Ecosystem model of the western Wadden Sea: A bridge between science and management?

This paper describes the construction, validation and application of a mathematical ecosystem model of the western Wadden Sea. This model is based upon the Ems-Dollard ecosystem model, but also includes a two-dimensional transport submodel, a benthic and epibenthic sublitoral submodel and nutrient regeneration. In 1986, an extensive field programme was executed to collect data for model validation. Apart from the subtidal processes the validation indicates that the model simulates the different state variables reasonably well. The applicability of the model for management purposes was tested in four case studies concerning: dredging activities, eutrophication, commercial mussel cultrue and oil spills and other calamities. It is concluded that the western Wadden Sea model in general is a useful tool in evaluating the impact of certain anthropogenic activities in the area. Presented at the VI International Wadden Sea Symposium (Biologische Anstalt Helgoland, Wattenmeerstation Sylt, D-2282 List, FRG, 1–4 November 1988). Publication nol 7. Applied Scientific Research NIOZ (BEWON)     

Human transformations of the Wadden Sea ecosystem through time: a synthesis

Todays Wadden Sea is a heavily human-altered ecosystem. Shaped by natural forces since its origin 7,500 years ago, humans gradually gained dominance in influencing ecosystem structure and functioning. Here, we reconstruct the timeline of human impacts and the history of ecological changes in the Wadden Sea. We then discuss the ecosystem and societal consequences of observed changes, and conclude with management implications. Human influences have intensified and multiplied over time. Large-scale habitat transformation over the last 1,000 years has eliminated diverse terrestrial, freshwater, brackish and marine habitats. Intensive exploitation of everything from oysters to whales has depleted most large predators and habitat-building species since medieval times. In the twentieth century, pollution, eutrophication, species invasions and, presumably, climate change have had marked impacts on the Wadden Sea flora and fauna. Yet habitat loss and overexploitation were the two main causes for the extinction or severe depletion of 144 species (~20% of total macrobiota). The loss of biodiversity, large predators, special habitats, filter and storage capacity, and degradation in water quality have led to a simplification and homogenisation of the food web structure and ecosystem functioning that has affected the Wadden Sea ecosystem and coastal societies alike. Recent conservation efforts have reversed some negative trends by enabling some birds and mammals to recover and by creating new economic options for society. The Wadden Sea history provides a unique long-term perspective on ecological change, new objectives for conservation, restoration and management, and an ecological baseline that allows us to envision a rich, productive and diverse Wadden Sea ecosystem and coastal society.     

Climatic change and the Wadden Sea,The Netherlands

The predicted increase in atmospheric carbon dioxide and the effects of global warming will influence the Wadden Sea, The Netherlands, an area of exceptional ecological value. The direct effect of elevated atmospheric CO₂ on terrestrial coastal ecosystems is either marginal or unknown. The slight acidification of the sea which is predicted might have an impact on primary producers and juvenile animals. The effect of CO₂ fertilization on marine primary production remains to be elucidated. Profound changes will occur if sea level rises at the predicted rate of 60 cm per century, as sedimentation rates will be insufficient to maintain the salt marshes on the barrier islands. The marshes of the mainland coast will be impoverished, as high and low marshes are not expected to continue to coexist at the same locations. As sediment supply to the Wadden Sea is sufficient to compensate for sea level rise, the estuarine character of the Wadden Sea, with sand- and mudflats, is expected to remain largely unchanged.     

General Patterns in Invasion Ecology Tested in the Dutch Wadden Sea: The Case of a Brackish-Marine Polychaetous Worm

The success of invasive aquatic species is determined by a variety of attributes such as wide environmental tolerance, high genetic variability, short generation time, early sexual maturity, high reproductive capacity, and a broad diet. Usually, introduced species, after some time lag since inoculation, show an exponential population increase and expansion. Maintenance of the immigrant species at a high population level will be dependent on interspecific competition with native species and availability of habitat and food. Eventually, the immigrant population may decline, for instance due to increased predation pressure, parasite infestation or loss of genetic vigour. These characteristic patterns in invasive species are reviewed for the case of the North American spionid polychaete Marenzelleria cf. wireni in the Dutch Wadden Sea. This species was first recorded in estuaries and coastal waters of the European continent in the Ems estuary (eastern Dutch Wadden Sea) in 1983. In the western part of the Dutch Wadden Sea the first specimens were found in 1989. The Ems estuary population showed the typical lag-phase, explosive increase, stabilisation, and eventual decline. In the western part of the Dutch Wadden Sea the latter two phases have not yet developed. The strong development and stabilisation of the population in the Ems estuary may have been caused by the availability of a yet not utilised food source. The species' final decline remains largely unexplained.     

Human impacts on fisheries resources and abundance in the Danish Wadden Sea, c1520 to the present

For the past 30 years, the Danish Wadden Sea area has been protected by fishery and hunting regulations, pollution controls and settlement restrictions. Nevertheless, todays Wadden Sea is a depleted ecosystem when we compare it with past abundance and diversity of marine animal populations. This review indicates that the abundance of finfishes has undergone a long-term decline since the seventeenth century. The review also indicates a trajectory of ecological decline since the early twentieth century that seems to be related to: (1) fishing extractions which focused on undersized fish throughout the first half of the twentieth century; (2) habitat destruction, which was most marked in the first half of the twentieth century; and (3) pollution, which was worst in the third quarter of the twentieth century. Historical investigation reveals that we need to fundamentally revise present-day baselines about the potential species richness and abundance of the Wadden Sea.     

Using an Ecosystem Modeling Approach to Explore Possible Ecosystem Impacts of Fishing in the Beibu Gulf,Northern South China Sea

Using the Ecopath with Ecosim software, a trophic structure model of the Beibu Gulf was constructed to explore the energy flows and provide a snapshot of the ecosystem operations. Input data were mainly from the trawl survey data collected from October 1998 to September 1999 and related literatures. The impacts of various fishing pressure on the biomass were examined by simulation at different fishing mortality rates. The model consists of 20 functional groups (boxes), each representing organisms with a similar role in the food web, and only covers the major trophic flows in the Beibu Gulf ecosystem. It was found that the food web of the Beibu Gulf was dominated by the primary producers path, and phytoplankton was the primary producer mostly used as a food source. The fractional trophic levels ranged from 1.0 to 4.02, and the marine mammals occupied the highest trophic level. Using network analysis, the ecosystem network was mapped into a linear food chain, and six discrete trophic levels were found with a mean transfer efficiency of 11.2%. The Finn cycling index was 9.73%. The path length was 1.821. The omnivory index was 0.197. The ecosystem had some degree of instability due to exploitation and other human activities, according to Odum’s theory of ecosystem development. A 10-year simulation was performed for each fishery scenario. The fishing mortality rate was found to have a strong impact on the biomass. By keeping the fishing mortality rate at the current level for all fishing sectors, scenario 1 had a drastic decrease in the large fish groups. The biomass of the small and medium pelagic fish would increase to some extent. The biomass of the small and low trophic level species, jellyfish, prawns and benthic crustaceans would be stable. The total biomass of the fishery resources would have a 10% decrease from the current biomass after 10 years. In contrast, the reduced fishing mortality rate induced the recovery of biomass (scenarios 2–4). In scenario 2, the biomass of the large demersal fish and the large pelagic fish would increase to over 16 times and 10 times, respectively, of their current level. In scenario 4, the biomass of the large pelagic fish would increase to over 3 times of its current level. The total biomass of the fish groups, especially the high trophic level groups, would become significantly higher after 10 years, which illustrates the contribution on biomass recovery by relaxing the fishing pressure. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Author contributions: Xiaoping Jia designed research; Zuozhi Chen and Yongsong Qiu performed research; Zuozhi Chen, Yongsong Qiu, and Shannan Xu analyzed data; and Zuozhi Chen and Shannan Xu wrote the article.     

Kudzu (Pueraria montana): History,Physiology, and Ecology Combine to Make a Major Ecosystem Threat

Kudzu, Pueraria montana (Lour.) Merr. variety lobata (Willd.) was introduced into the United States at the 1876 Centennial Exposition in Philadelphia, PA. Subsequently, it was planted widely to reduce soil erosion by the Soil Erosion Service and Civilian Conservation Corp. Over 85 million seedlings of kudzu were provided to landowners by government agencies in the southeast in the first half of the 20th century. In 1953, kudzu was removed from the list of approved plants for erosion control, in 1970 it was officially labeled a weed, and in 1997 it was placed on the Federal Obnoxious Weed List. Its rapid elongation rates, high leaf area indices, high photosynthetic rates, and frequent rooting at stem nodes make kudzu an aggressive competitor with native shrubs and trees. Estimates are that kudzu currently covers 3 million hectares throughout the eastern United States and is spreading by 50,000 ha per year. Despite widespread anecdotal statements, little quantitative information is available regarding the ecological effects of kudzu. The ability of kudzu to overtop and shade forest trees, fix atmospheric nitrogen, and emit isoprene suggest that it may have substantial effects on native forest biodiversity, forest nitrogen cycles, watershed nitrogen saturation, freshwater eutrophication, and regional air quality. Kudzu's growth rate increases strongly in response to increased CO₂, and without the constraint of allocation to woody tissue this response may increase the competitive dominance of kudzu as atmospheric levels of CO₂ increase. This fact, combined with its sensitivity to cold temperatures, implies that kudzu may increase its range in future warmer, high-CO₂ environments. The lack of quantitative investigations on the ecological effects of kudzu is a severe impediment to our understanding of its current and future effects on native plant and animal communities and forest ecosystems.     

Historical Changes in Marine Resources,Food-web Structure and Ecosystem Functioning in the Adriatic Sea,Mediterranean

The Mediterranean Sea has been strongly influenced by human activities for millennia. Although the environmental history of its surrounding terrestrial ecosystems has received considerable study, historical changes in its marine realm are less known. We used a multidisciplinary approach combining paleontological, archeological, historical, fisheries, and ecological data to reconstruct past changes in marine populations, habitats, and water quality in the Adriatic Sea. Then, we constructed binary food webs for different historical periods to analyze possible changes in food-web structure and functioning over time. Our results indicate that human activities have influenced marine resource abundance since at least Roman times and accelerated in the nineteenth and twentieth centuries. Today, 98% of traditional marine resources are depleted to less than 50% of former abundance, with large (>1 m) predators and consumers being most affected. With 37% of investigated species rare and 11% extirpated, diversity has shifted towards smaller, lower trophic-level species, further aggravated by more than 40 species invasions. Species providing habitat and filter functions have been reduced by 75%, contributing to the degradation of water quality and increased eutrophication. Increased exploitation and functional extinctions have altered and simplified food-web structure over time, especially by changing the proportions of top predators, intermediate consumers, and basal species. Moreover, simulations of species losses indicate that today’s ecosystems may be less robust to species extinctions than in the past. Our results illustrate the long-term and far-reaching consequences human activities can have on marine food webs and ecosystems.     

Coast of change: habitat loss and transformations in the Wadden Sea

In the southern North Sea, coastal people commenced with habitat conversions 1,000 years ago. Partly interrupted in late medieval times by large-scale inundations of marshland, progressive embankments transformed the landward half of the amphibic transition zone between a limno-terrestric and a brackish-marine ecosystem into arable land and freshwater lakes. Sea walls rigidly separated the land from the sea. Dynamic transitional habitats have vanished. Areal loss has diminished the capacity of the Wadden Sea to dissipate wave and tidal energy. A coastal ecosystem once rich in marsh plants, seagrass and diatoms on mud flats became transformed into one with less autochthonous phototroph production, dominated by sandy tidal flats, and dependent primarily on allochthonous plankton supply. The large estuaries have been dredged to serve as shipping canals, and have lost most of their former retention and filter capacity. Riverine loads are now flushed right into the North Sea. Symptoms of a syndromatic coastal habitat degradation are diagnosed, leading to a decline in natural habitat diversity. The conventional on-line coastal protection may not achieve a sustainable coastal habitat configuration. At sedimentary coasts immobilised by dikes and petrified shores, a more flexible response to sea level rise is recommended.     

Associations between macrobenthos and invasive cordgrass, Spartina anglica, in the Danish Wadden Sea

We conducted a survey to provide knowledge on mechanisms controlling spatial and temporal variability of macrobenthos in an intertidal Wadden Sea area partly covered by invasive Spartina anglica. Benthic macrofauna was collected seasonally at seven stations along a transect covering non-vegetated mudflat and vegetated marsh areas. Shannon diversity index was consistently higher for macrobenthos in the open mudflat compared to the marsh area. Infaunal species, like Arenicola marina, Tubificoides benedeni and Macoma balthica, were more abundant in the mudflat than the marsh, while the opposite was evident for epifaunal species, like Hydrobia ulvae and Littorina littorea. The infaunal crustacean, Corophium volutator, on the other hand, appeared particularly attracted to the mudflat–marsh boundary. The biomass of below-ground plant materials and macrodetritus was positively correlated with the total macrofaunal abundance, while the biomass of dead below-ground plant materials alone showed a significant negative correlation with infaunal abundance. Total macrofaunal abundance was highest during summer in the mudflat, while no seasonal pattern was evident in the marsh. The negative influence of S. anglica invasions on infaunal abundance may have serious implications for higher trophic levels, such as waterbirds, that forage in Wadden Sea areas and thus for the overall biodiversity and ecosystem functioning.     

Macrobenthos of the subtidal Wadden Sea: revisited after 55 years

During the years 1923–1926 Hagmeier & Kändler (1927) sampled the macrofauna of subtidal shallows and channels of the Wadden Sea close to the Island of Sylt (German Bight, North Sea). Reinvestigating this study area in 1980, a substantially altered faunal composition was recorded. An approach is made to quantify the comparison in terms of abundance, species richness and diversity of invertebrate taxa. Human interference is assumed to be responsible for the major changes. Natural oyster beds have been overexploited and the local population ofOstrea edulis has been driven to extinction. Subsequently, mussels(Mytilus edulis) spread in the entire region, promoted by shell fishery. Particularly barnacles and many polychaetes took advantage of the expansion of mussel banks which is substantiated by correlation analysis. Reefs of the colonial polychaeteSabellaria spinulosa stood in the way of shrimp trawling and became destroyed together with the associated fauna. A subtidalZostera marina bed was wiped out in 1934 by a natural epidemic disease but never succeeded in reestablishing itself. The associated fauna disappeared. Large epibenthic predators and scavengers (crabs, snails and starfish) survived all these changes. The total number of species remained approximately at the same level but molluscs experienced losses and polychaetes diversified. Overall abundance increased with a disproportionately large share of a few species(Mytilus edulis, Balanus crenatus, Cerastoderma edule, Scoloplos armiger). The subtidal fauna of the Wadden Sea proved to be vulnerable to human disturbance; thus, the present community can no longer be viewed as the outcome of entirely natural processes.     

Distribution patterns of bacterioplankton and chlorophyll-a in the German Wadden Sea

 In a first synoptic evaluation, the temporal and spatial distribution of bacterioplankton and chlorophyll-a were determined in the German Wadden Sea. Three surveys were undertaken in winter, spring, and summer of 1994 using up to eight ships simultaneously between the river Ems and Sylt island. Despite intensive hydrodynamic mixing of the Wadden Sea water, spatial gradients were obvious. The abundance of bacterioplankton ranged from 0.4 to 26×10⁵ ml^–1 and chlorophyll-a varied between <0.1 and 79 μg l^–1. In winter, relatively homogeneous distribution patterns of both parameters with small gradients were found. Highest chlorophyll-a values connected with a highly patchy structure were observed in spring, while in summer both total chlorophyll-a values and the complexity of the distribution pattern had decreased. In contrast, bacterial numbers increased steadily from January to July with the highest bacterial densities and greatest patchiness observed in summer. Moreover, in some regions of the Wadden Sea, a trophic succession of algae as carbon producers and bacteria as consumers was evident. Correlation analysis verified the relationship between bacteria and chlorophyll a, indicating bottom-up control of bacterial abundance in the northern part of the German Wadden Sea. Since the observed regression slope is remarkably low (0.12–0.46) compared to literature values (0.5–0.8), we suggest that the link between phytoplankton and bacteria found here is a special characteristic of the Wadden Sea as a transition zone between the coastal region and the outer North Sea. Received: 26 May 1998 / Accepted:12 November 1998     

Alien parasitic copepods in mussels and oysters of the Wadden Sea

Molluscan intestinal parasites of the genus Mytilicola, specifically M. intestinalis, were initially introduced into bivalves in the North Sea in the 1930s. It was presumably introduced from the Mediterranean with ship-fouling mussels, then attained epidemic proportions in Mytilus edulis in the 1950s and is now widely established in the North Sea region. Mytilicola orientalis was co-introduced with Pacific oysters to France in the 1970s and in the southern North Sea in the early 1990s. Its main host Crassostrea gigas has massively invaded the Wadden Sea with a concomitant decline in mussels. To explore whether introduced mytilicolid parasites could play a role in the shifting dominance from native mussels to invasive oysters, we analysed 390 mussels and 174 oysters collected around the island of Sylt in the northern Wadden Sea. We show that M. intestinalis has a prevalence >90% and a mean intensity of 4 adult copepods in individual mussels with >50 mm shell length at all sheltered sites. By contrast, none were found in the oysters. However, at one site, we found M. orientalis in C. gigas with a prevalence of 10% and an intensity of 2 per host individual (August 2008). This constitutes the most northern record in Europe for this Pacific parasite until now. Alignments of partial sequences of the mitochondrial cytochrome oxidase I (COI) gene and the nuclear internal transcribed spacers (ITS) and 18S rDNA sequences each show a distinct difference between the two species, which confirms our morphological identification. We suggest that the high parasite load in mussels compared to oysters may benefit the continued expansion of C. gigas in the Wadden Sea.     

Large-Scale Spatial Dynamics of Intertidal Mussel (Mytilus edulis L.) Bed Coverage in the German and Dutch Wadden Sea

Intertidal blue mussel beds are important for the functioning and community composition of coastal ecosystems. Modeling spatial dynamics of intertidal mussel beds is complicated because suitable habitat is spatially heterogeneously distributed and recruitment and loss are hard to predict. To get insight into the main determinants of dispersion, growth and loss of intertidal mussel beds, we analyzed spatial distributions and growth patterns in the German and Dutch Wadden Sea. We considered yearly distributions of adult intertidal mussel beds from 36 connected tidal basins between 1999 and 2010 and for the period 1968–1976. We found that in both periods the highest coverage of tidal flats by mussel beds occurs in the sheltered basins in the southern Wadden Sea. We used a stochastic growth model to investigate the effects of density dependence, winter temperature and storminess on changes in mussel bed coverage between 1999 and 2010. In contrast to expectation, we found no evidence that cold winters consistently induced events of synchronous population growth, nor did we find strong evidence for increased removal of adult mussel beds after stormy winter seasons. However, we did find synchronic growth within groups of proximate tidal basins and that synchrony between distant groups is mainly low or negative. Because the boundaries between synchronic groups are located near river mouths and in areas lacking suitable mussel bed habitat, we suggest that the metapopulation is under the control of larval dispersal conditions. Our study demonstrates the importance of moving from simple habitat suitability models to models that incorporate metapopulation processes to understand spatial dynamics of mussel beds. The spatio-dynamic structure revealed in this paper will be instrumental for that purpose.