When does local spatial structure hinder competitive coexistence and reverse competitive hierarchies?

Classical theory states that if conspecifics have a greater competitive effect on individuals than heterospecifics then coexistence should occur, and ecologists have spent much effort exploring ways to generate coexistence when this condition is not met. One process that has received particular attention in the last two decades is the effect of within-species aggregation and between-species segregation caused by limited dispersal. A number of theories have emerged as to how this common spatial pattern may help maintain biodiversity, and the general conclusion that has emerged is that spatial structure should almost always help competitors to coexist. But does spatial structure really always aid biodiversity? An individual-based model based on a spatial extension to the Lotka-Volterra competition equations and its mathematical approximation are presented to determine how local spatial structure may affect communities in which there is strong niche differentiation. Two main results emerge from analyses of the models. First, intraspecific competition being greater than interspecific competition coexistence may no longer be sufficient to generate coexistence when spatial structure is strong; and the species with the highest intraspecific competition coefficient is likely to be excluded. Second, dominance hierarchies may be reversed so that a competitor may become the subordinate species when dispersal and competitive interactions occur over short spatial scales. Both results emerge because, even though a species may be globally rare, intense clumping means most interactions occur between conspecifics, and if this is very intense it may be sufficient to stop a species from invading. However, long-range dispersal may ameliorate these effects by reducing the frequency of conspecific interactions, and this is especially important when a species is rare since it is very likely to land in an area dominated by heterospecifics. These results are most relevant to sessile organisms that produce relatively few viable offspring that survive to adulthood and that have relatively weak dispersal. The conclusion is that within-species aggregation may hinder coexistence when the toughest competitor an individual is likely to face is a member of its own species.