Description: 1. Tracer experiments with two diatoms labelled with 13C (Nitzschia palea) and 15N (Fragilaria crotonensis), were conducted to investigate feeding selectivity and interspecific competition between the grazers Asellus aquaticus (Isopoda, Crustacea) and Potamopyrgus antipodarum (Hydrobiidae, Gastropoda). Conventional methods, such as cell counts and estimated biovolume, were used first to detect feeding preferences within the different grazer treatments. 2. The results revealed a significant decline in algal biovolume in all grazer treatments and no indications of active selectivity were observed. In contrast to conventional methods, measurements based on isotope signatures showed strong differences in tracer uptake, thus indicating different degrees of assimilation and digestion by the two grazers. 3. The selectivity index Q, which provides information on the uptake ratio of 13C to 15N, showed a significant time effect for both grazer species and a significant difference between single- and mixed-grazer treatments for P. antipodarum. Thus, this technique enabled the direct quantification of the uptake by grazers and, therefore, served as an ideal tool for the detection of passive selectivity. 4. Our results indicate a shift in feeding preferences related to between-species competition and a potential divergence of trophic niches when species coexist.

Description: Diversity within distinct trophic groups is proposed to increase ecosystem functions such as the productivity of this group and the efficiency of resource use. This proposition has mainly been tested with plant communities, consumer assemblages, and multitrophic microbial assemblages. Very few studies tested how this diversity-productivity relationship varies under different environmental regimes such as disturbances. Coastal benthic assemblages are strongly affected by temporal instability of abiotic conditions. Therefore, we manipulated benthic ciliate species richness in three laboratory experiments with three diversity levels each and analyzed biomass production over time in the presence or absence of a single application of a disturbance (ultraviolet-B [UVB] radiation). In two out of three experiments, a clear positive relationship between diversity and productivity was found, and also the remaining experiment showed a small but nonsignificant effect of diversity. Disturbance significantly reduced the total ciliate biomass, but did not alter the relation between species richness and biomass production. Significant overyielding (i.e., higher production at high diversity) was observed, and additive partitioning indicated that this was caused by niche complementarity between ciliate species. Species-specific contribution to the total biomass varied idiosyncratically with species richness, disturbance, and composition of the community. We thus present evidence for a significant effect of consumer diversity on consumer biomass in a coastal ciliate assemblage, which remained consistent at different disturbance regimes.

Description: Most natural local systems exchange organisms with a regional pool of species through migration and dispersal. Such metacommunity processes of interconnected multispecies assemblages are likely to affect local dynamics of both species and processes. We present results from an artificial marine outdoor rock pool system in which we investigated the factors of (1) local grazer richness and composition, and (2) connectivity of local patches to a regional species pool, and their effects on algal biomass. Local species richness of six grazers was manipulated in both open and closed pools, which were embedded in a regional species pool containing all six grazers. Grazer richness showed significant net biodiversity effects on grazing in the closed, but not in the open, system. Grazer composition, on the other hand, showed significant effects on grazing in both open and closed systems, depending on which species were initially present. The two most efficient grazers were able to compensate for less efficient grazers in species mixtures, hence ensuring the function of grazing. The efficiency of top-down control of algal biomass in open systems thus depends on which particular species are lost. Further, differences in grazing between the open and closed system changed over time due to temporal dynamics in grazer composition. The results emphasize the importance of including system connectivity in experimental designs to allow an extrapolation of biodiversity ecosystem-functioning relationships to natural systems.

Description: Dispersal is a major factor regulating the number of coexisting species, but the relationship between species diversity and ecosystem processes has mainly been analysed for communities closed to dispersal. We experimentally investigated how initial local diversity and dispersal frequency affect local diversity and biomass production in open benthic microalgal metacommunities. Final local species richness and local biomass production were strongly influenced by dispersal frequency but not by initial local diversity. Both final local richness and final local biomass showed a hump-shaped pattern with increasing dispersal frequency, with a maximum at intermediate dispersal frequencies. Consequently, final local biomass increased linearly with increasing final richness. We conclude that the general relationship between richness and ecosystem functioning remains valid in open systems, but the maintenance of ecosystem processes significantly depends on the effects of dispersal on species richness and local interactions.

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: The global decline of biodiversity caused by human domination of ecosystems worldwide is supposed to alter important process rates and state variables in these ecosystems. However, there is considerable debate on the prevalence and importance of biodiversity effects on ecosystem function (BDEF). Here, we argue that much of the debate stems from two major shortcomings. First, most studies do not directly link the traits leading to increased or decreased function to the traits needed for species coexistence and dominance. We argue that implementing a trait-based approach and broadening the perception of diversity to include trait dissimilarity or trait divergence will result in more realistic predictions on the consequences of altered biodiversity. Second, the empirical and theoretical studies do not reflect the complexity of natural ecosystems, which makes it difficult to transfer the results to natural situations of species loss. We review how different aspects of complexity (trophic structure, multifunctionality, spatial or temporal heterogeneity, and spatial population dynamics) alter our perception of BDEF. We propose future research avenues concisely testing whether acknowledging this complexity will strengthen the observed biodiversity effects. Finally, we propose that a major future task is to disentangle biodiversity effects on ecosystem function from direct changes in function due to human alterations of abiotic constraints.