Dutch Wadden Sea Research Articles

SATELLITE DERIVED BATHYMETRY FOR MONITORING NEARSHORE DYNAMICS

Research on the physical, ecological, and environmental processes in the coastal zone traditionally rely on the availability of accurate seafloor topography (bathymetry) information. This information becomes increasingly important as coastal environments are under pressure by climate change and anthropogenic developments. Nonetheless, the bathymetry of shallow nearshore waters is only marginally monitored by cost- and time-intensive survey techniques that are dependent on in-field environmental conditions. Here, Satellite Derived Bathymetry (SDB) offers a valuable alternative to enrich available bathymetric data in both data sparse and data rich environments, reducing the need for in-situ measurements in the future. The high spatial and temporal resolution of satellite imagery yields a more comprehensive understanding of our coast and its dynamics. Yet, consistently mapping temporal change is still challenging, as the majority of currently available empirical SDB algorithms are heavily dependent on in-situ data for calibration purposes. In this study we present a methodology for assessing nearshore horizontal morphodynamics using the uncalibrated reflectance signal from satellite sensors while focusing on the Friese Zeegat, a tidal inlet of the Dutch Wadden Sea.

Who lives in a pear tree under the sea? A first look at tree reefs as a complex natural biodegradable structure to enhance biodiversity in marine systems

Hard substrates play an important role in global marine systems as settlement surface for sessile reef-forming species such as corals, seaweeds, and shellfish. In soft-sediment systems, natural hard substrates such as stones, bedrock and driftwood are essential as they support diverse assemblages of reef-associated species. However, availability of these hard substrates has been declining in many estuaries and shallow seas worldwide due to human impacts. This is also the case in the Dutch Wadden Sea, where natural hard substrates have gradually disappeared due to burial by sand and/or active removal by humans. In addition, driftwood that was historically imported from rivers has been nullified by upstream logging and coastal damming of estuaries. To investigate the historic ecological role of wood presence in the Wadden Sea as settlement substrate and fish habitat, we constructed three meter high artificial reefs made of felled pear trees. Results demonstrate that these reefs rapidly developed into hotspots of biodiversity. Within six months, the tree-reefs were colonized by sessile hard substrate associated species, with a clear vertical zonation of the settled species. Macroalgae and barnacles were more abundant on the lower parts of the reef, while bryozoans were more dominant on the upper branches. In addition, six fish species were observed on the reefs, while only two species were caught on sandy control sites. Moreover, the abundance of fish on the reefs was five times higher. Individuals of the most commonly caught species, the five-bearded rockling Ciliata mustela, were also larger on the reef. These patterns also hold true for common prawn, Palaemon serratus, which were also larger and ten times more numerous on the reefs. Present findings indicate that the reintroduction of tree-reefs as biodegradable, structurally complex hard substrates can increase local marine biodiversity in soft-sediment systems within relatively short time scales.

Atmospherically Driven Seasonal and Interannual Variability in the Lagrangian Transport Time Scales of a Multiple‐Inlet Coastal System

AbstractIntense short‐term wind events can flush multiple‐inlet systems and even renew the water entirely. Nonetheless, little is known about the effect of wind variations at seasonal and interannual scales on the flushing of such systems. Here, we computed two Lagrangian transport time scales (LTTS), the residence and exposure times, for a multiple‐inlet system (the Dutch Wadden Sea) over 36 years using a realistic numerical model simulation. Our results reveal pronounced seasonal and interannual variability in both system‐wide LTTS. The seasonality of the LTTS is strongly anti‐correlated to the wind energy from the prevailing directions, which are from the southwesterly quadrant and coincidentally aligned with the geographical orientation of the system. This wind energy, which is stronger in autumn‐winter than in spring‐summer, triggers strong flushing (and hence low values of the LTTS) during autumn‐winter. The North Atlantic Oscillation (NAO) and the Scandinavia Pattern (SCAN) are shown to be the main drivers of interannual variability in the local wind and, ultimately, in both LTTS. However, this coupling is much more efficient during autumn‐winter when these patterns show larger values and variations. During these seasons, a positive NAO and a negative SCAN induce stronger winds in the prevailing directions, enhancing the flushing efficiency of the system. The opposite happens during positive SCAN and negative NAO, when weaker flushing during autumn‐winter is observed. Thus, large‐scale atmospheric patterns strongly affect the interannual variability in flushing and are potential drivers of the long‐term ecology and functioning of multiple‐inlet systems.

Comparing traditional vs. biodegradable seed mussel collectors (SMCs) for seed settlement, seed density, and seed growth: Effect of deployment depth and location

Mussel bottom culture is historically based on transplanting wild mussel seed to designated culture plots. Seed mussel collectors (SMCs) that are deployed in the water column are gradually replacing benthic mussel beds for mussel seed resource provisioning. Traditional SMCs consist of weighted filamentous nylon ropes. The performance of SMCs are promising, but the major disadvantages are the increased cost, effort, and the use of non-sustainable materials. In this study, we developed an innovative SMC: the BioShell-SMC. It consists of a coconut core rope surrounded by empty cockle shells that are held in place by biodegradable socking. The advantage of this system compared to traditional SMCs is that it provides biodegradable and sustainable resource material suitable for on-bottom placement. We compared its relative performance to that of a traditional SMC at different deployment depths and locations used for SMC deployment in the Dutch Wadden Sea and Oosterschelde. The results from this experiment indicated that in six out of nine locations mussel seed biomass was comparable between the two collector types. On both collector types, mussel seed biomass was higher in the Wadden Sea than in the Oosterschelde. We also found that mussel seed biomass development was not affected by deployment depth, though mussels were more numerous and shorter in deep water. The results of the current study provide a promising start toward a more sustainable mussel seed collection for bottom cultivation.

Using salt marshes for coastal protection: Effective but hard to get where needed most

Abstract Salt marshes fronting coastal structures, such as seawalls and dikes, may offer important ecosystem‐based coastal defence by reducing the wave loading and run‐up levels during storms. We question (i) how the long‐term salt marsh development in the Dutch Wadden Sea relates to the tidal‐flat foreshore bathymetry and (ii) how the wave run‐up onto dikes, which enhances the risk of dike failure, depends on foreshore bathymetry, the presence/absence of marshes, marsh vegetation properties, tidal range and wind exposure. We analysed 15 years of vegetation and bathymetry maps along the entire Dutch Wadden Sea coast, in combination with detailed process‐based measurements at five locations during 3 years, to understand where salt marshes naturally form and what features determine their contribution to coastal protection. The horizontal extent of marshes along the dikes remained relatively stable over the past decade. The presence of marshes was associated with higher elevations of adjacent tidal flats (above ~0.5 m NAP), while landward‐directed marsh retreat was associated with surface erosion of the fronting tidal flats. Wave run‐up during storms was lower at sites with wider marshes and higher foreshore elevations. This was attributed to the marsh attenuation effect, which led to a reduction in wave heights at the dike toe. As the tidal range varies across the Dutch Wadden Sea, areas to the East with generally higher water levels experienced higher wave run‐up. Synthesis and applications. We found that (i) marshes, where present, effectively protected the dikes from wave loading and (ii) the sites where marshes typically do not develop spontaneously were the most vulnerable to high wave run‐up. This catch‐22 problem implies that increasing reliance on nature‐based coastal defences along soft‐bottom coasts may require human interventions to stimulate marsh formation at the locations where it is most needed. Alternatively, ‘hard engineering’ solutions may remain necessary where implementing nature‐based solutions are either too costly, unachievable, or at the expense of other ecological values, such as causing the loss of mudflats that are important for migratory birds.

Spatial design improves efficiency and scalability of seed‐based seagrass restoration

Abstract Coastal ecosystem restoration is often ineffective and expensive in practice. As a consequence, upscaling restoration efforts to functionally relevant spatial scales remains one of the largest hurdles for coastal restoration practice. On small scales, restoration success of vegetated ecosystems (i.e. salt marshes and seagrasses) can be amplified by spatial designs that harness positive interactions. However, it remains unknown if positive interactions can be harnessed with seed‐based approaches, that are considered to be more cost‐effective and scalable than traditional shoot‐based restoration methods. Here, we investigated with a full‐factorial seeding experiment if (1) restoration scale (4, 40 and 400 m2) and (2) seeding density (10 and 50 injections/m2) affected multi‐year recruitment efficiency (measured as restored plants/seed injection) of annual eelgrass Zostera marina in the Dutch Wadden Sea. We found that the largest restoration scale (400 m2) increased second‐generation recruitment efficiency by suppressing a sedimentation‐related negative feedback. With increased restoration scale, the inner parts of the restoration plots captured less sediment, which decreased the desiccation stress of the restored eelgrass during low tide. Due to this stress alleviation, plants grew larger and produced more seed‐bearing spathes, which the following year resulted in two and three times higher recruitment efficiency at the largest restoration scale compared to the smaller scales. Moreover, lower seeding density more than doubled second‐generation recruitment efficiency compared to the higher density, supporting recent work showing that the effectiveness of ‘clumped’ spatial designs is context dependent. Synthesis and applications. The efficiency of restoration efforts is seldom taken into account, but can offer restoration projects a valuable metric with which workload, donor material and cost‐requirements can be reduced. We demonstrate that simple modifications to seed‐based coastal restoration designs (e.g. scale and density) can have a substantial impact on recruitment efficiency and multi‐year restoration yields. Thus, optimised restoration designs can strongly contribute to the upscaling potential of coastal ecosystem restoration. However, optimal restoration designs are expected to be strongly context dependent and we therefore argue that investigating optimal designs should be adopted as common practice, providing a crucial steppingstone between ‘proof‐of‐concepts’ and true large‐scale restoration attempts (km2).

Space–time analyses of sediment composition reveals synchronized dynamics at all intertidal flats in the Dutch Wadden Sea

Intertidal mudflat systems are shaped by geological processes and an interplay of hydrodynamics, sediment availability and ecological processes. All around the world these systems are affected by relative sea level rise (RSLR), changing climate and by human activities such as sediment nourishments, dredging, hydrological engineering and bottom trawling. These kinds of perturbations cause changes in morphology and sediment composition which may cause shifts in the composition, spatial distribution and productivity of benthic communities. We analysed the spatial and temporal variability of the sediment grain size of more than 900 km2 intertidal flats in the Dutch Wadden Sea in the period 2009–2019. The large scale coverage was achieved by yearly grid sampling at more than 4000 stations. Spatial panel data models were used to analyse changes in median grain size and mud content between years and to estimate the effects of resuspension due to wind and the accumulation of silt during summer. We show that between years the sediments of the intertidal flats changed synchronously throughout the study area and that the flats became coarser during the period 2011–2015 and muddier again between 2015 and 2019. The system wide changes and the absence of clear local deviations leads to the hypothesis that a large scale factor like changing hydrodynamic regime (e.g. due to RSLR), variability in the composition of suspended sediment in the North Sea or changing microphytobenthic productivity were causally involved in the coarsening of intertidal flats. Our data and analysis provides a base for further scientific enquiry but longer time series on higher temporal resolution of both sediment data and the physical and ecological environment are required. Models simulating the environment may provide further insight into possible development of sediment composition of the intertidal flats of the Wadden Sea.

Adaptation timescales of estuarine systems to human interventions

Many estuaries and tidal basins are strongly influenced by various human interventions (land reclamations, infrastructure development, channel deepening, dredging and disposal of sediments). Such interventions lead to a range of hydrodynamic and morphological responses (a changing channel depth, tidal amplitude and/or suspended sediment concentration). The response time of a system to interventions is determined by the processes driving this change, the size of the system, and the magnitude of the intervention. A quantitative understanding of the response time to an intervention therefore provides important insight into the processes driving the response. In this paper we develop and apply a methodology to estimate the response timescales of human interventions using available morphological and hydraulic data. Fitting an exponential decay function to data with sufficient temporal resolution yields an adaptation timescale (and equilibrium value) of the tidal range and deposited sediment volumes. The method has been applied in the Dutch Wadden Sea, where two large basins were reclaimed and where long-term and detailed bathymetric maps are available. Exponential fitting the morphological data revealed that closure of a very large part of a tidal basin in the Wadden Sea initially led to internal redistribution and import of coarse and fine sediments, and was followed by a phase of extensive redistribution while only fine-grained sediments are imported. Closure of a smaller part of a smaller basin led to shorter response timescales, and these response timescales are also more sensitive to rising mean sea levels or high waters. The method has also been applied to tidal water level observations in the Scheldt and Ems estuaries. Exponential fits to tidal data reveal that adaptation timescales are shortest at the landward limit of dredging. The adaptation time increases in the landward direction because of retrogressive erosion (Scheldt) or lowering of the hydraulic roughness (Ems). The seaward increase in adaptation time is related to the seaward widening of both systems.

Macrozoobenthos as an indicator of habitat suitability for intertidal seagrass

Seagrass meadows form the foundation of many coastal ecosystems, but are rapidly declining on a global scale. To conserve and restore these key-ecosystems, improved understanding of drivers behind seagrass presence and recovery is needed. Many animals are known to both facilitate and inhibit seagrasses, but biotic factors are still rarely used as indicators of seagrass presence. Hence, we investigate if macrozoobenthos could be used as an indicator for intertidal seagrass (Zostera marina and Zostera noltii) habitat suitability in the international Wadden Sea. Additionally, we explore if macrozoobenthos can explain the differing seagrass recovery rates that have been observed between the Northern (Denmark and Schleswig Holstein) and Southern (Lower Saxony and Netherlands) regions of the Wadden Sea. To achieve this, we performed a Wadden Sea-wide survey at 36 intertidal locations, across three countries, and investigated the importance of 21 abiotic and biotic variables in explaining the presence and absence of intertidal seagrasses. Seagrass presence or absence could be reliably predicted (prediction error: 16.7%) with a multivariate logistic regression with only four variables; chlorophyll a, bivalve, ragworm and mudsnail biomass. We also found higher chlorophyll concentrations and ragworm biomass in the South compared to the Northern Wadden Sea, suggesting that eutrophication and associated community shifts might still inhibit seagrass recovery in the South. Our findings highlight the potential of using macrozoobenthos as indicators for seagrass habitat suitability. In areas, like the Dutch Wadden Sea, where macrozoobenthic surveys are common and where benthic data is readily available, our findings can be used to improve the understanding of seagrass recovery dynamics and the selection of suitable seagrass restoration sites.

Biodegradable artificial reefs enhance food web complexity and biodiversity in an intertidal soft‐sediment ecosystem

Abstract Reef‐forming species form integral aspects of coastal ecosystems, but are rapidly degrading world‐wide. To mitigate these declines, nature managers increasingly rely on the restoration of habitat‐structuring, reef‐forming species by, for example, introducing artificial reefs that may directly function as complex reef habitat. Since the use of biodegradable structures to restore biogenic reefs is becoming a popular technique, its effectiveness as reef habitat must be assessed. Therefore, we examine the trophic complexity on experimental large‐scale biodegradable artificial reefs using food web network analysis. We placed biodegradable artificial reefs on soft‐sediment intertidal flats in the Dutch Wadden Sea in a large‐scale (~650 m) and 2.5‐year‐long experiment. We compared food web networks and biodiversity indicators between biodegradable reefs and bare controls and quantified species composition inside and near the artificial reef community to assess the expansion of the reef community. During 2.5 years, we observed that artificial reefs changed food web networks compared to bare controls: in species richness (+76%), link density (the number of interactions per species; +15%) and the fraction of basal species (species of lowest trophic level; +40%), but lowered the connectance: the realized fraction of all possible links between species (−33%). Their effects on food web networks increased over time with a higher species richness (+22%) and more complex food web (link density +13%) on the artificial reef 2.5 years after deployment compared to 1.5 years. However, the effects of the reefs did not extend beyond the reef structures; the species composition and biodiversity of macrozoobenthos near the reefs were comparable to the control. Synthesis and applications. This study shows that biodegradable artificial reefs offer an effective tool for the restoration of food web complexity and biodiversity of intertidal soft‐sediment systems. However, application needs to be carefully considered as the reef‐building species did not expand beyond our structures, despite the ambitious spatial extent of this experiment. Therefore, we recommend restoration practitioners to design artificial reefs in such a way that they generate ecosystem connectivity (facilitation of higher trophic levels) and biogeomorphological effects on a landscape scale (reef expansion beyond the structures).

Bonamia-free flat oyster (Ostrea edulis L.) seed for restoration projects: non-destructive screening of broodstock, hatchery production and test for Bonamia-tolerance

Native (flat) oyster (Ostrea edulis) beds, once a major component of the North Sea, largely disappeared from the region in the late 19th century. Flat oyster restoration is taking place at a number of locations in the North Sea. When flat oyster beds are restored in areas where O. edulis is functionally extinct it is advised to treat these as disease-free areas. Adult oysters were collected in the Dutch Delta area which is infected with the pathogenic parasite Bonamia ostreae. The aim of this research was to obtain Bonamia-free seed from parents collected in a Bonamia-infected area. In addition, the oysters were analysed to identify candidate genomic regions related to bonamiosis tolerance and exposed to Bonamia in the field to assess survival compared to a naïve control group. With the aid of a non-destructive screening method, Bonamia-free broodstock were selected. These oysters produced Bonamia-free larvae and seed. For comparison, broodstock oysters were collected in the Dutch Wadden Sea, an area free of Bonamia. These oysters also produced Bonamia-free larvae and seed. To study if the Delta area oysters had developed a degree of resistance to the disease, while the naïve Wadden Sea oysters had not, seed of both groups was challenged in Lake Grevelingen where Bonamia occurs. Survival of the pre-selected Bonamia-free oysters was significantly higher than the naïve group. Samples of seed were analysed for association of candidate genetic markers related to bonamiosis tolerance. A higher percentage of individuals with tolerance-associated marker genotypes was found in the screened group compared to the naïve one. However, mortality of the naïve group could not be related to Bonamia presence. Further challenge tests are needed before firm conclusions regarding the genetic markers can be made. The results show that hatchery production of Bonamia-free and potentially Bonamia-tolerant flat oysters is possible.

Optimizing seed injection as a seagrass restoration method

Due to the major declines of seagrasses worldwide, there is an urgent need for effective restoration methods and strategies. In the Dutch Wadden Sea, intertidal seagrass restoration has proven very challenging, despite numerous restoration trials with different restoration methods. Recently, however, the first field trial performed with a newly developed “dispenser injection seeding” method (DIS) resulted in record‐high plant densities and seed recruitment. Here, we present the further development of the methodology and consequently improved restoration results. During two consecutive growing seasons, we honed the seeding technique and experimentally investigated how seeding depth (2/4 cm), injection density (25/100 injects/m2), and seed amount (2/20 seeds/inject) affected restoration of intertidal annual Zostera marina. We found that all variables had a significant impact on plant establishment. Seeding deeper (4 cm) had the largest positive effect on restored plant densities, while lowered seed densities (2 seeds/inject) had the largest positive impact on seed recruitment. The optimized DIS method, combined with an altered placement of the seeding hole, resulted in a 50‐fold increase in restored plant densities (from approximately 1 to 57 plants/m2) and a simultaneous increase in seed recruitment (from 0.3 to 11.4%). These improvements stem from the method's ability to counteract a recruitment bottleneck, where seeds are lost through hydrodynamic forcing. The methodological improvements described here are important steps toward restoring self‐sustaining seagrass populations in the future and our study demonstrates the high potential of the seed‐based DIS method for seagrass restoration.

Coastal environmental and atmospheric data reduction in the Southern North Sea supporting ecological impact studies

Coastal climate impact studies make increasing use of multi-source and multi-dimensional atmospheric and environmental datasets to investigate relationships between climate signals and the ecological response. The large quantity of numerically simulated data may, however, include redundancy, multi-colinearity and excess information not relevant to the studied processes. In such cases techniques for feature extraction and identification of latent processes prove useful. Using dimensionality reduction techniques this research provides a statistical underpinning of variable selection to study the impacts of atmospheric processes on coastal chlorophyll-a concentrations, taking the Dutch Wadden Sea as case study. Dimension reduction techniques are applied to environmental data simulated by the Delft3D coastal water quality model, the HIRLAM numerical weather prediction model and the Euro-CORDEX climate modelling experiment. The dimension reduction techniques were selected for their ability to incorporate (1) spatial correlation via multi-way methods (2), temporal correlation through Dynamic Factor Analysis, and (3) functional variability using Functional Data Analysis. The data reduction potential and explanatory value of these methods are showcased and important atmospheric variables affecting the chlorophyll-a concentration are identified. Our results indicate room for dimensionality reduction in the atmospheric variables (2 principle components can explain the majority of variance instead of 7 variables), in the chlorophyll-a time series at different locations (two characteristic patterns can describe the 10 locations), and in the climate projection scenarios of solar radiation and air temperature variables (a single principle component function explains 77% of the variation for solar radiation and 57% of the variation for air temperature). It was also found that solar radiation followed by air temperature are the most important atmospheric variables related to coastal chlorophyll-a concentration, noting that regional differences exist, for instance the importance of air temperature is greater in the Eastern Dutch Wadden Sea at Dantziggat than in the Western Dutch Wadden Sea at Marsdiep Noord. Common trends and different regional system characteristics have also been identified through dynamic factor analysis between the deeper channels and the shallower intertidal zones, where the onset of spring blooms occurs earlier. The functional analysis of climate data showed clusters of atmospheric variables with similar functional features. Moreover, functional components of Euro-CORDEX climate scenarios have been identified for radiation and temperature variables, which provide information on the dominant mode (pattern) of variation and its uncertainties. The findings suggest that radiation and temperature projections of different Euro-CORDEX scenarios share similar characteristics and mainly differ in their amplitudes and seasonal patterns, offering opportunities to construct statistical models that do not assume independence between climate scenarios but instead borrow information (“borrow strength”) from the larger pool of climate scenarios. The presented results were used in follow up studies to construct a Bayesian stochastic generator to complement existing Euro-CORDEX climate change scenarios and to quantify climate change induced trends and uncertainties in phytoplankton spring bloom dynamics in the Dutch Wadden Sea.

Ragworms (Hediste diversicolor) limit eelgrass (Zostera marina) seedling settlement: Implications for seed-based restoration

Seagrasses are globally declining and multiple restoration efforts are undertaken to reverse these losses. However, these efforts have proven to be challenging, facing a variety of bottlenecks. We studied how predation by macroinvertebrates may form a potential bottleneck for seed-based seagrass restoration. Specifically, we questioned if the omnivorous common ragworm (Hediste diversicolor) may act as a predator on eelgrass (Zostera marina) seeds and whether that could affect seed-based eelgrass restoration trials. In a controlled lab experiment, we studied (1) how seedling establishment was affected by ragworm biomass (0, 2, 8 g DW m−2), (2) if the absence or presence of an additional or alternative high-protein food source (Sanikoi ® Gold Protein Plus, 52% protein) prevented potential seed predation by ragworms and (3) how ragworm size (small: 0.0029 g and 3.3× bigger: 0.0095 g DW ragworm−1) affected eelgrass seedling establishment. Additionally, we questioned (4) if ragworms may provide a bottleneck for annual eelgrass restoration experiments in the Dutch Wadden Sea by combining data from a large-scale benthic survey (SIBES, Netherlands Institute for Sea Research (NIOZ), Texel) with an existing eelgrass habitat suitability map. We found that >2 g DW m−2 ragworms completely hampered eelgrass seedling establishment, even when fed an additional, protein-rich, food source. Ragworms only seemed to target sprouted seeds rather than intact seeds. Additionally, sprouted seed consumption by ragworms was size-dependent: sprouted seeds escaped predation by smaller ragworms even when present in high biomass (2 g DW m−2). By extrapolating our findings to the field, we showed that 52.8% of the potential eelgrass growth sites in the Dutch Wadden Sea overlap with impeding ragworm biomass (≥2 g DW m−2). By consuming sprouted eelgrass seeds, ragworms may consequently strongly impede seed-based eelgrass restoration efforts, especially since both species have highly overlapping distributions. We thus provided novel insights into an unknown bottleneck for seed-based eelgrass establishment, which may have restoration implications. Especially for annual eelgrass that fully depends on successful seedling establishment for their persistence and survival.

Imminent reversal of the residual flow through the Marsdiep tidal inlet into the Dutch Wadden Sea based on multiyear ferry-borne acoustic Doppler current profiler (ADCP) observations

Abstract. The Dutch Wadden Sea is a UN World Heritage Site connected to the North Sea by multiple tidal inlets. Although there are strong tidal currents flowing through these inlets, the magnitude and direction of the residual circulation in the western Dutch Wadden Sea is important for sediment, salinity and nutrient balances. We found that the direction of this residual flow is reversing. This residual circulation has been the subject of various studies since the 1970s, in which substantially different net volume fluxes were presented. Differences in tidal conditions in the main inlets, tidal rectification and meteorology were identified as driving mechanisms. Here we analysed almost 13 years of acoustic Doppler current profiler (ADCP) observations collected on the ferry crossing the Marsdiep tidal inlet in the Dutch Wadden Sea since 2009. The results are combined with earlier investigations covering the period 1998–2009. We find a significant trend in the magnitude of the residual volume flux, with decreasing export to the North Sea and with occasional imports observed in recent years. We hypothesise that this trend is related predominantly to changes in tides in the North Sea, which are caused by increased strength and duration of stratification in response to global warming. With warming projected to continue, we expect the residual flow in the Marsdiep to continue to reverse to full inflow within the current decade, with potential knock-on effects for the sediment budget and ecosystem of the western Wadden Sea.

WATLAS: high-throughput and real-time tracking of many small birds in the Dutch Wadden Sea

Tracking animal movement is important for understanding how animals interact with their (changing) environment, and crucial for predicting and explaining how animals are affected by anthropogenic activities. The Wadden Sea is a UNESCO World Heritage Site and a region of global importance for millions of shorebirds. Due to climate change and anthropogenic activity, understanding and predicting movement and space-use in areas like the Wadden Sea is increasingly important. Monitoring and predicting animal movement, however, requires high-resolution tracking of many individuals. While high-resolution tracking has been made possible through GPS, trade-offs between tag weight and battery life limit its use to larger species. Here, we introduce WATLAS (the Wadden Sea deployment of the ATLAS tracking system) capable of monitoring the movements of hundreds of (small) birds simultaneously in the Dutch Wadden Sea. WATLAS employs an array of receiver stations that can detect and localize small, low-cost tags at fine spatial (metres) and temporal resolution (seconds). From 2017 to 2021, we tracked red knots, sanderlings, bar-tailed godwits, and common terns. We use parts of these data to give four use-cases revealing its performance and demonstrating how WATLAS can be used to study numerous aspects of animal behaviour, such as, space-use (both intra- and inter-specific), among-individual variation, and social networks across levels of organization: from individuals, to species, to populations, and even communities. After describing the WATLAS system, we first illustrate space-use of red knots across the study area and how the tidal environment affects their movement. Secondly, we show large among-individual differences in distances travelled per day, and thirdly illustrate how high-throughput WATLAS data allows calculating a proximity-based social network. Finally, we demonstrate that using WATLAS to monitor multiple species can reveal differential space use. For example, despite sanderlings and red knots roosting together, they foraged in different areas of the mudflats. The high-resolution tracking data collected by WATLAS offers many possibilities for research into the drivers of bird movement in the Wadden Sea. WATLAS could provide a tool for impact assessment, and thus aid nature conservation and management of the globally important Wadden Sea ecosystem.

Comparing taxonomic and functional trait diversity in marine macrozoobenthos along sediment texture gradients

The rapid reorganization of global biodiversity has triggered an intense research effort to understand its consequences for ecosystem functioning. However, efforts to monitor biodiversity change and evaluate the outcomes for ecosystem states and processes are currently poorly aligned. While most monitoring programs evaluate ecosystem status by reporting measures of taxonomic diversity, it is not the number of species but rather the exhibited traits of these species that regulate function. Trait-based approaches assume that trait diversity and variability relate to changes in functions across environmental gradients, but this relationship remains to be explored for most marine benthic ecosystems. Using macrozoobenthic communities from the Dutch Wadden Sea as a model, we compiled information on traits related to animal-sediment relationships. This trait information was then combined with species’ abundance data from a 19 years-long database to calculate different taxonomic and functional metrics that reflect macrozoobenthic diversity, function, and community structure. Finally, we compared how these taxonomic and functional metrics change along with sediment texture gradients. Our analyses showed that the structure of macrozoobenthic communities and various diversity metrics all changed with sediment gradients. The observed changes in the communities’ species composition were associated with directional shifts in the relative presence of specific functional traits with increasing sediment grain size, from communities dominated by small body size, deposit-feeding, and short life span to communities characterized by large to medium body size, suspension-feeding, and long life span. We observed limited functional redundancy and high sensitivity of functional trait-based measures to changes in the community composition along sediment gradients. Our findings suggest that a trait-based approach provides valuable information about the ecological function of marine macrozoobenthic species complementary to traditional biodiversity measures (e.g., species richness, Simpson diversity, etc.). Hence, these measures may be used to characterize changes in ecosystem functioning in time and space using traditional monitoring datasets.

Compound-specific stable isotope analysis of amino acid nitrogen reveals detrital support of microphytobenthos in the Dutch Wadden Sea benthic food web

The Wadden Sea is the world’s largest intertidal ecosystem and provides vital food resources for a large number of migratory bird and fish species during seasonal stopovers. Previous work using bulk stable isotope analysis of carbon found that microphytobenthos (MPB) was the dominant resource fueling the food web with particulate organic matter making up the remainder. However, this work was unable to account for the trophic structure of the food web or the considerable increase in δ15N values of bulk tissue throughout the benthic food web occurring in the Eastern regions of the Dutch Wadden Sea. Here, we combine compound-specific and bulk analytical stable isotope techniques to further resolve the trophic structure and resource use throughout the benthic food web in the Wadden Sea. Analysis of δ15N for trophic and source amino acids allowed for better identification of trophic relationships due to the integration of underlying variation in the nitrogen resources supporting the food web. Baseline-integrated trophic position estimates using glutamic acid (Glu) and phenylalanine (Phe) allow for disentanglement of baseline variations in underlying δ15N sources supporting the ecosystem and trophic shifts resulting from changes in ecological relationships. Through this application, we further confirmed the dominant ecosystem support by MPB-derived resources, although to a lesser extent than previously estimated. In addition to phytoplankton-derived particulate, organic matter and MPB supported from nutrients from the overlying water column there appears to be an additional resource supporting the benthic community. From the stable isotope mixing models, a subset of species appears to focus on MPB supported off recycled (porewater) N and/or detrital organic matter mainly driven by increased phenylalanine δ15N values. This additional resource within MPB may play a role in subsidizing the exceptional benthic productivity observed within the Wadden Sea ecosystem and reflect division in MPB support along green (herbivory) and brown (recycled/detrital) food web pathways.

Characterizing bedforms in shallow seas as an integrative predictor of seafloor stability and the occurrence of macrozoobenthic species

AbstractIn soft‐bottom marine ecosystems, bedform variation is induced by wind‐ and tidal‐driven hydrodynamics. The resulting megaripples, sand waves and sandbanks form a spatially and temporally heterogeneous seafloor landscape. The strong physical forces imposed by the migration of these bedforms are important determinants for the occurrence of different macrozoobenthic species. Quantifying the effect of these forces can help in differentiating natural‐ and anthropogenically induced physical stressors. However, large‐scale mapping of seabed morphology at high resolution using multibeam echosounder is challenging, costly and time‐consuming, especially in shallow seas, prohibiting wide swaths. Instead, their bathymetry is typically studied using single‐beam transects that are interpolated to bathymetric grids with a relatively coarse resolution (20 m). However, this leaves out information on smaller scale (<20 m) bedforms that can be ecologically relevant. In the Dutch Wadden sea, a shallow tidal system, we characterized bedform variation at high resolution using single‐beam data for the first time. We calculated a 2‐D Terrain Ruggedness Index (TRI) at sub‐meter resolution along the single‐beam transects and interpolated the results to a full 3‐D grid. We then validated the result by relating TRI to independently modeled hydrodynamic parameters and to the distribution of macrozoobenthic species. We found that TRI successfully integrates the variation of tidal‐driven bed shear stress and wave‐driven orbital velocity. In addition, we found TRI to be a good predictor of the occurrence of macrozoobenthic species. The inferred small‐scale bedforms provide valuable information for separating the relative importance of natural dynamics versus anthropogenic disturbances such as dredging and bottom trawling activities. We discuss that by repurposing already available single‐beam data in this way, bedforms can be characterized at high resolution without the need for additional equipment or mapping campaigns, yielding novel input to decision‐making on marine management and conservation.

Subtle ecosystem effects of microplastic exposure in marine mesocosms including fish

For two months, communities in 5.8 m3 outdoor marine mesocosms were exposed to 700 μm sphere-shaped polystyrene (PS) beads in dosages between 0.08 and 80 g/m2. Barnacle (Semibalanus balanoides) densities were reduced at dosages of 0.8 g/m2 onwards without following a standard dose response curve. Lugworms and fish (Solea solea) ingested PS-beads without accumulating them. Lugworms (Arenicola marina) ingested the beads nonselective with the sediment without negative effects. The fish seemed to ingest the plastics only occasionally and at the final sampling day even in the highest dosed mesocosms (>30 beads/cm2) only 20% contained plastic. The condition index of the fish was slightly reduced in mesocosms with dosages of 0.8 g/m2 onwards. No difference in condition was found between fish with and without ingested plastic across mesocosms, illustrating the difficulty to relate plastic ingestion with condition from field data. The fish also ingested mollusks with shells exceeding the size of the PS-beads. Bivalves rejected the PS-beads as pseudofeces, without obvious impact on their condition. Mussel's (Mytilus edulis) pseudofeces present an effective matrix to monitor microplastic presence in the water column. Species richness and diversity of the pelagic and benthic community were not affected although, a trend was found that the lower microplastic dosages had a positive effect on the total abundance of benthic invertebrates. In general, the observed effects at even the highest exposure concentrations were that subtle that they will be obscured by natural variation in the field. This underlines the importance of experiments under semi-field conditions for meaningful assessment of the ecological impact of microplastics. This study was performed with the real life, non-toxic, sphere-shaped polystyrene beads as were lost during an actual spill near the Dutch Wadden sea in January 2019. We recommend future mesocosm studies with other types of microplastics, including microfibers, weathered microplastics from sea, and smaller sized particles down to nanoplastics.