Abstract
Background and Aims Androdioecy, the co-occurrence of males and hermaphrodites, is a rare reproductive system. Males can be maintained if they benefit from a higher male fitness than hermaphrodites, referred to as male advantage. Male advantage can emerge from increased fertility owing to resource reallocation. However, empirical studies usually compare sexual phenotypes over a single flowering season, thus ignoring potential cumulative effects over successive seasons in perennials. In this study, we quantify various components of male fertility advantage, both within and between seasons, in the long-lived perennial shrub Phillyrea angustifolia (Oleaceae). Although, owing to a peculiar diallelic self-incompatibility system and female sterility mutation strictly associated with a breakdown of incompatibility, males do not need fertility advantage to persist in this species, this advantage remains an important determinant of their equilibrium frequency. Methods A survey of >1000 full-sib plants allowed us to compare males and hermaphrodites for several components of male fertility. Individuals were characterized for proxies of pollen production and vegetative growth. By analysing maternal progeny, we compared the siring success of males and hermaphrodites. Finally, using a multistate capture–recapture model we assessed, for each sexual morph, how the intensity of flowering in one year impacts next-year growth and reproduction. Key Results Males benefitted from a greater vegetative growth and flowering intensity. Within one season, males sired twice as many seeds as equidistant, compatible hermaphroditic competitors. In addition, males more often maintained intense flowering over successive years. Finally, investment in male reproductive function appeared to differ between the two incompatibility groups of hermaphrodites. Conclusion Males, by sparing the cost of female reproduction, have a higher flowering frequency and vegetative growth, both of which contribute to male advantage over an individual lifetime. This suggests that studies analysing sexual phenotypes during only single reproductive periods are likely to provide inadequate estimates of male advantage in perennials.