How the storage effect and the number of temporal niches affect biodiversity in stochastic and seasonal environments

Temporal environmental variations affect diversity in communities of competing populations. In particular, the covariance between competition and environment is known to facilitate invasions of rare species via the storage effect. Here we present a quantitative study of the effects of temporal variations in two-species and in diverse communities. Four scenarios are compared: environmental variations may be either periodic (seasonal) or stochastic, and the dynamics may support the storage effect (global competition) or not (local competition). In two-species communities, coexistence is quantified via the mean time to absorption, and we show that stochastic variations yield shorter persistence time because they allow for rare sequences of bad years. In diverse communities, where the steady-state reflects a colonization-extinction equilibrium, the actual number of temporal niches is shown to play a crucial role. When this number is large, the same trends hold: storage effect and periodic variations increase both species richness and the evenness of the community. Surprisingly, when the number of temporal niches is small global competition acts to decrease species richness and evenness, as it focuses the competition to specific periods, thus increasing the effective fitness differences.Author summary: One of the major challenges of community ecology and population genetics is the understanding of the factors that protect biodiversity. Surprisingly, in many generic cases temporal environmental variations (and the abundance fluctuations associated with it) promote the coexistence of competing species and facilitate genetic polymorphism. Here we present a detailed and quantitative comparison between the stabilizing (and the destabilizing) effects of periodic (seasonal) and stochastic temporal variations. When the number of species is small, we show that persistence times under periodic variations are much longer than the persistence times in a stochastic environment. However, environmental variations facilitate coexistence only when the number of temporal niches is larger than the number of species, whereas in the opposite case the same mechanism acts to increase competition and to decrease species richness. Since it is reasonable to expect the number of temporal niches under seasonal variations to be typically smaller than the corresponding number in stochastic environments, stochastic variations provide a more plausible explanation for the apparent stability of high-diversity assemblages.