Trophic dynamics of a simple model ecosystem.

We have constructed a model of community dynamics that is simple enough to enumerate all possible food webs, yet complex enough to represent a wide range of ecological processes. We use the transition matrix to predict the outcome of succession and then investigate how the transition probabilities are governed by resource supply and immigration. Low-input regimes lead to simple communities whereas trophically complex communities develop when there is an adequate supply of both resources and immigrants. Our interpretation of trophic dynamics in complex communities hinges on a new principle of mutual replenishment, defined as the reciprocal alternation of state in a pair of communities linked by the invasion and extinction of a shared species. Such neutral couples are the outcome of succession under local dispersal and imply that food webs will often be made up of suites of trophically equivalent species. When immigrants arrive from an external pool of fixed composition a similar principle predicts a dynamic core of webs constituting a neutral interchange network, although communities may express an extensive range of other webs whose membership is only in part predictable. The food web is not in general predictable from whole-community properties such as productivity or stability, although it may profoundly influence these properties.