Comparative analysis of late floral development and mating-system evolution in tribe Collinsieae (Scrophulariaceae s.l.).

Species of Collinsia and Tonella, the two sister genera of self-compatible annuals that constitute tribe Collinsieae, show extensive variation in floral size and morphology and in patterns of stamen and style elongation during the life of the flower (anthesis). We used a nuclear ribosomal ITS phylogeny, independent contrasts, and phylogenetically corrected path analysis to explore the patterns of covariance of the developmental and morphological traits potentially influencing mating system. Large-flowered taxa maintain herkogamy (spatial separation of anthers and stigmas) early in anthesis by differential elongation of staminal filaments, which positions each of the four anthers at the tip of the "keel" upon dehiscence. Small-flowered taxa do not show this pattern of filament elongation. The styles of large-flowered taxa elongate late in the 2-5 d of anthesis, resulting in late anther-stigma contact and delayed self-pollination. Anther-stigma contact and self-pollination occur early in anthesis in small-flowered species/populations. Thus, we found complex covariation of morphological and developmental traits that can be interpreted as the result of multitrait adaptation for early selfing and high levels of autogamy, delayed selfing and higher levels of outcrossing, or intermediate levels of outcrossing. Continuous variation in these traits suggests the operation of continuous variation in selective optima or the combined effects of divergent selection and phylogenetic inertia.