POLLINATION OF RAVENALA-MADAGASCARIENSIS (STRELITZIACEAE) BY LEMURS IN MADAGASCAR - EVIDENCE FOR AN ARCHAIC COEVOLUTIONARY SYSTEM

Investigations of the floral biology of the traveler's tree (Ravenala madagascariensis) and the ecology of the ruffed lemur (Varecia variegata), both endemic to the island of Madagascar, suggest a plant-pollinator relationship. Ravenala exhibits many specializations for visitation by large nonflying animals: inflorescences placed below the crown of the plant and easily accessible to arboreal animals; large flowers enclosed in tough, protective bracts that require a strong pollinator to open; stiff, rodlike styles that withstand the rough handling of the visitors; and copious, sucrose-dominant nectar that provides a renewable reward for a sizable animal for a long time period. Our observations in the field also show that Varecia variegata: consistently and almost exclusively visit the flowers of Ravenala; carry pollen on their fur between flowers on the same plant and between conspecific plants; do not destroy the flowers while obtaining the nectar; and appear to be highly dependent on nectar as a food source during specific times of the year. The basal phylogenetic position of Ravenala in the family Strelitziaceae, as indicated by molecular sequence data, and the distribution of reproductive traits in the three extant genera are consistent with the hypothesis that pollination by nonflying mammals is an archaic system, whereas bird and bat pollination are derived.