Katharina Reichert

Changes in the macrozoobenthos of the intertidal zone at Helgoland (German Bight, North Sea): a survey of 1984 repeated in 2002

Changes in the presence and absence of invertebrates as well as in species conspicuousness were documented in a rocky intertidal community based on surveys in 1984 and 2002. In 2002 six vertically and/or morphologically different stations of an intertidal platform were sampled. Five of these six habitats had already been surveyed in 1984. Replicating precisely the method of the first assessment, presence/absence changes as well as changes in species conspicuousness of 83 invertebrate species were documented, indicating that this intertidal community changed considerably during the 18-year interval. Compared with the study in 1984, 27 species newly appeared, whereas 32 species disappeared. Furthermore, 16 species increased in conspicuousness, whereas eight invertebrates decreased. The total number of species in 2002 was 154 versus 158 in 1984. Although algal species were not recorded as thoroughly as invertebrates, a massive decline in cover of Halidrys siliquosa was noted. Conversely, two invasive algal species became established after 1984, Sargassum muticum (since 1988), a cosmopolitan fucoid alga that prefers shallow subtidal areas for colonization, and Mastocarpus stellatus (introduction in the 1980s) that particularly colonized areas in the mid intertidal. In 1984 the mid intertidal zone was dominated by the brown alga Fucus serratus, whereas in 2002 the blue mussel Mytilus edulis and the periwinkle Littorina littorea were the most conspicuous organisms. Annual mean sea surface temperature (BAH measurements) warmed by 1.1°C over the past four decades. Range-related community shifts, introductions of non-indigenous species and the input of pollutants, are considered to explain long-term ecological changes in the invertebrate community at Helgoland.

Scale-dependent patterns of variability in species assemblages of the rocky intertidal at Helgoland (German Bight, North Sea)

A growing body of literature shows that benthic communities are hierarchically structured on spatial and temporal scales. In two study locations at Helgoland (North Sea), the northern and the western locations, we: (i) investigated the variation in abundance of specific algae and invertebrates at two spatial scales; and (2) evaluated the relationship between elevation and specific species at these scales. We were also interested in using this information about the spatial pattern of individual algae and invertebrates as well as the patterns of elevation to help develop a monitoring programme of the rocky intertidal. We examined the variation of individual algae and invertebrates by means of a hierarchical nested design. Data were taken from five replicates per plot, with plots located in transects (two transects per location). At the northern location, the highest variability in cover of most algae and invertebrates occurred at the scale separated by about so m (scale: transect). This was a direct result of differences between the high- and the low-shore. Most species at high-shore showed a relatively low frequency of occurrence in contrast to a highest frequency of occurence (∼100%) and maximal values of cover at low-shore. However, neither a linear nor a non-linear relationship between elevation and the specific species occurred. At the western location, the highest variability in most macroalgae and invertebrates investigated was among replicates (los of centimetres apart). No relationship between elevation and individual species occurred at this location. Macroalgae at both locations were more consistent over time than invertebrate species. Our results suggest that the relevant processes shaping the individual macroalgae and invertebrates at the Helgoland rocky intertidal vary between locations and the specific species. The potential causes of variation in macroalgal and invertebrate species at different spatial scales are discussed and suggestions for a future monitoring programme are given. Temporal inconsistency in the spatial patterns, and the fact that some individual algae and invertebrates comprising the benthic assemblages vary at different scales, speak in favour of a multiple-scale sampling approach for monitoring change in the intertidal communities at Helgoland.

Community composition of the rocky intertidal at Helgoland (German Bight, North Sea)

At the rocky island of Helgoland (North Sea), the distribution and abundances of intertidal communities were assessed and the effects of wave exposure and tidal height on the spatial distribution patterns of the communities were evaluated. Macroalgae and invertebrates were sampled quantitatively along line transects in three intertidal locations, a semi-exposed, an exposed and a sheltered one. The semi-exposed location was characterised by (1) Ulva spp. at the high intertidal (Ulva-community), (2) mussels and periwinkles at the mid intertidal (Mytilus-community) and (3) Corallina officinalis and mainly the large brown alga Fucus serratus at the low intertidal (Fucus-community). The exposed location encompassed the mid and low intertidal; at both zones the Fucus-community occurred. The sheltered location was characterised by (1) barnacles (Balanus-community) and (2) bryozoans, hydrozoans and mainly the large brown alga Ascophyllum nodosum (Ascophyllum-community). At the semi-exposed, but not at the exposed location the communities changed with the intertidal position. A relationship between wave exposure and the occurrence of specific communities was shown for the sheltered location; in contrast, communities of the semi-exposed and the exposed location appear to be little influenced by wave exposure directly. The community concept and the potential causes of distribution patterns of the defined communities are discussed and suggestions for a future monitoring are given. Variations in the communities at different spatial scales speak in favour of a multiple scale sampling design to monitor changes in the intertidal communities at Helgoland.