Genetic and environmental integration of the hawkmoth pollination syndrome in Ruellia humilis (Acanthaceae).

Background and Aims: The serial homology of floral structures has made it difficult to assess the relative contributions of selection and constraint to floral integration. The interpretation of floral integration may also be clouded by the tacit, but largely untested, assumption that genetic and environmental perturbations affect trait correlations in similar ways. In this study, estimates of both the genetic and environmental correlations between components of the hawkmoth pollination syndrome are presented for chasmogamous flowers of Ruellia humilis , including two levels of control for serial homology. Methods: A greenhouse population for quantitative genetic analysis was generated by a partial diallel cross between field-collected plants. An average of 634 chasmogamous flowers were measured for each of eight floral traits that contribute to the hawkmoth syndrome. Genetic correlations (across parents) and environmental correlations (across replicate flowers) were estimated by restricted maximum likelihood. Key Results: Stigma height, anther height and floral tube length were very tightly integrated in their responses to both genetic and environmental perturbations. The inclusion of floral disc width as a control for serial homology suggests this integration is an adaptive response to correlational selection imposed by pollinators. In contrast, integration of non-homologous traits was low. Furthermore, when comparisons between the dimensions of serially homologous structures were excluded, the genetic and environmental correlation matrices showed little congruence. Conclusions: The results suggest that hawkmoths have imposed strong correlational selection on floral traits involved in the deposition and removal of pollen, and that this is a consequence of stabilizing selection on the relative positions of stigmas and anthers in the face of substantial flower size variation. Low integration of other floral traits, and conflicting patterns of genetic and environmental correlations among these traits, suggest weak or no correlational selection within the range of variability expressed within a population.

Low flower-size variation in bilaterally symmetrical flowers: Support for the pollination precision hypothesis.

PREMISE OF THE STUDY: The evolutionary shift from radial to bilateral symmetry in flowers is generally associated with the evolution of low flower-size variation. This phenomenon supports the hypothesis that the lower size variation in bilateral flowers can be attributed to low pollinator diversity. In this study, we propose two other hypotheses to explain low flower-size variation in bilateral symmetrical flowers. To test the three hypotheses, we examined the relative importance of pollinator diversity, composition, and bilateral symmetry itself as selective forces on low flower-size variation. METHODS: We examined pollinator diversity and composition and flower-size variation for 36 species in a seminatural ecosystem with high bee richness and frequent lepidopteran visitation. KEY RESULTS: Bilateral flowers were more frequently visited than radial flowers by larger bees, but functional-group diversity of the pollinators did not differ between symmetry types. Although bilateral flowers had significantly lower flower-size variation than radial flowers, flower-size variation did not vary with pollinator diversity and composition but was instead related to bilateral symmetry. CONCLUSIONS: Our results suggest that the lower size variation in bilateral flowers might have evolved under selection favoring the control of pollinator behavior on flowers to enhance the accurate placement of pollen on the body of the pollinator, independent of pollinator type. Because of the limited research on this issue, future work should be conducted in various types of plant-pollinator communities worldwide to further clarify the issue.