities of 0, 125, 250, 375 and 500 cockles (mean length 3 or 3.5 cm) were added to 1-m² plots within large depopulated squares in April. Subsequently, sediment samples were sieved once a month (from June to September) and the densities of all macrozoobenthic species were estimated. The presence of cockles significantly reduced the densities of juveniles of the bivalve species C. edule, Macoma balthica, Mya arenaria, Tellina (Angulus) tenuis, and Ensis directus and of the worm species Pygospio elegans, Lanice conchilega, Eteone longa, Anaitides spec., Nephtys hombergii, Heteromastus filiformis, Scoloplos armiger, and Tharyx marioni and of the amphipods Corophium volutator and C. arenarium. Non-significant effects were found only in Capitella capitata and Nereis diversicolor. For all species mentioned above a negative effect was already observed at the lower cockle densities of 125 - 250 per m². Reductions of about 50% or more were found at densities of 500 per m², which was more than could be expected based on the area occupied by the cockles (?16%), but close to the area occupied plus disturbed (?45%). It is concluded that (dense) assemblages of adult cockles can have a strong influence on the macrozoobenthic community.     

Exploring interactions among intertidal macrozoobenthos of the Dutch Wadden Sea using population growth models

Using data taken from three long-term monitoring programs, we modelled the population dynamics of 13 common macrozoobenthic species of the Balgzand, an approximately 50 km² intertidal area in the Dutch part of the Wadden Sea. In order to identify likely interactions among species, while accounting as much as possible for other factors influencing population dynamics, models included both ‘environmental’ (water temperature, and phytoplankton concentration) and ‘biological’ (biomass of potentially interacting species) variables. This approach appeared to be effective at identifying certain types of interactions acting over relatively short time-frames, such as predation and competition for food. Among the species we considered, the strongest effects of this kind appeared to be from what we presume to be predation by Nephtys hombergii, and from competition for food among bivalves. Several of the strongest results of our analysis corroborated expectations derived from previous small-scale surveys or experimental studies, but there were also a number of highly significant results indicating possible interactions which were not immediately explicable, including the apparent positive effect of Macoma balthica on Arenicola marina, and the apparent negative effect of Nephtys hombergii on Mya arenaria. Our results also suggest that year to year changes in Cerastoderma edule and Arenicola marina populations did not have substantial and widespread influences on the population dynamics of the majority of other common infauna, which is not to deny the likely critical importance of recruitment effects and biogenic transformation of the intertidal environment by these species over longer time frames. Our modelling approach could be applied to other long-term data sets, and will become more useful as longer time series become available.     

Overlapping distributions of cryptic Scoloplos cf. armiger species in the western Wadden Sea

Unknowingly lumping cryptic species in biological monitoring studies hinders progress in understanding their functioning in a wide range of research fields including population dynamics, ecophysiology and community ecology. The common polychaete worm Scoloplos cf. armiger is a cryptic species complex comprising entirely different developmental modes: holobenthic and pelago-benthic development. In the northeast Atlantic, three putative species have been described on the basis of molecular data and a breeding study. We report on the development of a fast genotyping assay and on the occurrence and distribution of different molecular types in the western Wadden Sea. The genotyping assay consists of PCR-RFLP analysis by two enzymes of a mitochondrial (cox3-trnQ-nad6) DNA segment. A new, fourth type was observed and this was the only type whose geographical distribution was not uniform but instead skewed toward the eastern part of the study area. All three hitherto known types were also observed and these three displayed a significant difference in depth distribution within the study area. This is the first ecological difference reported for the ‘Subtidal Clade’ (SC) versus the ‘Type Locality Clade’ (TLC). The new type (‘Intertidal Clade 2’) had a similar distribution to what is known as the Intertidal Clade (IC). However, the most striking observation is the large degree of overlap between the depth distributions in the western Wadden Sea, where the clade names ‘Intertidal’ and ‘Subtidal’ do not necessarily reflect local species composition. The necessity to genotype Scoloplos cf. armiger in monitoring programs is emphasized.     

A novel time-series station in the Wadden Sea (NW Germany): First results on continuous nutrient and methane measurements

In autumn 2002 a time-series station was installed in the tidal inlet between the Islands of Langeoog and Spiekeroog (Southern North Sea, NW Germany) to continuously measure physical, chemical, and meteorological parameters, even during extreme weather conditions (gale-force storms, drifting ice). Inside the pole of the station sensor tubes are installed in direction of the prevailing tidal currents. The tubes are equipped with hydrographic sensors (pressure, temperature, conductivity) and allow retrieval of water for nutrient analysis by automated instruments located inside the pole. Dissolved methane and the nutrients ammonia, nitrite, nitrate, phosphate, and silicate are measured at the station.     

Mussel fishery affects diet and reduces body condition of Eiders Somateria mollissima in the Wadden Sea

Although the Danish Wadden Sea is of international importance for several bird species, large-scale blue mussel Mytilus edulis fishing took place from 1984–1987, ceasing thereafter due to low mussel stocks. Mussel fishing removes much of the blue mussel biomass, especially larger individuals. Hence we predict that intensive mussel fishing will affect their predators, such as the Eider Somateria mollissima, which is predominantly a blue mussel feeder by, 1) reducing the amount of blue mussels in their diet relative to alternative prey items, 2) exploitation of smaller blue mussel shell classes, 3) loss of body condition, 4) changing feeding distribution to aggregate to the remaining mussel stocks, and 5) decreasing numbers. Before winter 1986/87 blue mussel biomass was estimated at 40,600 tons, decreasing to 15,400 tons in 1987/88 due to mussel fishery. We collected Eiders in both periods to monitor their diet and body mass and used aerial surveys to determine changes in numbers and distribution. Between the two periods, blue mussels declined in the Eiders diet, numbers of Eiders with empty stomachs increased and the mean length of blue mussel taken by Eiders decreased. Eider body condition declined from 1986/87 to 1987/88, mostly the result of the reduction in numbers of individuals with blue mussel remains in their gizzards and in better body condition compared to those taking alternative food items or having empty gizzards. Eiders shifted their distribution from the southern part of the Danish Wadden Sea to the northern part, where the remaining blue mussel stocks were situated. Eider numbers were lowest in 1987/88, the year of lowest blue mussel stocks. We conclude that intensive mussel fishery affected the Eider's diet, reduced their body condition and affected distribution and abundance. The results also showed that availability of blue mussels may have a key role in building up and maintaining body condition in Eiders during winter.     

Differential reproductive strategies of two bivalves in the Dutch Wadden Sea

Cerastoderma edule and Mya arenaria are two common bivalve species in European waters. Longevity and maximum size are much greater in the latter species. Because comparison of species life-history strategies states that a long life span (i.e. high annual survival) generally goes with lower fecundity, we hypothesise that reproductive output would be lower in M. arenaria than in C. edule. In the present paper, we studied the reproductive strategies of these two species in an intertidal and a subtidal area of the western Dutch Wadden Sea, by following seasonal changes in absolute and relative weights of somatic and gonadal tissues in these bivalves. Starting of spawning was similar in the two species, around May, except for intertidal M. arenaria, which initiated spawning in August. Individual energy investment in reproduction was similar for the two species but, unlike M. arenaria, C. edule spawned completely, releasing all energy of gonadal mass in the form of gametes. Mya arenaria used the gonad not only for reproduction but also for storage. In the intertidal area, we found a trade-off between longevity and reproduction, i.e. maximum reproductive output (expressed as a proportion of body mass) was higher in C. edule than in M. arenaria. However, since body size is larger and life span longer in M. arenaria than in C. edule, mean lifetime reproductive output per individual must be higher in the first than in the latter. Based on the differences in reproductive strategies of these two species, we hypothesise that the negative effects of warming climate on bivalve population dynamics in the Wadden Sea will be stronger in C. edule than in M. arenaria.     

Effects of suction-dredging for cockles on non-target fauna in the Wadden Sea

Suction dredging for cockles removes large cockles from tidal flats and may also cause mortality of non-target fauna and make the habitat less suitable for some species. This study examines whether suction dredging for cockles on tidal flats of the Dutch Wadden Sea had affected densities of non-target fauna, directly after fishing and one year later. Densities of non-target fauna in two randomly chosen undredged locations were compared to densities at the surrounding heavily commercially dredged area. A significant negative effect of cockle dredging on densities of 0-group Macoma balthica was observed and this effect persisted one year after dredging. The dredged area appeared to be less suitable for settlement of mussels Mytilus edulis. No significant effects of dredging on the mudsnail Hydrobia ulvae and on 0 and 1-group C. edule were found. For the mobile young Macoma balthica it seems unlikely that the effect found after one year was still due to the mortality caused by dredging and this suggests that the habitat was less suitable as a consequence of dredging. Thus, even in the highly dynamic ecosystem of the Wadden Sea, effects of bottom disturbance by cockle dredging may persist after one year.     

Nutrient dynamics in the Sylt-Rømø Bight ecosystem,German Wadden Sea: An ecological network analysis approach

Carbon, nitrogen, and phosphorus flow networks, consisting of 59 compartments, were constructed for the Sylt-Rømø Bight, a large shallow sea in the German Wadden Sea. These networks were analysed using ecological network analysis. Each network depicts the standing stock of each component in the ecosystem, and the flows between them. The trophic efficiency by which material is utilised in the Bight increase from 3%, to 6% to 17% for C, N and P, respectively. The number of cycles though which these elements pass increase from 1 197 for carbon, to 414?744 and 538?800 for nitrogen and phosphorus, respectively. The Finn Cycling Index, reflecting the amount of material recycled as a fraction of the total system activity, TST, increases from 17% for carbon, to 43% for nitrogen, to 81% for phosphorus. Other system level attributes such as the Average Path Length, the Average Internal Mutual Information, Relative Ascendancy, Relative and Normalized Redundancy, show an increase from the carbon to the nitrogen to the phosphorus networks. Phosphorus is tightly cycled over longer pathways than the other two elements, and also has the longest residence time in the Bight. Postulated differences between the behaviour of energy (or carbon) and biogeochemical networks in coastal ecosystems are evident from the results obtained from ecological network analysis.