Species richness effects on ecosystem multifunctionality depend on evenness, composition and spatial pattern

Recent studies have suggested that the simultaneous maintenance of multiple ecosystem functions (multifunctionality) is positively supported by species richness. However, little is known regarding the relative importance of other community attributes (e.g. spatial pattern, species evenness) as drivers of multifunctionality. We conducted two microcosm experiments using model biological soil crust communities dominated by lichens to: (i) evaluate the joint effects and relative importance of changes in species composition, spatial pattern (clumped and random distribution of lichens), evenness (maximal and low evenness) and richness (from two to eight species) on soil functions related to nutrient cycling (β-glucosidase, urease and acid phosphatase enzymes, in situ N availability, total N, organic C, and N fixation), and (ii) assess how these community attributes affect multifunctionality. Species richness, composition and spatial pattern affected multiple ecosystem functions (e.g. organic C, total N, N availability, β-glucosidase activity), albeit the magnitude and direction of their effects varied with the particular function, experiment and soil depth considered. Changes in species composition had effects on organic C, total N and the activity of β-glucosidase. Significant species richness×evenness and spatial pattern×evenness interactions were found when analysing functions such as organic C, total N and the activity of phosphatase. The probability of sustaining multiple ecosystem functions increased with species richness, but this effect was largely modulated by attributes such as species evenness, composition and spatial pattern. Overall, we found that model communities with high species richness, random spatial pattern and low evenness increased multifunctionality. Synthesis. Our results illustrate how different community attributes have a diverse impact on ecosystem functions related to nutrient cycling, and provide new experimental evidence illustrating the importance of the spatial pattern of organisms on ecosystem functioning. They also indicate that species richness is not the only biotic driver of multifunctionality, and that particular combinations of community attributes may be required to maximize it.