The situation regarding the distribution and abundance of seagrass, macroalgae and benthic fauna near the island of Sylt in the south-eastern North Sea during the period 1923 to 1940 is compared with that of the 1980s. Evidence of organic enrichment in recent times is provided by (1) massive growth of green algal mats on sheltered tidal flats, (2) a decline of red algae in the subtidal zone, (3) an expansion of mussel beds along low water line and down to 20 m depth, (4) increased abundance of polychaetes inhabiting intertidal and subtidal sandy bottoms. Seagrass beds have undergone complex changes which remain unexplained. Intensified erosion has contributed to the loss of habitats in the intertidal zone, and probably affected sessile epifauna in the deep channels. Here, direct removal and disturbance by the bottom-trawling fishery may also have contributed to the observed species impoverishment.
Intertidal sediments of K6nigshafen (Island of Sylt, North Sea) were sieved for mesofauna (> 0.25 ram) and macrofauna (> 1 ram) in spring and autumn 1990. Although sediments are coarser than in other parts of the Wadden Sea, the macrobenthic fauna was very similar but with a tendency towards higher species density, abundance and biomass. Taking into account the areal size of sandy flats, seagrass beds, mud flats and mussel beds, the average biomass is calculated to be 65 g ash-free dry weight m -2. The lugworm Arenicola marina dominates the biomass (28%), followed by the bivalves Mytilus edulis (21%), Mya arenaria (16 %), Cerastoderma edule (10 %) and the mudsnafl Hydrobia ulvae (9 %). While spring and autumn biomass are almost alike, abundance is highly variable and entirely dominated by H. ulvae. Mesofauna is mainly composed of ohgochaetes, small and juvenile polychaetes. Abundance is similar to that of macrofauna, while biomass is only about 1 g m -2. Macrophyte biomass amounted to 9 % of that of macrofauna. In the course of the century, mussel beds expanded while muddy areas declined. The concomitant effects on biomass presumably compensated each other.
Where, since the 1980s, patchy and variable green algal mats are prevailing, distinct belts of an amphipod (Corophium volutator) and seagrass (Zostera spp.) had dominated in the 1930s. The zonation between tide marks has been mapped in a sheltered sedimentary bay in the Wadden Sea near the island of Sylt (coastal eastern North Sea). Maps on vegetation from 1924 and on selected macrobenthos from 1932 and 1934 are compared with biannual surveys conducted from 1988 to 2006. Rising high water levels and eutrophication are suggested to be major causes of the observed long-term changes. In front of a saltmarsh, a sandy beach developed and partly displaced former cyanobacterial mats. Advancing sandiness may have inhibited C. volutator and facilitated lugworms, Arenicola marina, in the upper tidal zone. A variable occurrence of green algal mats arising in the 1980s aVected infauna and seagrass by smothering the biota underneath. This dissolved a coherent belt of Zostera noltii. In the lower tidal zone, natural disturbances had lasting eVects on the occurrence of mussels with attached fucoid algae. The spectrum of species became enriched by alien species (13% of macrobenthic taxa). A reversal to habitat structure and biotic zonation of the 1920-1930s does not seem possible. Aliens, in combination with climate change, are expected to further divert the ecological pattern to new conWgurations.
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.
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