Disturbance size and frequency mediate the coexistence of benthic spatial competitors.

Disturbance plays a key role in structuring community dynamics and is central to conservation and natural resource management. However, ecologists continue to debate the importance of disturbance for species coexistence and biodiversity. Such disagreements may arise in part because few studies have examined variation across multiple dimensions of disturbance (e.g., size, frequency) and how the effects of disturbance may depend on species attributes (e.g., competitiveness, dispersal ability). In light of this gap in understanding and accelerating changes to disturbance regimes worldwide, we used spatial population models to explore how disturbance size and frequency interact with species attributes to affect coexistence between seagrass (Zostera marina) and colonial burrowing shrimp (Neotrypaea californiensis) that compete for benthic space in estuaries throughout the west coast of North America. By simulating population dynamics under a range of ecologically relevant disturbance regimes, we discovered that intermediate disturbance (approximately 9-23% of landscape area per year) to short-dispersing, competitively dominant seagrass can foster long-term stable coexistence with broad-dispersing, competitively inferior burrowing shrimp via the spatial storage effect. When holding the total extent of disturbance constant, the individual size and annual frequency of disturbance altered landscape spatial patterns and mediated the dominance and evenness of competitors. Many small disturbances favored short-dispersing seagrass by hastening recolonization, whereas fewer large disturbances benefited rapidly colonizing burrowing shrimp by creating temporary refugia from competition. As a result, large, infrequent disturbances generally improved the strength and stability of coexistence relative to small, frequent disturbances. Regardless of disturbance size or frequency, the dispersal ability of the superior competitor (seagrass), the competitive ability of the inferior competitor (burrowing shrimp), and the reproduction and survival of both species strongly influenced population abundances and coexistence. Our results show that disturbance size and frequency can promote or constrain coexistence by altering the duration of time over which inferior competitors can escape competitive exclusion, particularly when colonization depends on the spatial pattern of disturbance due to dispersal traits. For coastal managers and conservation practitioners, our findings indicate that reducing particularly large disturbances may help conserve globally imperiled seagrass meadows and control burrowing shrimp colonies that can threaten the viability of oyster aquaculture.