Pollinator experience, neophobia and the evolution of flowering time.

Environmental changes, such as current climate warming, can exert directional selection on reproductive phenology. In plants, evolution of earlier flowering requires that the individuals bearing genes for early flowering successfully reproduce; for non-selfing, zoophilous species, this means that early flowering individuals must be visited by pollinators. In a laboratory experiment with artificial flowers, we presented captive bumble-bees (Bombus impatiens) with flower arrays representing stages in the phenological progression of a two-species plant community: Bees that had been foraging on flowers of one colour were confronted with increasing numbers of flowers of a second colour. Early flowering individuals of the second 'species' were significantly under-visited, because bees avoided unfamiliar flowers, particularly when these were rare. We incorporated these aspects of bee foraging behaviour (neophobia and positive frequency dependence) in a simulation model of flowering-time evolution for a plant population experiencing selection against late flowering. Unlike simple frequency dependence, a lag in pollinator visitation prevented the plant population from responding to selection and led to declines in population size. Pollinator behaviour thus has the potential to constrain evolutionary adjustments of flowering phenology.