title: Frequency-dependent pollinator discrimination acts against female plants in the gynodioecious Geranium maculatum.
abstract: OBJECTIVE
  Gynodioecy, the co-occurrence of female and hermaphroditic individuals, is thought to be an intermediate step between hermaphroditism and separate sexes, a major transition in flowering plants. Because retaining females in a population requires that they have increased seed fitness (to compensate for the lack of pollen fitness), factors that affect seed fitness are of great importance to the evolution of this mating system and have often been studied. However, factors negatively affecting female fitness are equally important and have been largely neglected. One such factor stems from female flowers being less attractive to insects than hermaphrodite flowers, thereby decreasing their relative fitness.
  
  
  METHODS
  To test the severity and consequences of this type of pollinator discrimination in Geranium maculatum, experimental populations with the range of sex ratios observed in nature were created, ranging from 13 % to 42 % females. Pollinators were observed in order to measure the strength of discrimination, and pollen deposition and seed production of both sexes were measured to determine the fitness consequences of this discrimination. Additionally a comparison was made across the sex ratios to determine whether discrimination was frequency-dependent.
  
  
  RESULTS
  It was found that female flowers, on average, were visited at half of the rate of hermaphrodite flowers, which decreased their pollen receipt and seed production. Additionally, females were most discriminated against when rare, due to both changes in the pollinators' behaviour and a shift in pollinator composition.
  
  
  CONCLUSIONS
  The results suggest that pollinator discrimination negatively affects females' relative fitness when they are rare. Thus, the initial spread of females in a population, the first step in the evolution of gynodioecy, may be made more difficult due to pollinator discrimination.
authors.affiliation: Institute of Agriculture and Environment, Massey University, Palmerston North, New Zealand mvanette@umich.edu.
authors.forename: Megan L
authors.initials: ML
authors.lastname: Van Etten
authors.affiliation: Plant Biology Department, University of Georgia, Athens, GA 30602, USA.
authors.forename: Shu-Mei
authors.initials: SM
authors.lastname: Chang
mesh_terms: D000818:Animals; D001516:Bees / Q000502:physiology*; D031315:Geranium / Q000033:anatomy & histology / Q000502:physiology*; D056771:Ovule / Q000033:anatomy & histology / Q000502:physiology*; D011058:Pollen / Q000502:physiology; D054817:Pollination / Q000502:physiology*; D012098:Reproduction; D012744:Sex Ratio
keywords: Geraniaceae; Geranium maculatum; frequency-dependent fitness; gynodioecy; minority disadvantage hypothesis; plant mating system; plant–pollinator interaction; pollinator discrimination