Hawkmoth pollination of aerangoid orchids in Kenya, with special reference to nectar sugar concentration gradients in the floral spurs

The African orchid flora has a high proportion of species with long-spurred white flowers. Few data exist to test the prediction that this floral syndrome pattern reflects an important role for hawkmoth pollination in the evolution and ecology of these orchids. The pollination biology of five aerangoid orchid species (Rangaeris amaniensis, Aerangis brachycarpa, A. confusa, A. thomsonii, and A. kotschyana) was investigated in Kenya. Four of these have long spurs (>10 cm) and were pollinated by Agrius convolvuli and Coelonia fulvinotata. Aerangis confusa, which has relatively short spurs (ca. 4 cm), was pollinated by the short-tongued hawkmoths Hippotion celerio and Daphnis nerii. Nectar frequently filled the entire spur in some of the study species, even at anthesis. Sugar concentration of the nectar of four species was found to vary from ca. 1% at the mouth of the spur to 20% at the tip. Gradients were expressed more strongly in species with long, straight spurs. Species with spirally twisted spurs showed both steep and shallow nectar gradients. These gradients, previously unknown in plants, may function as a "sugar trail," enticing longtongued hawkmoths to probe deeply into spurs without incurring the cost of filling an entire spur with concentrated nectar. In addition, the most concentrated nectar is kept out of reach of short-tongued pollinators.