Description: Recent experiments, mainly in terrestrial environments, have provided evidence of the functional importance of biodiversity to ecosystem processes and properties. Compared to terrestrial systems, aquatic ecosystems are characterised by greater propagule and material exchange, often steeper physical and chemical gradients, more rapid biological processes and, in marine systems, higher metazoan phylogenetic diversity. These characteristics limit the potential to transfer conclusions derived from terrestrial experiments to aquatic ecosystems whilst at the same time provide opportunities for testing the general validity of hypotheses about effects of biodiversity on ecosystem functioning. Here, we focus on a number of unique features of aquatic experimental systems, propose an expansion to the scope of diversity facets to be considered when assessing the functional consequences of changes in biodiversity and outline a hierarchical classification scheme of ecosystem functions and their corresponding response variables. We then briefly highlight some recent controversial and newly emerging issues relating to biodiversity-ecosystem functioning relationships. Based on lessons learnt from previous experimental and theoretical work, we finally present four novel experimental designs to address largely unresolved questions about biodiversity-ecosystem functioning relationships. These include (1) investigating the effects of non-random species loss through the manipulation of the order and magnitude of such loss using dilution experiments; (2) combining factorial manipulation of diversity in interconnected habitat patches to test the additivity of ecosystem functioning between habitats; (3) disentangling the impact of local processes from the effect of ecosystem openness via factorial manipulation of the rate of recruitment and biodiversity within patches and within an available propagule pool; and (4) addressing how non-random species extinction following sequential exposure to different stressors may affect ecosystem functioning. Implementing these kinds of experimental designs in a variety of systems will, we believe, shift the focus of investigations from a species richness-centred approach to a broader consideration of the multifarious aspects of biodiversity that may well be critical to understanding effects of biodiversity changes on overall ecosystem functioning and to identifying some of the potential underlying mechanisms involved.

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: The decline of biodiversity with latitude has received great attention, but both the concise pattern and the causes of the gradient are under strong debate. Most studies of the latitudinal gradient comprise only one or few organism types and are often restricted to certain region or habitat types. To test for significant variation in the gradient between organisms, habitats, or regions, a meta-analysis was conducted on nearly 600 latitudinal gradients assembled from the literature. Each gradient was characterized by two effect sizes, strength (correlation coefficient) and slope, and additionally by 14 variables describing organisms, habitats, and regions. The analysis corroborated the high generality of the latitudinal diversity decline. Gradients on regional scales were significantly stronger and steeper than on local scales, and slopes also varied with sampling grain. Both strength and slope increased with organism body mass, and strength increased with trophic level. The body mass-effect size relation varied for ecto- versus homeotherm organisms and for different dispersal types, suggesting allometric effects on energy use and dispersal ability as possible mechanisms for the body mass effect. Latitudinal gradients were weaker and less steep in freshwater than in marine or terrestrial environments and differed significantly between continents and habitat types. The gradient parameters were not affected by hemisphere or the latitudinal range covered. This analysis is the first to describe these general and significant patterns, which have important consequences for models aiming to explain the latitudinal gradient.

Description: In this article we review the indirect interactions prevailing in littoral communities and compare their importance to direct trophic interactions. We focus on the interaction between benthic algae and their herbivorous consumers, i. e. invertebrate grazers and herbivorous fish. In addition to the effects of grazers on periphyton biomass and production, we categorise and describe the types of indirect interactions observed and we present a quantitative meta-analysis to compare the effect magnitude of indirect and direct effects. Our review shows that herbivore-plant interactions comprise considerable complexity beyond consumption. We identify and review three different indirect interactions, which play important roles for periphyton-grazer interactions in benthic food webs. These are keystone predation, trophic cascading and habitat facilitation. Direct negative effects were stronger than indirect positive effect, because grazing leads in almost all studies to a decrease in algal biomass. We found positive effects of grazing on algal nutrient content, biomass-specific productivity and evenness. We found negative overall effects on species richness and area-specific productivity. Moreover, spatial heterogeneity was on average increased by grazing. The magnitudes of direct and indirect effects were often correlated

Description: Ecological stoichiometry describes the biochemical constraints of trophic interactions emerging from the different nutrient content and nutrient demand of producers and consumers, respectively. Most research on this topic originates from well-mixed pelagic food webs, whereas the idea has received far less attention in spatially structured habitats. Here, we test how light as well as grazing and nutrient regeneration by consumers affects growth and biomass of benthic primary producers. In the first laboratory experiment, we manipulated grazer presence (two different snail species plus ungrazed control), in the second experiment we factorially combined manipulation of grazer presence and light intensity. We monitored snail and periphyton biomass as well as dissolved and particulate nutrients (nitrogen and phosphorus) over time. Grazers significantly reduced algal biomass in both experiments. Grazers affected periphyton nutrient content depending on the prevailing nutrient limitation and their own body stoichiometry. In the nitrogen (N-) limited first experiment, grazers increased N both in the periphyton and in the water column. The effect was stronger for grazers with lower N-content. In the phosphorus (P-) limited second experiment, grazers increased the P-content of the periphyton, but the grazer with lower N-content had additionally positive effects on algal N. Light reduction did not affect periphyton biomass, but increased chlorophyll-, N- and P-content of the periphyton. These experiments revealed that the indirect effects of grazers on periphyton were bound by stoichiometric constraints of nutrient incorporation and excretion.

Description: In periphyton communities, autotrophic algae and prokaryotes live in close spatial proximity to heterotrophic components such as bacteria and micro- and meiofauna. In factorial field experiments, we manipulated grazer access and nutrient supply to periphyton communities and measured the effects on algal, ciliate, meiofaunal, and bacterial biomass. We tested whether grazing macrozoobenthos affects all periphytic components (generalist consumption), whether nutrient effects propagate through the community, and whether interactions between the different periphyton groups allow for indirect feedback mechanisms. The experiments were conducted during three different seasons in a meso-eutrophic lake in Sweden (Lake Erken) and at an adjacent coastal marine site (Väddö) of similar productivity, but with contrasting grazer fauna. We found strong direct effects of nutrients and grazing on algae at both sites. Algal biomass increased in fertilized treatments and was significantly reduced when grazers were present. The algae clearly dominated the system quantitatively and were positively correlated to the biomass of ciliates and meiofauna. The effects of grazing and nutrients were more complex for heterotrophs than for algae. Generally, the presence of grazers tended to increase the biomass of bacteria, ciliates, and meiofauna. Thus, macrograzers were not generalist consumers of the entire community, but mainly reduced algae. Furthermore, the results suggested strong indirect effects of grazing, presumably through changes in nutrient supply and algal size structure. Nutrient enrichment had weak and inconsistent effects on bacterial, ciliate, and meiofaunal biomass. There was thus no complete propagation of bottom-up effects through the community, and strong internal feedback mechanisms within the periphyton mediated the effects of macroconsumers and nutrient enrichment.

Description: A key question in ecology is which factors control species diversity in a community1, 2, 3. Two largely separate groups of ecologists have emphasized the importance of productivity or resource supply, and consumers or physical disturbance, respectively. These variables show unimodal relationships with diversity when manipulated in isolation4, 5, 6, 7, 8. Recent multivariate models9, 10, however, predict that these factors interact, such that the disturbance–diversity relationship depends on productivity, and vice versa. We tested these models in marine food webs, using field manipulations of nutrient resources and consumer pressure on rocky shores of contrasting productivity. Here we show that the effects of consumers and nutrients on diversity consistently depend on each other, and that the direction of their effects and peak diversity shift between sites of low and high productivity. Factorial meta-analysis of published experiments confirms these results across widely varying aquatic communities. Furthermore, our experiments demonstrate that these patterns extend to important ecosystem functions such as carbon storage and nitrogen retention. This suggests that human impacts on nutrient supply11 and food-web structure12, 13 have strong and interdependent effects on species diversity and ecosystem functioning, and must therefore be managed together.

Description: In most groups of organisms, the species richness decreases from the tropics to the poles. The mechanisms causing this latitudinal diversity gradient are still controversial. We present data from a comprehensive weighted meta-analysis on the strength of the latitudinal gradient in relation to body size. We sampled literature data on the correlation between species richness and latitude for a variety of organisms, ranging from trees to protozoa. In addition, own data on the presence of large-scale diversity patterns for diatoms were included, both for local and regional species richness. The strength of the latitudinal gradient was positively correlated to the size of the organisms. Strongest decreases of species richness to the poles was found for large organisms like trees and vertebrates, whereas meiofauna, protozoa and diatoms showed weak or no correlations between species richness and latitude. These results imply that latitudinal gradients are shaped by non-equilibrium (regional) processes and are persistent under conditions of dispersal limitation.