Richness-productivity relationships between trophic levels in a detritus-based system: significance of abundance and trophic linkage.

Most theoretical and empirical studies of productivity-species richness relationships fail to consider linkages among trophic levels. We quantified productivity-richness relationships in detritus-based, water-filled tree-hole communities for two trophic levels: invertebrate consumers and the protozoans on which they feed. By analogy to theory for biomass partitioning among trophic levels, we predicted that consumer control would result in richness of protozoans in the lower trophic level being unaffected by increases in productivity, whereas richness of invertebrate consumers would increase with productivity. Our data were consistent with this prediction: consumer richness increased linearly, but protozoan richness was unrelated to changes in productivity. The productivity-richness relationships for all taxa combined were not necessarily consistent with relationships within each trophic level. We used path analysis to investigate the mechanisms that may produce the observed responses of trophic levels to changes in productivity. We tested the importance of the direct effect of productivity on richness and the indirect effect of productivity mediated by effects on total abundance. For protozoans, only direct effects of productivity on richness were important, but both direct and indirect effects of productivity on richness were important for invertebrates. Protozoan richness was strongly affected by top-down impacts of abundance of invertebrates. These results are consistent with theory on biomass partitioning among trophic levels and suggest a strong link between richness and abundance within and between trophic levels. Understanding how trophic level interactions determine productivity-richness relationships will likely be necessary in order for us to achieve a comprehensive understanding of the determinants of diversity.