Direct evidence supporting Darwin's hypothesis of cross-pollination promoted by sex organ reciprocity.

The floral phenotype plays a main role in the attraction and fit of pollinators. Both perianth traits and the positioning of sex organs can be subjected to natural selection and determine nonrandom mating patterns in populations. In stylar-polymorphic species, the Darwinian hypothesis predicts increased mating success between individuals with sex organs at equivalent heights (i.e. with higher reciprocity). We used paternity analyses in experimental populations of a stylar-dimorphic species. By comparing the observed mating patterns with those expected under random mating, we tested the effects of sex organ reciprocity and perianth traits on mating success. We also analysed phenotypic selection on perianth traits through female and male functions. The (dis)similarity of parental perianth traits had no direct effects on the mating patterns. Sex organ reciprocity had a positive effect on mating success. Narrow floral tubes increased this effect in upper sex organs. Perianth traits showed little signs of phenotypic selection. Female and absolute fitness measures resulted in different patterns of phenotypic selection. We provide precise empirical evidence of the Darwinian hypothesis about the functioning of stylar polymorphisms, demonstrating that mating patterns are determined by sex organ reciprocity and only those perianth traits which are critical to pollinator fit.