Explaining variation in the effect of floral density on pollinator visitation

Pollinator responses to floral density have important implications for plant biology. In particular, a decline in pollinator visitation at low density can cause an Allee effect (a positive relation of fitness to density) in the plant population, which heightens that population's vulnerability to extinction. Empiricists have reported a variety of relations between flower or plant density and pollinator visitation rates. Here I develop and test a model that provides explanations for this diversity. The model assumes that pollinators distribute themselves between a focal patch of flowers and the surrounding environment so as to maximize foraging success. The resulting relation of per-flower visitation rate to focal-patch floral density is nonlinear, with positive effects at low floral densities and weaker or negative effects at higher densities. The relation is influenced by floral density in the surrounding environment and traits of both the plants and their pollinators. In a field experiment, floral density of Holocarpha virgata ssp. virgata had a nonlinear effect on per-flower visitation that was largely consistent with the model's predictions. By producing testable hypotheses based on biologically reasonable assumptions, this model serves as a starting point for explaining an important facet of plant-pollinator mutualisms.