Phenotypic gender in Hormathophylla spinosa (Brassicaceae), a perfect hermaphrodite with tetradynamous flowers, is variable

Angiosperms show an evolutionary trend from an indefinite to a fixed number of floral organs. When floral formula inconstancy in recent angiosperms is reported, it is often considered as a byproduct of stress and its fitness consequences remain mostly unexplored. We report substantial nonhomeotic meristic variation in stamen number (0-10 stamens per flower) in two populations of Hormathophylla spinosa during four years. This variation was plastic, suggesting its functional role in the adjustment of phenotypic gender. However, no correlations were found between phenotypic gender and plant size, pollinator or herbivore abundance. Effects on female reproductive success were inconsistent on a per-flower and on a per- plant basis, rendering adaptive explanations in terms of selfing or resource adjustment unsatisfac- tory with the data available. Nevertheless, individ- uals showing larger interannual variation in phenotypic gender showed higher female reproduc- tive success, suggesting an advantage for gender modification. Although our results do not easily conform to any adaptive explanation, this remark- able example of breakdown of trait canalization should stimulate the study of the mechanisms and ultimate causes responsible for the maintenance of fixed floral traits.