First Miocene fossils of Vivianiaceae shed new light on phylogeny,divergence times,and historical biogeography of Geraniales

The origin of Geraniales (approximately 900 species in three families: Geraniaceae, Melianthaceae, and Vivianiaceae) is traced back to the Cretaceous of Gondwana, yet their geotemporal history is largely unknown because of a limited fossil record and incomplete phylogenies. In the present study, we provide the first fossil record of Vivianiaceae and a highly resolved molecular phylogeny for all extant Geraniales genera. Our results support the hypothesis that five (instead of three) families should be recognized in the order Geraniales: Francoaceae A. Juss. (Francoa, Greyia, Tetilla), Geraniaceae Juss. (Erodium, Geranium, Monsonia, Pelargonium), Hypseocharitaceae Wedd. (monogeneric), Melianthaceae Horan. (Bersama, Melianthus), and Vivianiaceae Klotzsch (Balbisia, Rhynchotheca, Viviania). The four major lineages (i.e. Geraniaceae, Francoaceae + Melianthaceae, Hypseocharitaceae, Vivianiaceae) all originated within a narrow time frame during the Eocene (36.9–49.9 Mya) based on the five fossil calibration points. The divergence of most of the extant genera occurred much later, from the Miocene onwards. The South American–South African disjunction in Francoaceae apparently goes back to long distance dispersal with an estimated divergence time of the lineages in the Middle Miocene 11.2 (5.9–17.7) Mya]. Diversification in Melianthus appears to be much more recent than previously assumed starting approximately 3.4 (1.9–5.2) Mya rather than approximately 8–20 Mya]. However, divergence of the Andean Hypseocharis lineage 36.9 (31.9–42.8) Mya] significantly predates the main Andean uplift: Current distributions likely go back to northward migrations and subsequent extinctions in Patagonia. Similarly, Rhynchotheca, Balbisia, and Viviania have a current southern distribution limit > 10°N of the fossil finds, indicating a massive northward displacement. The present evidence suggests that niche conservatism likely played a major role in the historical biogeography of Geraniales. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ?? , ??–??.