EVOLUTION OF PLANT POLLINATION SYSTEMS: HYPOTHESES AND TESTS WITH THE NEOTROPICAL VINE DALECHAMPIA.

The results of pollination and mating-system studies were integrated with a phylogenetic study of 40 Neotropical species of Dalechampia L. (Euphorbiaceae) to reconstruct the history of evolutionary change in pollination systems. The results of this analysis were treated as a hypothesis and tested for circularity problems and robustness in the face of changes in the data set. The historical hypothesis was used to make specific predictions about details of pollination ecology and reward biochemistry; the predictions were supported by independent observations. I conclude that pollination systems in Dalechampia have been evolutionarily labile, relative to most morphological features, with repeated parallelisms and reversals. Transitions among the three pollination systems evolved by Dalechampia (pollination by resin-collecting bees, fragrance-collecting male euglossine bees, and pollen-collecting bees) have been facilitated by biochemical exaptation (preadaptation). Pollination by male euglossine bees is relatively rare in the genus but has originated independently three to four times. In contrast, pollination by resin-collecting female bees is very common, but has originated only once. Eighty-six to 97% of transitions between pollination systems involved an intermediate phase during which both old and new pollinators were effective, but 3 to 14% of transitions may have been "instantaneous," lacking the intermediate phase. Clades of species secreting resin rewards are about 10 times as species rich as clades of species secreting fragrance rewards; circumstantial evidence suggests that different extinction rates may be responsible for this difference. Relatively allogamous (cross-pollinating) species have evolved from more autogamous (self-pollinating) species up to 13 times, and autogamous species have evolved from more allogamous ones up to 11 times. Species occurring in disturbed habitats are facultatively autogamous, whereas species of undisturbed habitats are often highly allogamous.