Ecological mechanisms of evolution by natural selection: causal processes generating density-and-frequency dependent fitness.

The current theory of natural selection explains that adaptive evolution occurs because genotypes with greater survival or reproductive tendencies, due to their particular biological properties, tend to increase in frequency over the lesser ones in a common environment; therefore, the former will eventually replace the latter. In nature, such a selection process most often occurs in a local population which is nested in a community involving local ecological dynamics which are not clearly articulated in the explanatory scheme of the theory. This paper seeks to explicate such an ecological process giving rise to the evolution of a local population with a particular focus on dynamic effects of an increase in the number of invasive, new types on the fate of old ones. Arguments using the ecological-mechanistic model, representing negative interactions among alternative types of organisms, suggest major ecological mechanisms by which the new replace the old; a selective increase in the number of one type leads to a decrease in the equilibrial abundance of a limiting resource, an increase in the density of conspecifics, and/or an increase in the density of predators, which would in turn lower the per capita reproductive rate, or raise the morality rate of another and make it extinct. Thus, replacement due to selection is associated with such dynamic shifts in equilibria occurring in a local community. The analysis of three (a resource, a prey and a predator) and four species (those plus a top predator) models suggests that evolutionary processes cannot be predicted without reference to the web structure of the community, that some fitness components causing a selective increase in a particular type can have, in some cases, nothing to do with factors causing selective decreases in alternatives, and that evolution of some traits can occur without resource competition.