Ecosystem flux and biotic modification as drivers of metaecosystem dynamics

The fluxes of energy, matter, and organisms are important structuring forces of metaecosystems. Such ecosystem fluxes likely interact with environmental heterogeneity and differentially affect the diversity of multiple communities. In an aquatic mesocosm experiment, we tested how ecosystem flux and patch heterogeneity affected the diversity of bacteria, phytoplankton, and zooplankton metacommunities, and the structure and functioning of metaecosystems. We built metaecosystems consisting of three mesocosms that were either connected by flux of living organisms, organic material, and nutrients (alive ecosystem flux) or only by flux of organic material and nutrients (dead ecosystem flux). The three patches of each metaecosystem were either homogeneous or heterogeneous in nutrient loading. We found that the three groups of organisms responded differently to our treatments: flux of living organisms increased bacterial diversity irrespective of nutrient heterogeneity, while flux effects on phytoplankton diversity depended on nutrient heterogeneity, potentially indicating source-sink effects. Although zooplankton diversity was largely unaffected by our manipulations, subtle changes of community composition in response to ecosystem flux had strong effects on lower trophic levels, highlighting the importance of indirect flux effects via alterations in trophic interactions. Furthermore, differential effects of communities on the mean and spatial variability of local abiotic environments influenced the development of metaecosystem heterogeneity through time. Despite identical nutrient loading at the scale of the metaecosystem, abiotic conditions diverged between homogeneous and heterogeneous metaecosystems. For example, concentrations in dissolved organic carbon (DOC) were higher in homogeneous than heterogeneous metaecosystems, possibly because of differential responses of the algal community to local environmental conditions. Similarly, we found that flux effects on organisms translated into effects on DOC concentrations at the patch level, suggesting that flux-mediated changes in abundances of species can alter abiotic conditions. Our study shows that the dynamics of biotic and abiotic compartments of spatially structured ecosystems are intricately linked, highlighting the importance of integrating metacommunity and metaecosystem perspectives.