Timing of disturbance alters competitive outcomes and mechanisms of coexistence in an annual plant model

Ecological disturbance is inherently a multi-faceted phenomenon; disturbance events can differ in distinct quantifiable aspects, such as intensity, duration, spatial extent, and time since last disturbance. Though effects of disturbance timing (specifically, time within a season) have been investigated empirically, theoretical work is lacking, in part because effects of disturbance may depend on the timing relative to the life cycle of the species in question. To demonstrate a theoretical basis for the effects of timing, we develop a model of annual plants subject to soil disturbance. We show that timing of disturbance can have significant effects on community composition. In addition, we quantify the mechanisms of coexistence acting under different timing regimes and show that differences in timing lead to different coexistence mechanisms. Specifically, we find that early disturbance (which enhances germination from the seed bank) generates the storage effect, whereas coexistence under late disturbance (which reduces adult fecundity and contributions to the soil seed bank) depends more on relative nonlinearity of competition. We discuss these two distinct mechanisms within the context of the underlying ecological processes, and we also briefly consider the broader implications of our analyses for disturbance timing in real communities. Our findings extend ecological disturbance theory by linking timing to specific competitive outcomes and can be applied to a wide range of disturbance-prone communities. Because we identify the underlying mechanisms resulting from different disturbance timings, our results can potentially inform theory for conservation and invasive species management practice.