Out of Amazonia and Back again: Historical Biogeography of the Species-Rich Neotropical Genus Philodendron (Araceae)1

Abstract The origin of Neotropical species diversity is strongly associated with the geological history of South America. Since the Miocene, a number of species radiations across different Neotropical lineages coincided with the rise of the Andes and the formation of the Isthmus of Panama. The species-rich genus Philodendron Schott (Araceae) is widely distributed across Neotropical rainforests, originating in the Late Oligocene and diversifying more intensely from the Miocene onward. It is likely that its diversification process and distribution patterns are associated with recent geological changes in the Americas. To test this hypothesis, we sampled the species diversity of Philodendron across its entire geographic range and used a combination of three non-coding plastid regions (petD, rpl16, and trnK/matK) to obtain a comprehensive time-calibrated phylogeny. We then inferred geographic range evolution and explored the impact of the Andean orogeny on speciation, extinction, and dispersal. The genus Philodendron originated ∼29 million years ago (mya) and experienced the earliest diversification events ∼25 mya in the Pan-Amazonian rainforests. From the Middle Miocene onward, multiple geographic range expansion events occurred from Amazonia to southeast Brazil and to the area which would become the Chocó and the northern Andes. From the Pliocene onward, Philodendron reached Central America and the Caribbean islands, and Andean lineages recolonized and diversified in Amazonia. In Philodendron, higher diversification rates are found in the adjacent lowland rainforests of the northern Andes compared with other regions in the Neotropics, demonstrating a potential indirect impact of the Andean uplift on species radiations in lowland regions.