Description: An experimental laboratory set-up was used to study the influence of different grain size compositions and temperatures on the growth of benthic cyanobacteria and diatoms, and on the competition between these 2 groups. Monospecific cultures of 3 species of cyanobacteria (Merismopedia punctata, Microcoleus chthonoplastes, Oscillatoria limosa), and of 2 species of benthic diatoms (Phaeodactylum tricornutum and Nitzschia sp.) were used. The organisms were cultured in 100 ml flasks filled with medium and 3 different kinds of sediment: (1) Sand (fine sand, 63 to 200 µm), (2) Mud-I (mixed fine sand and mud 〈63 µm in the ratio 80:20 wt %), (3) Mud-II (mixed fine sand and mud in the ratio 50:50 wt %). Experimental temperatures were 10, 15 and 25°C. At 10°C and 15°C, both diatom species achieved the highest biomass on the sediments of the finest grain size (50 wt % 〈 63 µm) while cyanobacteria achieved low biomass levels. Coarsening of sediments at the same temperature levels revealed a gradually lower biomass of the diatoms. Particularly on sand, the diatoms never reached the same concentrations of chlorophyll a as on mud. The cyanobacteria, on the other hand, had the highest biomass on sand at 15°C. In the competition experiments the benthic diatom species Nitzschia sp. dominated all types of sediments at 10°C and 15°C. The experiments at 25°C were dominated by the filamentous cyanobacterium M. chthonoplastes. This indicates the importance of abiotic conditions for the distribution and abundance of benthic phototrophic micro-organisms.

Description: The response of epilithic microphytobenthos to in situ nutrient enrichments was studied in the Kiel Fjord, Western Baltic Sea. For this purpose an experimental setup with continuous nutrient supply was designed and installed. Experiments followed the colonization of artificial substrates and the responses of benthic algae to different concentrations and combinations of nitrogen and phosphate. They revealed nitrogen limitation of epilithic microflora from late spring to autumn, such that there was higher biovolume with increasing nitrogen concentrations. Diatoms became dominant in all experiments except one in which the rhodophyte Ceramium strictum prevailed. Species composition was altered by nutrient treatments; one to several species were highly favoured by nutrient enrichment. Consequently, diversity was negatively correlated with final yield. These findings support the hypothesis that competition is an important factor structuring microphytobenthic communities.

Description: Generalisations on the combined effects of consumers and resources on autotrophs in aquatic food webs largely rely on freshwater studies. In this study, we tested these general concepts with marine benthic microalgae, which are important components of coastal food webs. We manipulated nitrogen availability and herbivore presence in a factorial field experiment in the Western Baltic Sea. Moreover, we investigated how herbivore control varied among 3 sites and 2 seasons and tested for trophic cascades by enhancing demersal fish density at 2 sites. Nitrogen availability and herbivore presence had strong and antagonistic effects on microalgal biomass, species composition and diversity. Herbivores significantly reduced algal biomass, whereas nutrient enrichment led to an increase in biomass. Herbivore effects on microalgal biomass increased with increasing nitrogen availability, indicating a functional response of herbivores to nutrient enrichment. The response of microalgae at the species level suggested a trade-off between nutrient use and grazing resistance which appeared to be linked to algal growth form. Compared to other growth forms, large erect species were most responsive to both nitrogen loading and herbivory. Grazing reduced microalgal diversity at low nutrient supply, but enhanced it at high nutrient supply. Herbivore effects varied considerably among different sites and were stronger in spring than in summer. Manipulations of fish density during summer did not have any effects on microalgal community structure. In conclusion, our results demonstrate that herbivores and nutrients have strong and balancing effects on marine microbenthic community structure.

Description: Four in situ experiments were conducted to examine the potential top-down and bottom-up control of epibenthic ciliate communities. The experiments were run in the littoral of Lake Erken and at a brackish water site on the island of Väddö on the Baltic coast of Sweden, during the spring of 2000. The experimental manipulations were the presence/absence of the natural macrozoobenthos grazer community, cross-classified with the presence/absence of additional nutrients. Epibenthic ciliates responded to both manipulation of grazers and resources, but the response was group specific. Total ciliate abundance decreased when macrozoobenthos (largely chironomids, gastropods, trichopteran larvae, isopods and amphipods) were removed, thus excluding a direct predation effect of the macrozoobenthos community on ciliates. Total ciliate biomass, but not abundance, tended to increase in the presence of additional nutrients; an effect weakly dependent on season and site. The disparity between effects of nutrients on biomass and abundance was due to effects on heterotrichs, a group of large but relatively rare algivorous ciliates. The manipulations altered the ciliate community composition, and between lakes there were differences in species richness and diversity and experiments. However, neither the removal of macrozoobenthos nor the addition of nutrients changed species richness or diversity. This runs counter to work with other taxonomic groups, which shows maximal diversity at an intermediate level of resources or predation. This can only be partially explained by the lack of direct predation effects and the open nature of the experimental system.

Description: Cellular nutrient ratios are often applied as indicators of nutrient limitation in phytoplankton studies, especially the so-called Redfield ratio. For periphyton, similar data are scarce. We investigated the changes in cellular C: N: P stoichiometry of benthic microalgae in response to different levels and types of nutrient limitation and a variety of abiotic conditions in laboratory experiments with natural inocula. C: N ratios increased with decreasing growth rate, irrespective of the limiting nutrient. At the highest growth rates, the C: N ratio ranged uniformly around 7.5. N: P ratios 〈13 indicated N limitation, while N: P ratios 〉22 indicated P limitation. Under P limitation, the C: P ratios increased at low growth rate and varied around 130 at highest growth rates. For a medium with balanced supply of N and P, an optimal stoichiometric ratio of C: N: P = 119 : 17 : 1 could be deduced for benthic microalgae, which is slightly higher than the Redfield ratio (106 : 16 : 1) considered typical for optimally growing phytoplankton. The optimal ratio was stable against changes in abiotic conditions. In conclusion, cellular nutrient ratios are proposed as an indicator for nutrient status in periphyton.

Description: Ecological stoichiometry can be a powerful tool to understand food web consequences of altered biogeochemical cycles as well as consequences of biodiversity loss on biogeochemistry and has proved to be a suitable framework to predict effects of consumers on the nutrient content of their prey. However, predictions from ecological stoichiometry have mainly been tested using single consumer species, whereas in most natural ecosystems several consumer species coexist. We conducted 2 outdoor mesocosm experiments with marine rock pool communities to test whether species richness and species combination of benthic invertebrates affected the nutrient content of periphyton. We independently manipulated 12 different consumer combinations ranging from 0 to 6 (2004) or 0 to 4 (2005) grazer species and measured the biomass and nutrient content of the algae. Grazers included 3 gastropods and 3 crustaceans. In 2005, we additionally analyzed animal nutrient content and N excretion rate. Algal biomass and C:N ratios decreased in the presence of grazers in both years, indicating that the remaining algae had higher internal N content. Also, both biomass (2004 and 2005) and C:N ratios (only 2004) decreased even further when grazer richness increased. In 2004, significant net diversity effects of grazer richness on periphyton C:N ratios indicated that periphyton N content under multispecies grazing could not be predicted from the effect of single species. In 2005, significant net diversity effects on C:N ratios were rare, but periphyton C:N ratios consistently decreased with increasing grazer excretion rate, indicating that higher nitrogen regeneration by grazers led to higher N incorporation by algae. The effects of species richness were mainly affected by the presence of one efficient grazer, the gastropod Littorina littorea. Our experiments indicate that non-additive intraguild interactions may qualitatively alter the stoichiometric effects of multispecies consumer assemblages.

Description: Latitudinal declines of species richness from the tropics to the poles represent a general spatial pattern of diversity on land. For the marine realm, the generality of this pattern has frequently been questioned. Here, I use a database with nearly 600 published gradients (198 of which were marine) to assess whether there is a marine latitudinal diversity gradient of similar average strength and slope as that for terrestrial organisms. Using meta-analysis techniques, I also tested which characteristics of organisms or habitats affected gradient strength and slope. The overall strength and slope of the gradient for marine organisms was significantly negative and of similar magnitude compared to gradients for terrestrial organisms. Marine gradients were on average stronger as well as steeper than freshwater gradients. Latitudinal gradients were clearly a regional phenomenon, with stronger gradients and steeper slopes for diversity assessed on regional than on local scales. The gradient parameters differed also between oceans and between different habitats, with steeper gradients related to the pelagial rather than the benthos. There were on the other hand no significant differences between hemispheres and between different gradient ranges, although such differences have often been presumed. The most important organismal characteristic related to gradient structure was body mass, with significant gradients related to large organisms. A significant increase in gradient strength with increasing trophic level was observed. The meta-analysis also revealed strongest gradients for nekton and mobile epifauna, whereas the gradients were weak for sessile epifauna and for infauna. In conclusion, marine biota reveal a similar overall decline in diversity with latitude to that observed in terrestrial realms, but the strength and slope of the gradient are clearly subject to regional, habitat and organismal features.