Wadden Sea Research Articles

Vessel noise exposures of harbour seals from the Wadden Sea

The North Sea faces intense ship traffic owing to increasing human activities at sea. As harbour seals (Phoca vitulina) are abundant top predators in the North Sea, it is hypothesised that they experience repeated, high-amplitude vessel exposures. Here, we test this hypothesis by quantifying vessel noise exposures from deployments of long-term sound and movement tags (DTAGs) on nine harbour seals from the Wadden Sea. An automated tool was developed to detect intervals of elevated noise in the sound recordings. An assessment by multiple raters was performed to classify the source as either vessels or other sounds. A total of 133 vessel passes were identified with received levels > 97 dB re 1µPa RMS in the 2 kHz decidecade band and with ambient noise > 6 dB below this detection threshold. Tagged seals spent most of their time within Marine Protected Areas (89 ± 13%, mean ± SD) and were exposed to high-amplitude vessel passes 4.3 ± 1.6 times per day. Only 32% of vessel passes were plausibly associated with an AIS-registered vessel. We conclude that seals in industrialized waters are exposed repeatedly to vessel noise, even in areas designated as protected, and that exposures are poorly predicted by AIS data.

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).

Mass mortality among colony-breeding seabirds in the German Wadden Sea in 2022 due to distinct genotypes of HPAIV H5N1 clade 2.3.4.4b.

Mass mortality was observed among colony-breeding seabirds in the German Wadden Sea area of the North Sea during the summer months of 2022. Several species' colonies were affected, most notably sandwich terns (Thalasseus sandvicensis), common terns (Sterna hirundo) and Germany's only northern gannet (Morus bassanus) colony on the island of Heligoland. Mortality in some tern colonies reached 40%, while other colonies were almost spared. In all cases, infections with the high-pathogenicity avian influenza virus (HPAIV) subtype H5N1 of clade 2.3.4.4b were identified to have caused the epidemic. Phylogenetic analysis of whole-genome sequences revealed that the outbreaks were dominated by two genotypes, Ger-10-21 N1.2 and Ger-10-21 N1.5, previously identified in Germany. Spatiotemporal analyses of phylogenetic data suggested that these viruses could have entered the continental North Sea coastal region via the British Isles. A close linkage of viruses from tern colonies in the German Wadden Sea was evident with further connections to breeding colonies in Belgium and the Netherlands, and further spread to Denmark and Poland. Several of the affected species are endangered, such that negative effects of epizootic HPAIV infections on populations are feared, with uncertain long-term consequences.

Impact of mussel seed fishery on subtidal sediment and macrozoobenthos in the western Wadden Sea

Impact of mussel seed fishery on subtidal sediment and macrozoobenthos in the western Wadden Sea

Prey choice and ingestion of microplastics by common shelducks and common eiders in the Wadden Sea World Heritage Site

Top predators such as coastal birds are essential components of marine food webs, and understanding their trophic interactions forms an essential basis of food web models. At the same time, the proportion of plastic debris in marine food webs has constantly increased while the degree of plastic ingestion by marine birds is still poorly known. In this study, the diets and microplastic uptakes in two numerous benthivorous bird species in the Wadden Sea were examined microscopically, i.e. the common eider (Sommateria mollissima) indicative for the subtidal and the common shelduck (Tadorna tadorna) indicative for the intertidal area. Eiders (n = 42 carcasses; n = 120 faecal samples) mainly ingested common cockles (Cerastoderma edule). Blue mussels (Mytilus edulis) ranged second and—same as the invasive razor clam (Ensis leei)—occurred more frequently than reflected by historic data. Proportions of gastropods, bristle worms and crustaceans were low. Shelducks (n = 20 carcasses; n = 98 faecal samples) fed mainly on small molluscs, especially C. edule, mud snails Peringia ulvae, and amphipods. Plastic debris was found in 92.9% of the stomachs and 74.2% of the faeces from eiders as well as in 95% of the stomachs and 85.7% of the faeces from shelducks. Filaments in shelduck prey remains were significantly larger, whereas there was no species-specific difference in abundance of filaments. Most plastic consisted of brightly coloured filaments ˂ 5 mm. These findings indicate regular uptake and excretion of plastic debris by coastal benthivorous seabirds, both in the subtidal and intertidal realm. The origin and mode of uptake of microplastics are discussed.

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.

Minerogenic salt marshes can function as important inorganic carbon stores

AbstractStocks and fluxes of soil inorganic carbon have long been ignored in the context of coastal carbon sequestration, and their implications for the climate cooling effect of blue carbon ecosystems are complex. Here, we investigate the role of soil inorganic carbon in five salt marshes along the northern coast of the European Wadden Sea, one of the world's largest intertidal areas, harboring ~ 20% of European salt‐marsh area. We demonstrate a substantial contribution of inorganic carbon (average: 29%; range: 7–57%) to the total soil carbon stock of the top 1 m. Notably, inorganic exceeded organic carbon stocks in one of the studied sites; a finding that we ascribe to site geomorphic features, such as proximity to marine calcium carbonate sources and hydrodynamic exposure. Contrary to our hypothesis that inorganic carbon stocks would decline along the successional gradient from tidal flat to high marsh, as carbonate deposits would progressively dissolve in increasingly organic‐rich rooted sediments, our findings demonstrate the opposite pattern—an increase in inorganic carbon stocks along the successional gradient. This suggests that the dissolution of calcium carbonates in the root zone is counterbalanced by other processes, such as trapping of sedimentary carbonates by marsh vegetation and calcium carbonate precipitation in anaerobic subsoils. In the context of blue carbon, it will be critical to develop an improved understanding of these plant‐ and microbiota‐mediated processes in calcium carbonate cycling.

An evaluation of long-term physical and hydrochemical measurements at the Sylt Roads Marine Observatory (1973–2019), Wadden Sea, North Sea

Abstract. The Sylt Roads pelagic time series covers physical and hydrochemical parameters at five neighboring stations in the Sylt–Rømø bight, Wadden Sea, North Sea. Since the beginning of the time series in 1973, sea surface temperature (SST), salinity, ammonium, nitrite, nitrate, and soluble reactive phosphorus (SRP) have been measured twice a week. The other parameters were introduced later (dissolved silicate (Si) since 1974, pH since 1979, dissolved organic nitrogen (DON) since 1996, dissolved organic phosphorus (DOP) since 2001, chlorophyll a since 1979, and suspended particulate matter (SPM) since 1975), and in the case of dissolved oxygen, were already discontinued (1979–1983). In the years 1977, 1978, and 1983, no sampling took place. Since the start of the continuous sampling in 1984, the sea surface temperature in the bight has risen by +1.11 ∘C, with the highest increases during the autumn months, while the pH and salinity decreased by 0.23 and 0.33 units, respectively. Summer and autumn salinities are generally significantly elevated compared to spring and winter conditions. Dissolved nutrients (ammonium, nitrite, nitrate, and SRP) have displayed periods of intense eutrophication (1973–1998) and de-eutrophication since 1999. Silicate has shown significantly higher winter levels since 1999. Interestingly, phytoplankton parameters did not mirror these large changes in nutrient concentrations, as a seasonal comparison of the two eutrophication periods showed no significant differences with regard to chlorophyll a. This phenomenon might be triggered by an important switch in nutrient limitation during the time series. With regard to nutrients, the phytoplankton was probably primarily limited by silicate until 1998, while, since 1999, the SRP limitation has become increasingly important. All data are available in Rick et al. (2017b–e, 2020a–o) from https://doi.org/10.1594/PANGAEA.150032, https://doi.org/10.1594/PANGAEA.873549, https://doi.org/10.1594/PANGAEA.873545, https://doi.org/10.1594/PANGAEA.873547, https://doi.org/10.1594/PANGAEA.918018, https://doi.org/10.1594/PANGAEA.918032, https://doi.org/10.1594/PANGAEA.918027, https://doi.org/10.1594/PANGAEA.918023, https://doi.org/10.1594/PANGAEA.918033, https://doi.org/10.1594/PANGAEA.918028, https://doi.org/10.1594/PANGAEA.918024, https://doi.org/10.1594/PANGAEA.918034, https://doi.org/10.1594/PANGAEA.918029, https://doi.org/10.1594/PANGAEA.918025, https://doi.org/10.1594/PANGAEA.918035, https://doi.org/10.1594/PANGAEA.918030, https://doi.org/10.1594/PANGAEA.918026, https://doi.org/10.1594/PANGAEA.918036, and https://doi.org/10.1594/PANGAEA.918031.

Tidally-modulated stratification in a channel-shoal estuary

This study integrates field observations and numerical simulations in the Delaware Estuary. The geometry of the Delaware Estuary is similar to that of South San Francisco Bay, the German Wadden Sea, and the mouth of the Changjiang Estuary. Both the observations and model results demonstrate that the water column was stratified (Δs > 15 psu), in contrast to the prior designations of the Delaware Estuary as a “weakly-stratified” estuary (Garvine et al., 1992). This study implements the salinity variance method to qualify the contributions of different mechanisms to initiating estuarine stratification. The analysis indicates that the tendency for stratification is initialized by the interplay between tidal straining and mixing from the end of spring tides. Although the longitudinal straining decreases during flood tides, the lateral straining contributes to a positive tendency for estuarine stratification. The lateral salinity gradient is not significant under the annual mean discharge condition. The strong vertical shear of flow and deviation from the mean salinity are favorable for the generation of lateral tidal straining. In contrast, a prior study showed that strong lateral salinity gradients enhance vertical stratification during the flood. The lateral straining at the end of the flood provides initial stratification, resulting in enhanced the longitudinal tidal straining during the early ebb due to the inhibition of turbulence and thereby enhanced tidal shear.

Regime shift and changes in sediment morphology driven by sea level rise affect abundance of migratory waders

Regime shifts are likely to have strong impacts on all trophic levels and their interactions. In addition, weather conditions and sea level rise together with tidal currents may change sediment morphology in coastal areas and estuaries. Here, we studied the effects of these drivers of ecosystems on abundance of seven wader species feeding on macrozoobenthos in the German sector of the Wadden Sea. We hypothesized that regime shift caused decreases in wader abundance and that changes in sediment morphology driven by sea level rise and tidal amplitude affected these changes. We analyzed numbers of waders on a short term, 1998–2016, that matched a large-scale geographical study of sediment morphology in the German sector, and at a long term, 1987–2019. Changes in numbers of waders were estimated as slope of coefficients of the trends of wader species numbers in the two periods. The average (se) year of decreases in numbers of waders was initiated for most species and tidal basins in the year 1992 (± 0.4) that followed a regime shift in the Southern North Sea and in the Wadden Sea in 1988–1989. There were no statistical differences between the years when numbers of waders started to decrease between the northern and the southwestern tidal basins of the German sector, indicating that the drives affected both coastal sections simultaneously. Significant relationships were found between slopes of coefficients of wader species and sediment morphology and physical features as sea level rise and tidal amplitude. Relationships were most pronounced for the long term, 1987–2019 compared to the short term, 1998–2016. We conclude that regime shift and changes in sediment morphology driven by sea level rise and sediment change can affect abundance of migratory wader species in large estuaries and shallow coastal waters.

Features of protection of transboundary objects of the world natural heritage: Russian and foreign experience (Part 2)

This article is a continuation of the analysis of the fulfillment by Benin, Niger and Burkina Faso of international obligations and the application of national law norms for the protection of the transboundary object of the V-Arly-Panjari Complex, as well as the implementation by Germany, Denmark and the Netherlands of a set of measures for the protection and conservation of the Wadden Sea. The authors identified the main provisions for the protection and management of the outstanding universal value, considered the foundations for the formation of systems of bodies that perform the functions of protecting objects, analyzed the main measures for coordinating the actions of national authorities to eliminate the main threats and maintain the state of unique natural transboundary objects in order to fulfill obligations under the Convention. The novelty of this study is predetermined by the fact that at the present time there are practically no legal studies of the practice of protecting World Heritage sites in general and transboundary (transnational) sites in particular. The analysis leads the authors to the conclusion that the extrapolation of the main provisions to the current situation in the field of conservation of such Russian transboundary natural objects as the Ubsunur basin and the Landscapes of Dauria gives rise to the need to develop appropriate individual adjustments in the legislation of our country, regardless of the relevant UNESCO measures. The authors make proposals to improve the situation in the field of ensuring its protection of a Russian transboundary natural objects.

Groundwater springs in the German Wadden Sea tidal flat: A fast-track terrestrial transfer route for nutrients and dissolved organic matter

Submarine groundwater discharge (SGD) connects fresh groundwater and marine ecosystems and conveys terrestrially derived dissolved organic matter (DOM) and nutrients from land to sea. The connectivity of terrestrial and marine ecosystems via SGD depends strongly on local environmental settings. For example, SGD composition is modified on its transit through the coastal aquifer, with spring-type SGD from highly permeable aquifers presumably being less affected than diffuse discharge systems from sedimentary environments. In our study, we investigated spring-type SGD near Sahlenburg/Cuxhaven, Northern Germany, which passes through fine, unconsolidated tidal sediments before entering the coastal ocean. We characterized groundwater, surface water and seawater endmembers from different seasons and assessed the potential of tidal sediments impacting the biogeochemistry of “fast-track”, point-source groundwater discharge systems. In addition to physicochemical parameters and nutrients, we analyzed the DOM molecular composition via ultrahigh-resolution mass spectrometry (FT-ICR-MS). Our data revealed a widespread physicochemical and geochemical influence of the groundwater springs on the tidal flat, producing low salinity and low dissolved organic carbon (DOC), and high nitrate and high oxygen concentrations not only in the springs, but also in adjacent porewater. From near- to offshore, salinity and DOC concentrations in springs decreased whereas nitrate and oxygen concentrations increased, resembling an inverse estuarine pattern. Furthermore, high nitrate values suggest anthropogenic sources (e.g., agricultural influence) in the surrounding watershed and may stimulate primary productivity in the tidal flat. Humic-like fluorescent DOM (FDOM) abundances and DOM molecular fingerprints indicated inputs of terrestrial DOM from nearshore saltmarsh plants, as well as from the nearby Elbe and Weser estuaries. Our study demonstrated that SGD had a strong geochemical impact even in the vicinity of large rivers, with productive springs actively hindering sulfate and nitrate reduction by flushing otherwise anoxic systems with oxygen. We posit that the geochemical influence of groundwater springs in tidal flats is underestimated because it can extend far beyond their visual discharge points.

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.

Long-term response of coastal macrofauna communities to de-eutrophication and sea level rise mediated habitat changes (1980s versus 2018)

Since the last decades, previous long-term Wadden Sea studies revealed significant changes in the abundance, biomass and spatial distribution of characteristic macrofauna communities in response to environmental changes and anthropogenic stressors. In this study, we performed statistical community analysis for the East-Frisian Wadden Sea (EFWS, southern North Sea) on two reference datasets across a period with severe climatic and environmental changes (1980s-2018). Therefore, historical macrofauna data from the Quantitative Sensitivity Mapping (1980s, SENSI 1) were reanalyzed and compared with data from the Synoptic Intertidal Benthic Survey (SIBES/SENSI 3) collected in 2018. Our results revealed significant quantitative and spatial changes in the characteristic macrofauna communities between the 1980s and 2018, most likely in response to de-eutrophication and sea level rise mediated habitat changes. Since the 1980s, the total number of taxa remained relatively stable (1980s: 90, 2018: 81), but the total abundance decreased by ca. -31% and the total biomass decreased by ca. -45%, particularly in the eastern regions of the study site probably due to de-eutrophication processes. Thereby, the mean abundances/m2 of ≥ -80% (1980s-2018) in the EFWS of several dominant species decreased: e.g. the gastropod Peringia ulvae, the polychaete Lanice conchilega and the bivalve Mya arenaria. In contrast, the mean abundance/m2 of one dominant species increased by ≥ +80% (1980s-2018): the invasive bivalve Ensis leei. The mean biomass [g/m2] of three dominant species decreased by ≥ -80% (1980s-2018): P. ulvae, L. conchilega and the amphipod Corophium volutator. In contrast, the mean biomass [g/m2] of one dominant species increased by ≥ +75% (1980s-2018): the polychaete Arenicola marina. In the western part of the EFWS, not only higher abundances of A. marina, but also of L. conchilega and P. ulvae were found in 2018, caused i.a. by sea level rise mediated decreasing mud contents and sand accretion on tidal flats. At the community level, the Oligochaeta/Heteromastus community increased in spatial distribution in the western EFWS in 2018 in the vicinity of increasing mussel/oyster beds.

Prey ingestion rates revealed by back-mounted accelerometers in Eurasian spoonbills

BackgroundQuantifying foraging success in space and time and among individuals is essential for answering many ecological questions and may guide conservation efforts. However, collecting this information is challenging for species that forage on mobile prey and are difficult to observe visually, for example, because they forage in inaccessible areas or at night. In such cases, the use of tracking devices that simultaneously collect location and acceleration data may provide a solution if foraging success can be extracted successfully. The aim of this study was to assess how well searching for and ingesting prey, among other behaviours, could be distinguished from 20 Hz acceleration data collected by GPS/ACC-trackers mounted on the back of Eurasian spoonbills Platalea leucorodia. Upon capturing a prey, spoonbills make a distinct movement with their head and back to throw the prey from the tip of the bill into the throat.MethodsWe compared the behavioural classification performance of random forest models that were trained and tested on video-annotated acceleration data segments of different (fixed or flexible) lengths. The best-performing model was then applied to 4 years of data of spoonbills foraging in the Wadden Sea during the breeding season, to explore seasonal and annual variation in prey ingestion rates.ResultsHighest classification accuracies (as indicated by the F-measure, a balanced measure of precision and sensitivity) of foraging behaviours were achieved by analysing short fixed-length segments (0.4–0.8 s) or “flexibly-cut” segments. The F-measure was very high (> 0.90) for searching, standing, sitting and flying (distinguishing active and passive flight), 0.73 for ingesting prey and 0.65 for walking. False positive and negative prey ingestions were equally likely and most often confused with searching, resulting in a close match between the predicted and observed prey ingestion rates. Application of the best-performing model revealed strong seasonal patterns in prey ingestion rates in the Wadden Sea that varied between years.ConclusionsWe show that prey ingestion rates of spoonbills can be fairly accurately estimated from acceleration data. These results are promising for the use of spoonbills equipped with GPS/ACC-trackers as monitors of spatial and temporal variation in the availability of small fish and shrimp, which is key to understand the foraging and migratory movements of spoonbills and provides information on the quality of (coastal) wetlands.

Features of protection of transboundary objects of the world natural heritage: Russian and foreign experience (part 1)

This work is the first part of the article, the subject of which is the practice of fulfillment by Benin, Niger and Burkina Faso of international obligations to protect the transboundary object of the world natural heritage of the "W-Arly-Panjari" Complex, as well as the practice of managing activities for the protection of such an object as Wadden Sea, located in the territories of Germany, Denmark and the Netherlands. The authors highlight the general problems of protecting world heritage sites in the framework of the implementation of the provisions of the Convention, as well as review and analyze the main activities for coordinating the actions of national authorities to preserve and maintain unique natural complexes, in particular, in terms of the implementation of management plans. The analysis of the experience of cooperation between the authorities of Benin, Niger and Burkina Faso with the authorities of the world heritage protection system in relation to the W-Arly-Panjari Complex, as well as Germany, Denmark and the Netherlands in relation to the Wadden Sea, leads to the conclusion that the interaction of national and international funds is effective protection of nature and the need to develop measures aimed at strengthening cooperation between states, including within the framework of the functioning of supranational systems and legal institutions in order to preserve transboundary natural heritage sites.

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.

Introduced species in a tidal ecosystem of mud and sand: curse or blessing?

For about a century, biodiversity in the tidal Wadden Sea (North Sea, European Atlantic) has increased by more than one hundred introduced species from overseas. Most originate from warmer waters and could facilitate the transformation of this coastal ecosystem to comply with climate warming. Some introduced species promote sediment stabilization and mud accretion. This could help tidal flats to keep up with sea level rise. Although some introduced species also entail negative effects, introductions have diversified lower food web levels, and may benefit foraging birds. So far, no resident populations have gone extinct because an introduced species had established. Rather than degrading the ecosystem, the establishment of introduced species seems to have raised the capacity to follow environmental change. We support increasing efforts against introductions to avoid risk. However, once species are integrated, the common condemnation attitude against “non-natives” or “aliens” ought to be reconsidered for tidal ecosystems of low biodiversity.

Mollusc shell detritus affects benthic subtidal community dynamics in the Northern Wadden Sea

A shallow subtidal area in the northern Wadden Sea was monitored over 17 years (from 2003 to 2019) for sediment parameters and macrobenthic fauna. Due to the sheltered position of the study area, sediment composition remained rather stable with only minor annual and seasonal variations in sediment granulometry. An intermittend storm (‘Kyrill’) had no significant effect on sediment composition parameters; the construction of an artificial dune along the southern border of the study area had minor transient but no lasting effects on sediment composition. Faunal species richness and total abundance showed the typical seasonality with minimum abundance in late winter and a peak after recruitment in summer. Variations between years in autumn (post-recruitment) abundances were best explained by the number of days with a seawater temperature <1°C during the preceding winter. Temperature during other seasons, salinity, chlorophyll concentration and NAO showed no significant correlations with total abundance, nor did storm (‘Kyrill’) or construction of the artificial dune. Within-site faunal variability was best explained by water depth and velocity of the tidal currents while sediment granulometry was of minor importance. However, the amount of bivalve shell detritus mixed with the sandy sediment proved to be the strongest covariant of species numbers, total abundance, and species-specific abundances. At the sediment surface, shells provide the only anchorage for epibenthic species which in turn attract associated fauna. Shells inside the sediment hamper movement of infaunal predators and epibenthic predators are handicapped in rooting about for prey. Thus, shell material is a highly significant structural factor for the macrozoobenthos in these shallow waters. In a future with increasing ocean acidification, the availability of benthos as a food source for higher trophic levels will depend on the balance between pH-driven shell destruction and the compensatory power of shell-building species.

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).