EXPERIMENTAL STUDIES OF COMMUNITY EVOLUTION I: THE RESPONSE TO SELECTION AT THE COMMUNITY LEVEL.

Coevolution generally refers to the process of two or more organisms adapting to each other as a result of individual selection. Another possibility, however, is that coevolution may result from selection acting directly at the community level. Certain types of multispecies associations, such as lichens, which are a symbiotic association between an alga and a fungus, are examples of simple two species communities that may be units of selection. The study presented here uses two species communities of Tribolium castaneum and T. confusum in an investigation of selection acting at the community level. Selection at the community level is performed on one trait measured in one species and correlated responses in other traits measured both within species and among species are monitored. I demonstrate that community selection, defined as the differential survival and or reproduction of communities, can result in significant changes in the phenotype of a community. The observed changes in the phenotype of a community as a result of community selection included changes in the trait under selection (direct effects of selection), as well as changes in traits that are not under selection (correlated responses to selection). Furthermore, two types of correlated responses to selection were observed. The first, within-species correlated responses to selection, are changes in a trait measured in one species as a result of community selection acting on another trait measured in the same species. The second, between-species correlated responses to selection, are changes in a trait measured in one species as a result of community selection acting on a trait measured in another species. Between species correlated responses to selection are of particular interest because they cannot be mediated by pathways of gene action that are internal to an individual, rather they can be mediated only through ecological pathways. In other words, between-species correlated responses to selection suggest that genetically based interactions among individuals are contributing to the response to community selection. These among species ecological pathways of gene action cannot contribute to a response to selection at a lower level; thus community selection may be able to bring about a response to selection that is qualitatively different from the response selection that would occur as a result of selection acting at a lower level.