Erodibility of a mixed mudflat dominated by microphytobenthos and Cerastoderma edule, East Frisian Wadden Sea,Germany

Sediment erodibility and a range of physical and biological parameters were measured at an intertidal site in the German Wadden Sea area in June, September and November 2002 and February and April 2003 in order to examine the influence of macrozoobenthos and microphytobenthos on sediment erodibility and the temporal variation. The study site was a mixed mudflat situated in the mesotidal Baltrum–Langeoog tidal basin at the East Frisian barrier coast. The mud content at the site was about 35% and the filter-feeding cockle Cerastoderma edule was the dominating macrozoobenthic species (by biomass). The erodibility of the sediment showed strong temporal variation with high erosion thresholds in spring and late summer and significantly lower thresholds during the rest of the study period. The erosion thresholds were strongly dependent on the contents of chlorophyll a (chl a) and colloidal carbohydrates, both indicators of the content of microphytobenthos, in this environment primarily benthic diatoms. The content of microphytobenthos was high in September 2002 and April 2003, and regression analysis indicated that this was the only likely reason for the low erodibility found at these times. A biostabilisation index of about 4.5 was found for a situation with both abundant biofilms and cockles.     

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.     

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.     

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.     

Metazoan parasites in intertidal cockles Cerastoderma edule from the northern Wadden Sea

At four intertidal sites near the island of Sylt (eastern North Sea), 13 metazoan parasite taxa were found in 1400 cockles investigated, with digenean trematodes being dominant. Almost all cockles were infected by parasites and most individuals harboured more than one parasite species. We observed four conspicuous patterns: (1) Adult cockles harboured a two times higher species richness (2003: 6.1 ± 0.7 species/host; 2004: 7.1 ± 0.7) than juveniles (2003: 2.9 ± 0.8; 2004: 3.4 ± 0.8) and total parasite community composition significantly differed between age groups. (2) Infection levels were 2–52 times higher in adult cockles than in juveniles both in trematode species and in non-trematode species. In the dominant trematodes, species utilising cockles as first intermediate host (Gymnophallus choledochus, Labratrema minimus, Monorchis parvus) only occurred in adult cockles, and prevalences were low (2–12%). Prevalences of up to 100% were reached by trematodes using cockles as second intermediate host (Himasthla elongata, H. continua, H. interrupta, Renicola roscovita, Psilostomum brevicolle, Meiogymnophallus minutus, Gymnophallus gibberosus). Metacercariae of these species were segregated between body parts within cockles. (3) High spatial heterogeneity in parasite community composition and infection levels occurred between sampling sites. However, communities in juveniles were more similar than communities in adults. (4) Temporal variation in parasite community composition was low between two consecutive years, especially for adult cockles. The omnipresence of parasites in this dominant bivalve species has important implications for sampling designs and as a potentially confounding variable in e.g. physiological studies. It suggests strong and cumulative negative effects on the cockle hosts.     

Effects of current exposure on habitat preference of mobile 0-group epibenthos for intertidal seagrass beds (Zostera noltii) in the northern Wadden Sea

In the northern Wadden Sea, the extent of intertidal seagrass beds, their plant biomass and shoot density highly depends on local current regimes. This study deals with the role of intertidal Zostera noltii beds as nursery for mobile epibenthic macrofauna and the impact of seagrass bed characteristics on their abundance and distribution patterns. According to their exposure to the main tidal gullies, sampling sites were separated into exposed, semi-exposed and sheltered. Dominant species of crustaceans and demersal fish were studied in respect of their abundances within seagrass beds and adjacent unvegetated areas. Quantitative sampling was performed at day and night high tide using a portable drop trap. In general, species composition varied little between seagrass beds and bare sand. However, the presence of vegetation had a quantitative effect increasing individual numbers of common epifaunal species. Abundances of 0-group shore crabs (Carcinus maenas), common gobies (Pomatoschistus microps) and brown shrimps (Crangon crangon) were highest within sheltered seagrass beds. With decreasing plant density habitat preference of epibenthos changed on species level. By regulating the habitat complexity the currents regime is profoundly influencing the nursery function of intertidal seagrass beds in the Wadden Sea.     

Factors involved in growth plasticity of cockles Cerastoderma edule (L.), identified by field survey and transplant experiments

In Arcachon Bay, a macrotidal coastal lagoon in southwest France, growth rates of cockles Cerastoderma edule (L.) differ greatly between locations. Sampling of populations at different tidal levels showed that the mean shell length was significantly and positively correlated with immersion time, whereas no correlation was found with population density and microphytobenthos biomass. Transplants of cockles between two intertidal sites were used to examine the relative importance of habitat and site of origin for growth rate and condition index. Artifacts due to manipulation were assessed, i.e. the impact of enclosures on growth, conditon index and mortality. During a 5-mo reciprocal transplant experiment, growth rates of the transplanted cockles and the cockles already present were similar, whereas the sites of origin did not affect growth rate. Condition index, however, displayed significant differences in relation to both transplant and origin sites. During a further 6-mo experiment in which cockles were transplanted from one site to four new sites, growth was mainly influenced by tidal level. These results indicate that net growth started when cockles were immersed 30% of the time. It is concluded that, on the scale of Arcachon Bay (156 km²), differences in growth and condition between cockle populations are largely phenotypic. Tidal level (i.e. food supply and emersion stress) accounts for most of these differences, but the existence of genetically or physiologically different populations cannot be excluded.     

Observations of a mass occurrence of Macoma balthica larvae in midsummer

In 1995 the seasonal development of concentrations of both phytoplankton and larvae of the bivalve Macoma balthica was studied in the coastal zone behind the back-barrier island of Spiekeroog (German Wadden Sea). In July/August larvae reached maximum concentrations of about 1000 to 4200 ind. m⁻³ (depending on the sampling site and tidal period), probably in relation to a phytoplankton bloom in July. This observation of an unusually late maximum of Macoma larvae is discussed in detail in connection with the recent literature available about spawning of M. balthica in the Wadden Sea.     

Reduced survivorship of Himasthla (Trematoda, Digenea)-infected cockles (Cerastoderma edule) exposed to oxygen depletion

Bivalve populations from inshore waters often accommodate a diverse trematode fauna that may have a variety of effects on host specimens. In particular, larval trematodes that grow or reproduce within their host are known to be severe pathogens, whereas trematodes utilising bivalves only for encystment are thought to be relatively benign. Yet this may depend on the environmental conditions, and it can be expected that such trematodes in concert with other stress agents can be detrimental to host organisms. To examine the impact of such larval trematodes on hosts subjected to stress, we studied the digenetic trematode Himasthla elongata and one of its second intermediate hosts, the bivalve Cerastoderma edule. Experimentally infected cockles and non-infected cockles were exposed to oxygen depletion, whereupon we measured their burrowing ability and survivorship. After 30 h of hypoxia, the survival of infected cockles was significantly reduced compared to non-infected cockles, whereas no effect of parasites on cockles under normoxic conditions was found. In addition, parasites tended to reduce the burrowing ability of cockles exposed to hypoxia but the effect was not clear. The effect of parasites and possible ecological consequences are discussed and it is suggested that the combined effects of parasites and oxygen deficiency may explain some hitherto unexplained cases of mass mortalities in bivalve populations.     

Influence of chronic exposure to silver and mercury in the field on the bioaccumulation potential of the bivalve Macoma balthica

In order to assess the adaptation to metals previously observed in the bioindicator organism, Macoma balthica, subjected to chronic contamination by silver and mercury in the French Loire estuary, the bioaccumulation potential of individual organisms originating from the contaminated Loire estuary and a relatively uncontaminated control estuary (Somme) was evaluated using both radiotracers and stable isotopes of Ag (80 μg Ag litre⁻¹) and Hg (100 μg Hg litre⁻¹). Clams from the contaminated estuary were more sensitive to Ag (LT₅₀ = 9d) than those originating from the Somme estuary (LT₅₀ > 15d), even though the former bioaccumulated Ag to a significantly lower degree. This is attributed to a consequence of the chronic stress induced by Ag while clams were living in their natural environment. Therefore, past history of trace metal contamination should be considered when evaluating the susceptibility of M. balthica to heavy metal exposure. Lower uptake rates obtained for Hg (during the initial uptake phase only) and for Ag in clams from the polluted estuary suggest the presence of an adaptive trait for survival in contaminated areas. However, the lower degree of bioconcentration observed for Ag was not sufficiently low to reduce the sensitivity of the organisms to Ag and allow them to resist the toxic stress. Clams that survived Ag or Hg exposure at LT₅₀ did not protect themselves against metal toxicity by accumulating a significantly lesser amount of these metals than clams which did not survive metal stress. The results suggest that the bioaccumulation potential of each individual was not a factor which can explain the survival ability of M. balthica exposed to chronic Ag and Hg contamination in estuaries. In this case, cellular, biochemical and genetic levels of adaptation are presumed to be of greater importance.     

Contribution of different generations of the brown shrimp Crangon crangon (L.) in the Dutch Wadden Sea to commercial fisheries: A dynamic energy budget approach

In this paper a contribution is made to the ongoing debate on which brown shrimp generation mostly sustains the autumn peak in coastal North Sea commercial fisheries: the generation born in summer, or the winter one. Since the two perspectives are based on different considerations on the growth timeframe from settlement till commercial size, the Dynamic Energy Budget (DEB) theory was applied to predict maximum possible growth under natural conditions. First, the parameters of the standard DEB model for Crangon crangon L. were estimated using available data sets. These were insufficient to allow a direct estimation, requiring a special protocol to achieve consistency between parameters. Next, the DEB model was validated by comparing simulations with published experimental data on shrimp growth in relation to water temperatures. Finally, the DEB model was applied to simulate growth under optimal food conditions using the prevailing water temperature conditions in the Wadden Sea. Results show clear differences between males and females whereby the fastest growth rates were observed in females. DEB model simulations of maximum growth in the Wadden Sea suggest that it is not the summer brood from the current year as Boddeke claimed, nor the previous winter generation as Kuipers and Dapper suggested, but more likely the summer generation from the previous year which contributes to the bulk of the fisheries recruits in autumn.     

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.     

Effect of macrofaunal bioturbation on bacterial distribution in marine sandy sediments, with special reference to sulphur-oxidising bacteria

We have studied the impact of the bioturbating macrofauna, in particular the lugworm Arenicola marina and the bivalve Cerastoderma edule, on abundances and distribution patterns of total bacteria and of bacteria of selected functional groups in sandy intertidal sediments. The selected groups comprised the colourless sulphur-oxidising bacteria and the anoxygenic phototrophic bacteria, which are expected to occupy small zones at the oxygen–sulphide interface in stable (non-bioturbated) sediments. The presence of a wooden wreck buried in the sediment at 10 cm depth within a large area of intertidal sand flat colonised by lugworms provided a unique opportunity to confront field observations with laboratory simulations. The site with the wooden wreck, which was used as control site, was devoid of both A. marina and C. edule, while the composition of the rest of the zoobenthic community was rather similar to that of the surrounding area. In the field, the density of total bacteria was approximately one order of magnitude higher in the control site than in the natural (bioturbated) site. This can be explained by the higher contents of silt and clay particles (higher surface-area/volume ratio) and higher total organic-carbon contents found at the control site. It appears that the presence of macrofauna affects sedimentation processes, which indirectly influence bacterial dynamics. Samples from the control site have been incubated in the laboratory with A. marina and C. edule added (bioturbated core), while an unamended core served as a control. The laboratory experiments contrasted with the field observations, because it was found that total bacteria were actually higher in the deeper layers of the bioturbated core. Moreover, the populations were more homogeneous (less stratified) and colourless sulphur bacteria were on average less numerous in the bioturbated core. In general, laboratory incubations resulted in a decrease of total bacteria with a concomitant increase of colourless and phototrophic sulphur-oxidising bacteria and thus in modifications of the bacterial community structure. Hence, our results demonstrate that care must be taken in extrapolating results from laboratory experiments (e.g. mesocosm research) to field situations.     

Rapid invasion of Crassostrea gigas into the German Wadden Sea dominated by larval supply

Invasions of non-indigenous species into coastal habitats have been a frequent phenomenon in the last decades, sometimes with significant impact on the receiving ecosystem. However, the understanding of the entire process and especially the relative importance of larval supply and local recruitment remains unclear. In this study, we simulate the invasion of a benthic invertebrate into a previously uncolonised habitat over several years and validate the results with field data. Therefore, we present field data from a monitoring programme revealing the rapid invasion of the oyster Crassostrea gigas into the East Frisian Wadden Sea, North Sea, between 2003 and 2005. The applied model combines a simple, spatially-explicit population dynamics model for the adult stage with a particle tracking model for the larval stage of the life cycle. Simulation results are able to reproduce the large-scale pattern of the field data and indicate a domination of larval supply on the population dynamics in the early stage of the invasion. Though monitoring and simulations suggest a single larval source outside the study area in the west, the population dynamics in the eastern part is only explainable with an additional source within the study area attributed to an unintentional input of juveniles by mussel fishery. High sensitivities to uncertain parameters result in distinct deviations between monitoring and simulations at particular sites. Especially the impact of site-specific variations of the post-settlement mortality underlines the variability of local recruitment conditions and indicates the need for spatially resolved information for exact predictions.     

The Effect of Macrofauna on Porewater Profiles and Nutrient Fluxes in the Intertidal Zone of the Humber Estuary

Macrofauna, nutrient fluxes, porewater chemistry and sediment characteristics were measured at six intertidal mudflat sites in the Humber Estuary, U.K., during the different seasons. Nereis diversicolor , Macoma balthica and Corophium volutator were found to be the dominant macrofauna. Salinity was the baseline control on macrofauna distribution but this was overprinted by periodic impoverishment due to sediment mobilization. High resolution gel probe porewater samplers provided direct evidence for the impact of burrows on porewater chemistry. The macrofauna modified nutrient fluxes during periods of mud flat stability. Nereis caused a decrease in silicate and phosphate effluxes but enhanced ammonia release and nitrate uptake. Macoma enhanced ammonia and nitrite release. The impact of Corophium was not possible to discern. The Humber is a large, highly dynamic macrotidal estuary in which sediment resuspension has a large impact on porewater profiles, nutrient fluxes and macrofaunal communities. Simple patterns and inter-relationships which are seen in small sheltered estuaries are not observed in the Humber.