Biodiversity effects on ecosystem functioning can be partitioned into complementarity effects, driven by many species, and selection effects, driven by few. Selection effects occur through interspecific abundance shifts (dominance) and intraspecific shifts in functioning. Complementarity and selection effects are often calculated for biomass, but very rarely for secondary productivity, that is, energy transfer to higher trophic levels. We calculated diversity effects for three functions: aboveground biomass, insect herbivory and pathogen infection, the latter two as proxies for energy transfer to higher trophic levels, in a grassland experiment (PaNDiv) manipulating species richness, functional composition, nitrogen enrichment, and fungicide treatment. Complementarity effects were, on average, positive and selection effects negative for biomass production and pathogen infection and multiple species contributed to diversity effects in mixtures. Diversity effects were, on average, less pronounced for herbivory. Diversity effects for the three functions were not correlated, because different species drove the different effects. Benefits (and costs) from growing in diverse communities, be it reduced herbivore or pathogen damage or increased productivity either due to abundance increases or increased productivity per area were distributed across different plant species, leading to highly variable contributions of single species to effects of diversity on different functions. These results show that different underlying ecological mechanisms can result in similar overall diversity effects across functions.

The ability of an ecosystem to deliver multiple functions at high levels (multifunctionality) typically increases with biodiversity but there is substantial variation in the strength and direction of biodiversity effects, suggesting context-dependency. However, the drivers of this context dependency have not been identified and understood in comparative meta-analyses or experimental studies. To determine how different factors modulate the effect of diversity on multifunctionality, we conducted a large grassland experiment with 216 communities, crossing a manipulation of plant species richness (1-20 species) with manipulations of resource availability (nitrogen enrichment), plant functional composition (gradient in mean specific leaf area [SLA] to manipulate abundances of fast vs. slow species), plant functional diversity (variance in SLA) and enemy abundance (fungal pathogen removal). We measured ten functions, above and belowground, related to productivity, nutrient cycling and energy transfer between trophic levels, and calculated multifunctionality. Plant species richness and functional diversity both increased multifunctionality, but their effects were context dependent. Species richness increased multifunctionality, but only when communities were assembled with fast growing (high SLA) species. This was because slow species were more redundant in their functional effects, whereas fast species tended to promote different functions. Functional diversity also increased multifunctionality but this effect was dampened by nitrogen enrichment, however, unfertilised, functionally diverse communities still delivered more functions than low diversity, fertilised communities. Our study suggests that a shift towards exploitative communities will not only alter ecosystem functioning but also the strength of biodiversity-functioning relationships, which highlights the potentially complex effects of global change on multifunctionality.

Biodiversity affects ecosystem functioning through complementarity effects, driven by many species, and selection effects, driven by few. Selection effects occur through interspecific abundance shifts (dominance) and intraspecific shifts in functioning. Complementarity and selection are often calculated for biomass, but we know little about how diversity affects other ecosystem functions. We calculated diversity effects for aboveground biomass, insect herbivory, pathogen infection and two leaf traits as nutrient cycling proxies, in a grassland experiment (PaNDiv) manipulating species richness, functional composition, nitrogen