Red List of macrofaunal benthic invertebrates of the Wadden Sea

Parasites were observed in medium- and small-sized fish taken from the discards of a commercial shrimper during seven different cruises in the tidal channels of the North Frisian Wadden Sea (Süderaue, North Sea) from April to September 1991. In total, 442 fish comprising four species (Sprattus sprattus, Hyperoplus lanceolatus, Ammodytes tobianus, Pomatoschistus minutus) were investigated. The parasite fauna consisted of 22 species. The parasite community structure of the 4 hosts was compared. The diet of the hosts seemed to be the main factor determining the structure of the parasite community. Other factors could not be assessed. Eight species of parasites occurred as larval stages. This indicated that fish were intermediate or paratenic hosts in their life cycle. The nematodeHysterothylacium sp. (Anisakidae) and the digeneanCryptocotyle lingua (Heterophyidea) were the dominant parasites, reaching their highest prevalence and density in sprat and sand eel. Sprat and sand eel play a very important role in parasite transmission to predacious fish and seabirds.     

Chemical monitoring in the Dutch Wadden Sea by means of benthic invertebrates and fish

In monitoring, it is of utmost importance to carefully define the purpose, the sampling strategy, as well as the analytical chemical and statistical requirements. Surveys are appropriate for describing the geographical variation in environmental contaminant levels. Repeated surveys and recurrentdata collection at permanent locations provide means of detecting temporal trends. Results are presented here of surveys on pollution by trace metals, polychlorinated biphenyls and organochlorine pesticides in the Ems Estuary and Dutch Wadden Sea usingMytilus edulis, Mya arenaria, Arenicoia marina, Nereis diversicolor andCrangon crangon as test organisms. Trends towards decreasing pollution by mercury are illustrated by monitoring data onMytilus edulis andZoarces viviparus. It is stressed that the results of chemical monitoring in organisms may be interpreted only in termser the biological effects on the basis of relevant toxicological knowledge and/or additional bio-assays. Presented at the VI International Wadden Sea Symposium (Biologische Anstalt Helgoland Wattenmeerstation Sylt, D-2282 List, FRG, 1–4 November 1988)     

Effects of herbivorous birds on intertidal seagrass beds in the northern Wadden Sea

During autumn migration (September to December), brent geese (Branta b. bernicla) and wigeon (Anas penelope) feed on the seagrass Zostera noltii in the nearshore, upper tidal zone leeward of the island of Sylt (eastern North Sea). To graze on leaves and shoots above the sediment and on rhizomes and roots below, these birds reworked the entire upper 1 cm layer of sediment eight times within this 3-month period. In addition, brent geese excavated pits 3–10 cm deep by trampling in order to feed on below-ground phytomass. About 12% of the seagrass beds became pitted to an average depth of 4.5 cm. Using net exclosures, it was estimated that birds removed 34 g dry weight m^–2 of above-ground and 28 g of below-ground phytomass. This corresponds to 45% of the phytomass in September. Of the overall loss of phytomass from September to December, 63% was caused by birds. Roughly half of the leaves fell off anyway until December and the other half were taken by the birds. Below the ground, phytomass remained almost constant where birds were excluded, while with birds phytomass of rhizomes and roots was halved. In spite of this strong effect, in the next vegetation period the blade density was lower at former exclosure sites compared to the ambient seagrass bed. The underlying process seems to be a self-inhibition of dense overwintering seagrass by mud accretion. Assuming our experimental results can be scaled up to the entire seagrass bed, we hypothesize that in the sheltered upper intertidal zone, seasonal erosion caused by herbivorous geese and ducks is necessary for the persistence of Z. noltii. Received: 7 January 1999 / Received in revised form: 23 August 1999 / Accepted: 25 August 1999     

The effects of mechanical and manual digging for lugworms (Arenicola marina L.) on the benthic fauna of the tidal flats in the Dutch Wadden Sea

Effects of manual bait digging and mechanical cropping ofArenicola marina (L.) on the macrobenthic animals in the Dutch Wadden Sea were investigated. Samples were taken on two tidal flat areas: the Balgzand and the Vlakte van Kerken, from March to November 1981. Only the impacts on major species and on the total biomass are discussed.The numbers of most of the major species are severely reduced immediately after mechanical digging. Some species (e. g. Macoma balthica, Scoloplos armiger) show a fast migration to the depopulated area. Mechanical cropping appears to be more efficient, catching moreArenicola per m². However, manual digging causes less mortality of other benthic animals perArenicola. Recovery of the diminished abundance varies per species; in general the areas harvested show a larger recruitment compared with the non-disturbed areas.     

Management of the Wadden Sea

The Wadden Sea situated along the North Sea coasts of Denmark, the Federal Republic of Germany and The Netherlands represents one of the world's largest bar-built type of estuaries. The area is a typical sedimentation and mineralization basin, with a large influx of organic matter from the adjoining North Sea, consequently a delicate oxygen balance and a rich benthic macrofauna, poor in species, which serves as food for juveniles of some commercially important North Sea fishes and for large numbers of migrating and wintering waders and waterfowl. Past and present activities of the human society in the area include fisheries (mainly for shrimp and mussels, semi-culture), shipping, land reclamation, recreation, dredging for sand and shells, and waste discharge from industries and human communities. Until the present these activities, although sometimes conflicting, did not fundamentally affect the area and its biota (pollution excluded), but future claims, including the construction of large deep-sea harbours, drilling for natural gas and oil, large-scale land reclamation and increased industrialization etc., might gradually induce degradation. For instance, area reduction by continued land reclamation could lead to irreversible losses of specific biotopes (e. g. salt-marshes, mud-flats), which could affect the size of bird and fish populations in a much wider region. Increased pollution, which has already inflicted damage on bird and seal populations, could reduce the fauna and hence the value of the area as a natural sanctuary. In the event of a proposal for a new human activity in the area, the present standing practice in the countries concerned requires an evaluation of its safety and economic aspects and its environmental impact. However, the various plans are considered separately and there is a general need for integrated management of the area.     

Now an empty mudflat: past and present benthic abundances in the western Dutch Wadden Sea

The benthic fauna of two areas in the western Dutch Wadden Sea, Posthuiswad and Staart van Schieringhals, was described in 1930–1960 and again between 1996 and 2005. Here, we document the changes. Whereas both areas formerly had high densities of species that biogenically structured the intertidal mudflats such as mussels Mytilus edulis and cockles Cerastoderma edule, by 1996 they had shown a tenfold decrease in the densities of molluscs, with no recovery till 2005. Although the number of species of polychaetes and crustaceans may not have changed much, their relative abundance did. Nowadays, more polychaete species are common than before. We briefly discuss whether the changes in benthic community composition could be due to industrial fishery practices or eutrophication effects.     

Environmental influences on benthic fauna associations of the Kara Sea (Arctic Russia)

Macrobenthic faunal associations, hydrography and sediment structure were examined at 14 stations in the Kara Sea. The stations were located in an area influenced by huge runoff from the Ob and Yenisei Rivers and in areas influenced by Barents Sea water. Sampling depths varied from 17 to 43 m, with one station at 195 m. The sediments were predominantly muddy but some stations were sandy. Three hundred and eighty-seven taxa were identified and Polychaeta, Crustacea and Mollusca were the most conspicuous. Species number, abundance and biomass varied widely among stations, and were generally higher in the more marine waters. Boreal-arctic species predominated, but an increase of arctic species from marine to the estuarine areas was evident. Five faunal associations were delineated by cluster analysis and suggested quite heterogeneous sampling areas. The most conspicuous species of each faunal association were Spiochaetopterus typicus, Tridonta borealis, Serripes groenlandicus, Portlandia arctica, and Marenzelleria arctia, respectively. The sedimentation rate, as well as depth, sediment structure and salinity, apparently influenced the main differences in the fauna. Accepted: 29 June 1999     

Community structure and spatial distribution of benthic fauna in the Bellingshausen Sea (West Antarctica)

The structure and spatial distribution of the macrofauna community of the Bellingshausen Sea in the western sector of Antarctica was studied during the ‘BENTART–06’ oceanographic expedition. This is one of the least explored Antarctic seas. A total of 20 box cores were sampled at 11 stations ranging from 157 to 3,304 m depth, using an USNEL-type box corer (BC) dredge. Representatives of 25 higher taxa of invertebrates were collected. Deeper sampling sites were less rich in taxa (4–7 taxa), whereas the figures were higher at shallower sites (up to 17 taxa). Faunal density on the sea bottom revealed a horizontal spatial gradient from the western sites with extremely low figures (90 indiv./m²) towards the eastern ones with the highest figures (1,360 indiv./m²) close to the Antarctic Peninsula. Several abiotic factors (depth, redox, organic matter, carbonates and particle size of surficial sediments) were measured simultaneously on the sea floor to characterise the substrate preferences of the fauna. Positive correlations were found between the faunal distribution and a combination of depth, redox values, and organic matter content of sediments. This indicates decreasing availability of food in the deeper bottoms of the Bellingshausen Sea with a prevalence of depauperated bottoms dominated almost exclusively by a foraminiferans community.     

Notes on growth and behaviour of the American razor clamEnsis directus in the Wadden Sea and the predation on it by birds

Ensis directus (Conrad), a bivalve found in Europe only in the past few years, has filled an empty niche and spread rapidly over the intertidal flats in the Wadden Sea. The growth of the animals was investigated using a few samples taken near Langeness in February 1984. The age of the animals could be determined by reading the clearly visible year marks on the shells. Notwithstanding the low density ofE. directus, some oystercatchers appear to have already specialized in feeding on this new food source.     

Eutrophication and mussel culture in the western Dutch Wadden Sea: Impact on the benthic ecosystem; a hypothesis

Since 1950, two large-scale changes have taken place in the western Dutch Wadden Sea, namely the eutrophication of the area and the introduction of an extensive mussel culture. Although eutrophication in the fresh waters started already around 1950, nutrient concentrations in the western Wadden Sea remained fairly constant until about 1970, due to the retention of nutrients in Lake IJssel, the main source. From 1970–1980 concentrations increased strongly, and during the last years the situation has stabilized. Mussel culture was introduced in 1950 and expanded during the next decade to an area of 70 km², all situated in the sublittoral area. From 1960 the area of mussel culture remained about constant with fluctuating yields of between 35 and 120 millions of kg fresh weight. Due to a lack of data for the period until 1970 the impact of eutrophication and mussel culture on the ecosystem cannot be assessed. From 1970 onwards an increased biomass and production of the macrofauna in the intertidal zone has been observed, which is attributed to eutrophication. The hypothesis is postulated that the introduction of mussel culture between 1950 and 1960 has resulted in an increased food competition in the area, leading to a decreased stock of the macrofauna in the intertidal. Eutrophication from about 1970 onwards has improved the food conditions and as a result both the macrofauna in the intertidal and the mussel in the sublittoral area would have increased in biomass, allowing higher maximum yields of the mussel culture. The importance of monitoring programs is stressed to follow these trends in the near future and to check the above hypothesis in areas where it is decided to introduce or intensify mussel culture. Presented at the VI International Wadden Sea Symposium (Biologische Anstalt Helgoland, Wattenmeerstation Sylt, D-2282 List, FRG, 1–4 November 1988) Publication No. 30 of the project “Ecological Research of the North Sea and Wadden Sea” (EON)     

Epibenthic fauna dredged from tidal channels in the Wadden Sea of Schleswig-Holstein: spatial patterns and a long-term decline

The epibenthic fauna dredged in the channels of the Wadden Sea of Schleswig-Holstein is dominated by crustaceans. Mean species richness is lower in a southern region (7.6±2.2 species haul⁻¹) which is part of the Elbe estuary and lacks protective barrier islands. A northern region is more marine and is protected by a seaward barrier of high sands. Here mean species richness is 10.9±2.8. Within channels, there is no significant inshore-offshore gradient, and species number does not increase with depth. Stratified sampling near the island of Sylt revealed that abundance and diversity are high along the gentle upper slope (5 m depth) of channels when mussels are present, but low in the absence of mussels. Abundance and diversity are also low along the steeper slope below (at 10 and 15 m depth). In a channel with scattered stones and boulders, highest diversity (16.0±5.1) and dense epibenthic assemblages were encountered along the upper and middle slopes (5 and 10 m). Comparisons with historical surveys suggest that a decline of nearly fifty percent of all epifaunal species within the last hundred years may be attributed to fishery disturbances.     

Moderate predation on meiofauna by the macrobenthos of the Wadden Sea

The role of meiofauna in the trophic web of marine ecosystems is a controversial topic. During an experimental study on tidal flats in the eastern part of the North Sea, potential predators on meiofauna were kept in enclosures. A gobiid fish, shrimp, crabs, an amphipod, four polychaete worms and a nemertine were tested for their ability to decimate nematodes, turbellarians, ostracods and benthic copepods. Only few macrobenthic species preyed heavily on permanent meiofauna: juveniles of the shore crabCarcinus maenas, the hermit crabPagurus bernhardus with the colonial hydrozoanHydractinia echinata on its shells, and the rag wormNereis diversicolor. When benthic infauna was protected from these predators with exclosures, juvenile macrofauna responded with a marked increase in number, while the permanent meiofauna remained unaltered except for a limited increase in nematodes. It is concluded that in the Wadden Sea abundance of permanent meiofauna (Nematoda, Turbellaria, Ostracoda, Copepoda) is only locally or temporarily regulated by macrobenthic predators.I acknowledge a grant from the Deutsche Forschungsgemeinschaft (DFG).     

Increasing species richness of the macrozoobenthic fauna on tidal flats of the Wadden Sea by local range expansion and invasion of exotic species

A 40-y series of consistently collected samples (15 fixed sampling sites, constant sampled area of 15 × 0.95 m², annual sampling only in late-winter/early-spring seasons, and consistent sieving and sorting procedures; restriction to 50 easily recognizable species) of macrozoobenthos on Balgzand, a tidal flat area in the westernmost part of the Wadden Sea (The Netherlands), revealed significantly increasing trends of species richness. Total numbers of species annually encountered increased from ~28 to ~38. Mean species density (number of species found per sampling site) increased from ~13 to ~18 per 0.95 m². During the 40 years of the 1970–2009 period of observation, 4 exotic species invaded the area: (in order of first appearance) Ensis directus, Marenzelleria viridis, Crassostrea gigas, and Hemigrapsus takanoi. Another 5 species recently moved to Balgzand from nearby (subtidal) locations. Together, these 9 new species on the tidal flats explained by far most of the increase in total species numbers, but accounted for only one-third of the observed increase in species density (as a consequence of the restricted distribution of most of them). Species density increased particularly by a substantial number of species that showed increasing trends in the numbers of tidal flat sites they occupied. Most of these wider-spreading species were found to suffer from cold winters. During the 40-y period of observation, winter temperatures rose by about 2°C and cold winters became less frequent. The mean number of cold-sensitive species found per site significantly increased by almost 2 per 0.95 m². Among the other species (not sensitive to low winter temperatures), 6 showed a rising and 2 a declining trend in number of occupied sites, resulting in a net long-term increase in species density amounting to another gain of 1.6 per 0.95 m². Half of the 50 studied species did not show such long-term trend, nor were invaders. Thus, each of 3 groups (local or alien invaders/winter-sensitive species/other increasing species) contributed to a roughly similar extent to the overall increase in species density.     

Planktonic primary production in the German Wadden Sea

By combining weekly data of irradiance, attenuation and chlorophylla concentrations with photosynthesis (P) versus light intensity(E) curve characteristics, the annual cycle of planktonic primaryproduction in the estuarine part of the Northfrisian WaddenSea was computed for a 2 year period. Daily water column particulategross production ranged from 5 to 2200 mg C m^–2 day^–1and showed a seasonal pattern similar to chlorophyll a. Budgetcalculation yielded annual gross particulate primary productionsof 124 and 176 g C m^–2 year^–1 in 1995 and 1996,respectively. Annual amounts of phytoplankton respiration, calculatedaccording to a two-compartment model of Langdon [in Li,W.K.W.and Maestrini,S.Y. (eds), Measurement of Primary Productionfrom the Molecular to the Global Scale. International Councilfor the Exploration of the Sea, Copenhagen, 1993, pp. 20–36],and dissolved production in 1996, were both in the range of24–39 g C m^–2 year^–1. Annual total net productionwas thus very similar to particulate gross production (127 and177 g C m^–2 year^–1 in 1995 and 1996, respectively).Phytoplankton growth was low or even negative in winter. Inspring and summer, production/biomass (Pr/B) ratios varied from0.2 up to 1.7. Phytoplankton growth during the growth seasonalways surpassed average flushing time in the area, thus underliningthe potential of local phytoplankton bloom development in thispart of the Wadden Sea. The chlorophyll-specific maximum photosyntheticrate (P^B_max) ranged from 0.8 to 9.9 mg C mg^–1 Chl h^–1and was strongly correlated with water temperature (r² = 0.67).By contrast, there was no clear seasonal cycle in ^B, which rangedfrom 0.007 to 0.039 mg C mg^–1 Chl h^–1 (µmolphotons m^–2 s^–1)^–1. Its variability was muchless than P^B_max and independent of temperature. The magnitudeand part of the variability of P^B_max and ^B are presumably causedby changes in species composition, as evidenced from the rangeof these parameters found among 10 predominant diatom speciesisolated from the Wadden Sea. The ratio of average light conditionsin the water column (E_av) to the light saturation parameterE_k indicates that primary production in the Wadden Sea regionunder study is predominantly controlled by light limitationand that nutrient limitation was likely to occur for a few hoursper day only during 5 (dissolved inorganic nitrogen) to 10 (PO₄,Si) weeks in the 2 year period investigated.     

Effects of the severe winter 1995/96 on the benthic macrofauna of the Wadden Sea and the coastal North Sea near the island of Sylt

The development of benthic macrofauna in the Wadden Sea and in the coastal North Sea after the severe winter of 1995/96 is compared with the preceding years with mild to moderate winters. In the intertidal of the Wadden Sea, ice-drift and low temperature caused the expected changes in species composition by increasing winter mortality in sensitive species, and by exceptionally high recruitment of some species during the succeeding summer. In the shallow subtidal (10–20 m depth), similar winter effects were observed. However, recovery of many subtidal populations was still incomplete until the summer of 1997. It is suggested that this was due to hydrographic conditions that carried many larvae or drifting juveniles into more distant offshore areas. This may have limited larval supply and may have delayed recovery at the onshore sites. Since in the eastern North Sea severe winters are accompanied by frequent easterly winds, it is not clear whether decreasing winter abundances in some species were due to increased mortality, or to a seaward dislocation of organisms. Received in revised form: 7 May 2001 Electronic Publication