Selection for restraint in competitive ability in spatial competition systems.

The absence of 'super competitors' in nature is usually attributed to organisms facing trade-offs in resource allocation. Here we identify another mechanism, dependent on indirect interactions among species and non-random spatial organization, in which selection favours restraint in competitive ability. In simple spatial models of a three-species intransitive network, indirect interactions favour slower growth and selection limits the difference in growth rate among species. The mechanism involves a trade-off between selection at the individual level, which selects for increased growth rate, and at the community level, which acts to limit growth rate to less than the maximum possible. If the difference in growth rates among species becomes too large, then the community becomes unstable and collapses to a monoculture of the slowest growing species. The mechanism requires both the intransitive network structure and self-organized spatial structure in the system. Similar behaviours arise in more complex systems of more than three species, and where there are reversals in interaction outcomes between species pairs. The work suggests that spatial self-structuring, indirect interactions and selection acting on community properties can be important in evolution. It provides a partial explanation of the high level of species coexistence and apparent restraint in interspecific interactions evident in some assemblages of sessile marine colonial organisms.