Does spatiotemporal nutrient variation allow more species to coexist?

Temporal heterogeneity in nutrient availability is known to increase phytoplankton diversity by allowing more species to coexist under different resource niches. Spatial heterogeneity has also been positively correlated with species diversity. Here we investigated how temporal and spatial differences in nutrient addition together impact biodiversity in metacommunities varying in the degree of connectivity among the patches. We used a microcosm experimental design to test two spatiotemporal ways of supplying nutrients: synchronously (nutrients were added regionally-to all four patches at the same time) and asynchronously (nutrients were added locally-to a different patch each time), combined with two different degrees of connectivity among the patches (low or high connectivity). We used three species of algae and one species of cyanobacteria as the primary producers; and five ciliate and two rotifer species as the grazers. We expected higher diversity in metacommunities receiving an asynchronous nutrient supply, assuming stronger development of heterogeneous patches with this condition rather than with synchronous nutrient supply. This result was expected, however, to be dependent on the degree of connectivity among patches. We found significant effects of nutrient addition in both groups of organisms. Phytoplankton diversity increased until the fourth week (transiently) and zooplankton richness was persistently higher under asynchronous nutrient addition. Our results were consistent with our hypothesis that asynchronicity in nutrient supply would create a more favorable condition for species to co-occur. However, this effect was, in part, transient and was not influenced by the degree of connectivity.