The maintenance of outcrossing in predominantly selfing species: Ideas and evidence from cleistogamous species

Cleistogamous species present strong evidence for the stability of mixed mating, but are generally not considered in this context. Individuals of cleistogamous species produce both obligately selfing cleistogamous flowers (CL) and potentially outcrossed chasmogamous flowers (CH) with distinct morphologies. Greater energetic economy and reliability of CL relative to CH suggest that forces that maintain selection for outcrossing may be stronger in these species than in mixed maters with monomorphic flowers. We reviewed data from 60 studies of cleistogamous species to evaluate proposed explanations for the evolutionary stability of mixed cleistogamous and chasmogamous reproduction and to quantify the magnitude of selection necessary to account for the maintenance of CH. We found circumstantial support for existing hypotheses for the stability of cleistogamy, and that forces that maintain CH must account for a 15-342% advantage of reproduction via CL. We suggest that heterosis and the effects of mass action pollination should be considered.