Does species richness drive community production or vice versa? Reconciling historical and contemporary paradigms in competitive communities

Studies examining the relationship between species richness and the productivity of ecological communities have taken one of two opposite viewpoints, viewing either productivity as a primary driver of richness or richness as a driver of productivity. Recently, verbal and graphical hypotheses have been proposed that attempt to merge these perspectives by clarifying the causal pathways that link resource supply, species richness, resource use, and biomass production. Here we present mathematical models that formalize how these pathways can operate simultaneously in a single ecological system. Using a metacommunity framework in which classic consumer-resource competition theory governs species interactions within patches, we show that the mechanisms by which resource supply influences species richness are inherently linked to the mechanisms by which species richness controls resource use and biomass production. Unlike prior hypotheses, our models show that resource supply can affect species richness and that richness can affect productivity simultaneously at a single spatial scale. Our models also reproduce scale-dependent associations between species richness and community biomass that have been reported elsewhere. By detailing the pathways by which resource supply, species richness, biomass production, and resource use are connected, our models move closer to resolving the nature of causality in diversity-productivity relationships.