FLORAL STRUCTURE AND EVOLUTION IN LOPEZIEAE (ONAGRACEAE)

The Lopezieae present an interesting mixture of ancestral and derived characters: some members of the tribe retain the basic onagraceous chromosome number (n = 11), but the flowers are advanced in that they are mostly zygomorphic and always have a two‐merous androecium. Species differ in the position of the nectaries, also in the way in which floral parts are united above the inferior ovary. These differences, when analyzed with information from a new monograph of the Lopezieae, provide the basis for a phylogenetic tree. It is inferred that ancestral Lopezieae were bird‐pollinated woody perennials with regular flowers, two fertile stamens, and no floral tube distal to the ovary. Evolutionary events accompanying the emergence of modern taxa included abortion of the abaxial stamen (all surviving Lopezieae except Lopezia lopezioides), development of an epigynous floral tube (L. riesenbachia, L. semeiandra), decrease in floral symmetry without conversion to insect pollination (in two independent lines), and decrease in floral symmetry with conversion to insect pollination (in at least two independent lines). The prominent tubercles on upper petals of certain insect‐pollinated species apparently evolved from the less prominent swollen areas still present in some of the bird‐pollinated species. The tubercles and an associated snapping mechanism arose in response to increasing selection for fly pollination. Densely staining areas in some specimens may be osmophores; if so, scent plays a supplementary role in the orienting of insects to the upper petals. Interstaminal nectaries and the absence of a floral tube link the Lopezieae to Ludwigia; the relationship of these two taxa to Epilobium is presently unclear. Fossil records indicate that the Onagraceae had evolved by the beginning of the Tertiary Period and that the Ludwigia line is very old. The family's ancestral features are retained to a greater degree in Fuchsia, however, than in Ludwigia.