Do hermaphrodites of gynodioecious Phacelia linearis (Hydrophyllaceae) trade off seed production to attract pollinators?

Abstract

Sexual selection theory predicts that hermaphroditic plants might trade off seed production to attract pollinators. This paper reports a test of this prediction in gynodioecious Phacelia linearis (Hydrophyllaceae), a species in which attractiveness to pollinators increases with corolla diameter. The relationship between corolla diameter and seed production was determined in three natural and three experimental populations. Phenotypic selection analysis was used, with lifetime seed production as a surrogate for fitness. Negative directional selection was expected on hermaphrodite corolla diameter. No directional selection (but possibly stabilizing selection) was expected on corolla diameter in females, which have smaller corollas than hermaphrodites. Shoot biomass and flowering time were included in selection analyses so that the effects of corolla size could be assessed independently of these correlated characters. A parent-offspring study of the quantitative genetics of these characters also was performed. High seed production was associated with large shoot biomass (especially in natural populations) and early flowering (especially in experimental populations). The effect of hermaphrodite corolla size on seed production was not as predicted. Directional selection on hermaphrodite corolla diameter was generally positive. Directional selection on female corolla diameter was relatively weak and variable; stabilizing selection was found in one population. Most of the phenotypic selection on corolla diameter was caused by its (environmental) correlation with shoot biomass. Although these findings do not support the hypothesis that hermaphrodites trade off seed production to attract pollinators, a cautious interpretation is warranted. In one natural population, insect herbivory generated positive covariance between hermaphrodite corolla diameter and seed number, leading to a spurious estimate of positive directional selection for corolla diameter. Other, undetected sources of positive covariance between corolla size and seed production also might have obscured the expected tradeoff. Corolla diameter was found to be heritable, but shoot biomass and flowering time were not. The genetic correlation between hermaphrodite corolla diameter and female corolla diameter was positive and significant, but it was significantly less than 1, so corolla size could evolve independently in the two genders.