Leptosporangiate ferns, with more than 9000 extant species, are truly exceptional among the non-flowering lineages of vascular plants. However, this rather remarkable diversity was not simply a consequence of being able to "hold on" as flowering plants rose to dominance. Instead, it appears to be the result of an ecological opportunistic response to the establishment of more complex, angiosperm-dominated ecosystems. The proliferation of flowering plants across the landscape undoubtedly resulted in the formation of a plethora of new niches into which leptosporangiate ferns could diversify. Many of these were evidently on shady forest floors, but many others were actually within the new angiosperm-dominated canopies. Today, almost one third of leptosporangiate species grow as epiphytes on angiosperm trees. My dissertation aims to demystify the evolution and diversification of epiphytic ferns in order to more fully understand the leptosporangiate success story. By assembling and analyzing the most inclusive molecular dataset for leptosporangiate ferns to date, I provide unprecedented insight into overall fern relationships and a solid and balanced phylogenetic framework within which the evolution of epiphytism can be examined. By employing this phylogeny and numerous constraints from the fern fossil record, I uncover the timing of epiphytic fern diversification and examine the origin of the modern tropical rain forest biome in which these ferns reside. / Dissertation