Aliciella heterostyla represents the only documented case of the unusual breeding system heterostyly in Polemoniaceae. Two competing hypotheses, explaining the origin of heterostyly, have been proposed. These hypotheses differ with respect to the predicted order in which characteristics associated with the heterostyly syndrome arise (e.g., Darwin 1877; Lloyd and Webb 1992 versus Charlesworth and Charlesworth 1979). This syndrome includes reciprocal herkogamy, self-incompatibility and a host of ancillary morphological characteristics, such as variation in pollen size and shape. Sequence data from the internal transcribed spacer (ITS) region of nuclear ribosomal DNA are used to estimate the phylogenetic relationships among A. heterostyla and its relatives. The resulting molecular phylogeny is used as an evolutionary setting to estimate the order of development of physiological and morphological characters associated with heterostyly. Although reciprocal herkogamy is uniquely derived in A. heterostyla , compatibility is more complicated. Both self-compatibility (SC) and self-incompatibility (SI) are present in Aliciella . Within A. heterostyla , however, a mixed mating system (cryptic self-incompatibility and intra-morph compatibility) is exhibited. The ancestral condition of this trait is greatly influenced by assumptions about the relative ease of deriving SC from SI versus deriving SI from SC. As a consequence, both the Lloyd and Webb and the Charlesworth and Charlesworth models may be consistent with the Aliciella data, depending upon assumptions made about the evolution of compatibility.

Key words: Aliciella, heterostyly, Polemoniaceae, self-incompatibility