The role of secondary pollinators in the evolution of complex colour signals in a bimodal pollination system

Flower colour is recognized as a key trait influencing pollinator visitation and behaviour, evolved to match the sensory system of a particular pollinator group. However, some flowers combine colours associated with different pollinators, suggesting bimodal adaptation. The perennial herb Cyrtanthus obliquus produces red corollas with yellow tips and is pollinated by birds and bees. To assess the role of the two pollinator categories in shaping selection on the bicoloured corollas, we examined bee choices in arrays where the colour signals of plants had been manipulated, and we quantified total as well as bird-mediated phenotypic selection on flower colour in an unmanipulated natural population, using bird and bee visual models. Bees preferred flowers with a yellow signal over all-red flowers. The analysis of phenotypic selection indicated that both birds and bees influence net selection on flower colour, resulting in conflicting selection on the colour contrast between corolla and corolla tip. Our findings show that the optimal phenotype should depend on the relative importance of the two categories of pollinators in a given population and are consistent with bimodal adaptation. This study fills an important gap by showing that secondary pollinators can contribute to selection on complex colour signals in angiosperm flowers.