Evolution of fruit and seed characters in the Diervilla and Lonicera clades (Caprifoliaceae, Dipsacales).

BACKGROUND AND AIMS The Diervilla and Lonicera clades are members of the family Caprifoliaceae (Dipsacales sensu Donoghue et al., 2001, Harvard Papers in Botany 6: 459-479). So far, the intergeneric relationships of the Lonicera clade and the systematic position of Heptacodium remain equivocal. By studying fruit and seed morphology and anatomy, an attempt is made to clarify these issues. In addition, this study deals with the evolution of fruit and seed characters of the Diervilla and Lonicera clades with reference to allied taxa. METHODS Light and scanning electron microscopy were used for the morphological and anatomical investigations. Phylogenetic analyses were carried out by applying the parsimony and Bayesian inference optimality criteria. Character evolution was studied by means of parsimony optimization and stochastic character mapping. KEY RESULTS Diervilla and Weigela (Diervilla clade) are characterized by several unique traits in Dipsacales, including capsules with numerous seeds, seed coats without sclerified outer tangential exotestal cell walls, and dehiscent fruits. Seeds with completely sclerified exotestal cells and fleshy fruits characterize the Lonicera clade. Leycesteria and Lonicera have berries, ovaries without sterile carpels and several seeds per locule, whereas Symphoricarpos and Triosteum have drupes, ovaries with one or two sterile carpels and a single seed per locule. Heptacodium shares several characteristics with members of the Linnina clade, e.g. achenes, single-seeded fruits and a compressed, parenchymatous seed coat. CONCLUSIONS The results confirm the monophyly of the Diervilla and Lonicera clades and allow us to hypothesize a close relationship between Leycesteria and Lonicera and between Symphoricarpos and Triosteum. Fruit and seed morphology and anatomy point to a sister relationship of Heptacodium with the Linnina clade, rather than with the Lonicera clade.