Dynamics of Phaeocystis pouchetii blooms in the Wadden Sea of Sylt (German Bight,North Sea)

The colony-forming alga Phaeocystis pouchetii (Haptophyceae) forms regular blooms in spring and early summer in the German Wadden Sea. This paper gives a summary of observations of Phaeocystis blooms in 1975, 1976, and 1981 in the Wadden Sea of Sylt (German Bight, North Sea).The onset of Phaeocystis pouchetii blooms started after the decline of diatoms at low inorganic nutrient concentrations, when particularly inorganic phosphate had dropped to minimal concentrations. Phaeocystis blooms did not start unless water temperatures were above 7°C in the investigation area, and the admixture of organic substances seemed to be an important factor in controlling the onset and the size of Phaeocystis blooms.As diatom populations increased again at the end of the Phaeocystis blooms, a causal relationship between the successive development of both algal groups, possibly in combination with other abiotic factors, cannot be excluded.     

Predatory Promesostoma species (Plathelminthes, Rhabdocoela) in the Wadden Sea

Laboratory experiments were performed on the food ecology of four congeneric species of free-living plathelminths, Promesostoma caligulatum, P. marmoratum, P. rostratum, and P. meixneri, all inhabiting an intertidal sandflat near the island of Sylt (North Sea). Their prey spectrum is within the microcrustaceans: P. caligulatum preferred ostracods, while the other three species favoured copepods, with species-specific differences for copepod species and size classes. Daily consumption of prey species varied with the size of both the predator and the prey. On average, P. marmoratum consumed 0.76 Harpacticus flexus per day while this rate decreased to 0.06 in P. meixneri, the smallest predator. When these Promesostoma species were fed with Tachidius discipes, a smaller prey species, their predation rates were about 25% higher. While the larger predators preferred the larger harpacticoids as prey, the small P. meixneri preferred small cyclopoids over larger harpacticoids. In terms of biomass, P. marmoratum's mean consumption of T. discipes per day was about half the predator's own weight. This average varied with prey density and temperature. A comparison of these consumption rates with the field densities of the predators and their prey shows that the plathelminth predators may consume as much as 10% per day of their copepod prey populations, thus strongly influencing these prey populations on these sandflats. The predation pressure of P. caligulatum on ostracods was about 1% per day of the prey population. Since ostracods usually have fewer generations per year, the total effect on the population dynamics may be similar to that on copepods. Therefore, nocturnal swimming of copepods in the water column may be interpreted as an attempt to escape plathelminth predators.     

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.     

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.     

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.     

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.     

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.     

Functional structure of laminated microbial sediments from a supratidal sandy beach of the German Wadden Sea (St. Peter-Ording)

Hidden for the untrained eye through a thin layer of sand, laminated microbial sediments occur in supratidal beaches along the North Sea coast. The inhabiting microbial communities organize themselves in response to vertical gradients of light, oxygen or sulfur compounds. We performed a fine-scale investigation on the vertical zonation of the microbial communities using a lipid biomarker approach, and assessed the biogeochemical processes using a combination of microsensor measurements and a ¹³C-labeling experiment. Lipid biomarker fingerprinting showed the overarching importance of cyanobacteria and diatoms in these systems, and heterocyst glycolipids revealed the presence of diazotrophic cyanobacteria even in 9 to 20 mm depth. High abundance of ornithine lipids (OL) throughout the system may derive from sulfate reducing bacteria, while a characteristic OL profile between 5 and 8 mm may indicate presence of purple non-sulfur bacteria. The fate of ¹³C-labeled bicarbonate was followed by experimentally investigating the uptake into microbial lipids, revealing an overarching importance of cyanobacteria for carbon fixation. However, in deeper layers, uptake into purple sulfur bacteria was evident, and a close microbial coupling could be shown by uptake of label into lipids of sulfate reducing bacteria in the deepest layer. Microsensor measurements in sediment cores collected at a later time point revealed the same general pattern as the biomarker analysis and the labeling experiments. Oxygen and pH-microsensor profiles showed active photosynthesis in the top layer. The sulfide that diffuses from deeper down and decreases just below the layer of active oxygenic photosynthesis indicates the presence of sulfur bacteria, like anoxygenic phototrophs that use sulfide instead of water for photosynthesis.     

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.     

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.     

Reproductive investment of the American razor clam Ensis americanus in the Dutch Wadden Sea

The American razor clam Ensis americanus is a marine bivalve indigenous from the American coast that was introduced in Europe in the late 1970s and has managed to build up strong populations along the European coast. In the present paper, we studied energy investment into soma and gonads by following the seasonality in growth and reproduction of this species in a subtidal area of the western Dutch Wadden Sea. Body and somatic mass indices showed a seasonal pattern with maximal values in August and minimal in April. Gonadal mass index showed the opposite trend with highest values in April. Spawning started in April/May and a second, but weaker, spawning event apparently occurred in August/September. Gonadosomatic ratio values were very low. Just before the spawning period, only 2.5% of the body mass of E. americanus consisted of gonads.     

Pseudo-cryptic species Arenicola defodiens and Arenicola marina (Polychaeta: Arenicolidae) in Wadden Sea,North Sea and Skagerrak: Morphological and molecular variation

The polychaete Arenicola defodiens, or black lug, was recently described as a morphologically highly similar species alongside the blow lug Arenicola marina. So far it was only known from the British Isles. A double spawning peak was observed earlier in lugworms of the western Wadden Sea. Here, we test the hypothesis that the two spawning peaks represent the two species in sympatry. This hypothesis is refuted on the basis of both morphological and mitochondrial DNA data; both spawning peaks are attributed to A. marina. In spite of this, we confirm, on the basis of new collections as well as re-examination of museum collections, the presence of A. defodiens in the western Wadden Sea, North Sea and also the Skagerrak; its distribution is restricted to habitats which are either submerged or have short emersion times, have relatively coarse sediments and high and stable salinities. Sympatry is common. The species differ strongly in the mitochondrial DNA fragment examined; the observed 14% uncorrected minimum difference amounts to an estimated 33–63 million years since sequence divergence. The amount of intraspecific molecular variation is larger for A. defodiens than for A. marina. This is evidence to suggest that A. marina may have undergone more recent and/or more severe population size bottlenecks.     

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.     

Net dispersal of harbour seals within the Wadden Sea before and after the 1988 epizootic

Harbour seals in the Wadden Sea (The Netherlands, Germany and Denmark) have been monitored by aerial surveys since the 1980s. Annual maximum figures reported in the various regions (38 separate tidal basins) and the number of pups counted were used to estimate vital population parameters and to quantify the net migration of harbour seals within the Wadden Sea before (1980–1987) and after an epizootic (1990–1994). The total pre-whelping population increased at a mean annual rate of 9 and 14%, respectively, and varied considerably among the various regions of the Wadden Sea. The overall mean annual survival was found to increase from 0.88 during the pre-epizootic period to 0.93 during the post-epizootic period. The distribution of pups and of older seals over the 38 separate sub-areas was highly uneven. Based on between-year population changes and the number of pups reported, net dispersal fluxes among the four regions of the Wadden Sea were estimated for the two time intervals. During the pre-epizootic period, only Schleswig-Holstein, Germany, produced a surplus of animals which counter-balanced the low recruitment of the other regions. During the post-epizootic period, the net dispersal fluxes were at a much lower level. Nevertheless, in The Netherlands, a continuous influx of seals was required because the recruitment was insufficient to explain the observed numerical increase. At the level of the 38 separate tidal basins, the actual pattern of dispersal showed that the majority of sub-areas were not self-supporting with respect to the local pup production and thus dependent on the influx of animals. More than 65% of all immigrants originated from only 7 sub-areas, which are considered `key areas' of vital importance for the Wadden Sea harbour seal population.     

Sources and fate of manganese in a tidal basin of the German Wadden Sea

Dissolved and particulate Mn concentrations were investigated on a seasonal scale in surface waters of the NW German Wadden Sea (Spiekeroog Island) in 2002 and 2003. As the Wadden Sea forms the transition zone between the terrestrial and marine realms, Mn was analysed in coastal freshwater tributaries and in the adjoining German Bight as well. Additionally, sediments and porewaters of the tidal flat sediments were analysed for Mn partitioning and microbial activity.Dissolved Mn concentrations show strong tidal and seasonal variation with elevated concentrations during summer at low tide. Summer values in the Wadden Sea (av. 0.7 μM) are distinctly higher than in the central areas of the German Bight (av. 0.02 μM), suggesting a possible impact of the Wadden Sea environment on the Mn budget of the North Sea. Seasonality is also observed for particulate Mn in the Wadden Sea (winter av. 800 mg kg^− 1; summer av. 1360 mg kg^− 1). Although particles are relatively Mn-poor during winter, the high SPM load during this season causes elevated excess concentrations of particulate Mn, which in part exceed those of the dissolved phase. Therefore, winter values cannot be ignored in balance calculations for the Wadden Sea system.Porewater Mn concentrations differ depending on sediment type and season. Maximum concentrations are found in surface sediments at a mixed flat site (190 μM) during summer, while winter values are distinctly lower. This indicates that enhanced microbial activity owing to higher temperature during summer leads to increased reduction of Mn-oxides in surface sediments and enhances the corresponding diffusive and advective Mn flux across the sediment-water interface. Draining of Mn-rich porewaters from sediments is also documented by analyses of tidal creek waters, which are highly enriched in Mn during summer.Furthermore, an important Mn source is freshwater discharged into the Wadden Sea via a flood-gate. The concentration of dissolved Mn in freshwater was highly variable during the sampling campaigns in 2002 and 2003, averaging 4 μM. In contrast, particulate Mn displayed a seasonal behaviour with increasing contents during summer. On the basis of salinity variations in the Wadden Sea, the total amount of Mn contributed to the Wadden Sea via freshwater was estimated. This balance shows the importance of the freshwater environment for the Mn inventory of the Wadden Sea. During winter the total Mn inventory of the Wadden Sea water column may be explained almost completely by freshwater discharge, whereas in summer the porewater system forms the dominating source.