Boulders are an important substrate for Baltic Sea benthic communities. Although previous studies have examined benthic species on sublittoral boulders in the Baltic Sea, information on benthic assemblages and how they vary in relation to structuring factors is limited. The aim of the present study was to describe the benthic communities, including both algae and animal taxa, on sublittoral boulders in relation to spatial positions on the boulders (4 levels) and differences in wave exposure (2 levels). The multivariate results, including 29 algae and 25 animal taxa, from the field sampling showed a significant interaction between the 2 main factors. Changes in the biomass of both annual an perennial macroalgae as well as Mytilus edulis and Balanus improvisus, were found among different positions on wave-exposed boulders, a pattern which was less clear on boulders from sheltered sites, showing that water movement influences small-scale spatial patterns on sublittoral boulders. Apart from water movement, the physical factors substrate slope and light intensity were also associated with the small-scale patterns found in benthic communities. The difference in these factors may influence both settlement and post-settlement processes for both algae and animals, and possible mechanisms leading to the patterns found are discussed.
KEY WORDS: Spatial patterns · Physical factors · Macroalgae · Mytilus edulis · Balanus improvisusResale or republication not permitted without written consent of the publisher algae and animals on sublittoral boulders (Kautsky 1989, Gilek et al. 2001), but to our knowledge no study has described the community composition or the small-scale patterns of occurrence for the whole macrobenthic community (including both macroalgae and animals) on sublittoral boulders in the Baltic Sea. The aim of the present study was, therefore, to describe the benthic plant and animal community composition, on a small scale, on sublittoral boulders in the Baltic Sea in relation to different positions on boulders and in relation to different degrees of wave exposure.
MATERIALS AND METHODSThe study was carried out in June 2004 in the Askö area, in the northern Baltic proper (Fig. 1). The salinity in this area is relatively constant at around 7. During the winter months the watersurface temperature is ~0°C, while in July to August the temperature may exceed 20°C.On each of 3 wave-exposed (Sites 1 to 3) and 3 sheltered sites (Sites 4 to 6) ( Fig. 1) 3 randomly selected boulders were sampled.All the sites are located in a sparsely populated area. There is no freshwater or sewage water outflow close to any of the sites, and they are homogenous in salinity. The sites were chosen on a marine chart of the area. To determine the degree of wave exposure, a model developed by Isaeus (2004), based on effective fetch, wind direction and wind force, was used. The 3 wave-exposed boulders, which are open to waves from the south (the dominating wind direction), are all homogenous in wave exposure, as are the 3 sheltered s...