Fission as a source of variation for group selection

Without heritable variation natural selection cannot effect evolutionary change. In the case of group selection, there must be variation in the population of groups. Where does this variation come from? One source of variation is from the stochastic birth-death processes that occur within groups. This is where variation between groups comes from in most mathematical models of group selection. Here, we argue that another important source of variation between groups is fission, the (generally random) group-level reproduction where parent groups split into two or more offspring groups. We construct a simple model of the fissioning process with a parameter that controls how much variation is produced among the offspring groups. We then illustrate the effect of that parameter with some examples. In most models of group selection in the literature, no variation is produced during group reproduction events; that is, groups "clone" themselves when they reproduce. Fission is often a more biologically realistic method of group reproduction, and it can significantly increase the efficacy of group selection.