Clades, clocks, and continents: Historical and biogeographical analysis of Myrtaceae, Vochysiaceae, and relatives in the Southern Hemisphere

Some of the most interesting but still most contentious disjunct biogeographical distributions involve Southern Hemisphere tropical and warm temperate families. The PHMV clade of Myrtales includes four families (Psiloxylaceae, Heteropyxidaceae, Myrtaceae, and Vochysiaceae) that exhibit a number of these biogeographical patterns. The related Psiloxylaceae and Heteropyxidaceae are small families restricted in distribution to the recent volcanic Mascarene Islands to the east of Madagascar and to southeast Africa, respectively. Myrtaceae are found on three major Gondwanan regions (South America, Australasia, and Africa). Because the New World taxa are almost exclusively fleshy fruited, it is unclear whether the family distribution is a classic Gondwanan vicariance pattern or results from one or more long-distance dispersal events over ocean barriers. The Vochysiaceae represent one of a handful of families with amphi-Atlantic distributions vigorously argued to support both long-distance dispersal over the Atlantic and vicariance of western Gondwanan biota by Atlantic seafloor spreading. Molecular phylogenetic relationships, fossil dating of nodes, and penalized likelihood rate smoothing of maximum likelihood trees were employed for a Myrtales-wide analysis using rbcL and ndhF and an analysis of the PHMV analysis using ndhF and matK. The results indicate that the PHMV differentiated during the late Cretaceous. The African lineage of Vochysiaceae is nested within a South American clade and probably arose via long-distance dispersal in the Oligocene at a time when the Atlantic had already rifted 80 m. yr. at the equatorial region. The African/Mascarene Island families, most closely related to Myrtaceae, differentiated during the late Eocene, with subsequent but recent long-distance dispersal from Africa to the Mascarenes. Myrtaceae show a rapid differentiation of a basal, paraphyletic subf. Leptospermoideae in Australasia. Fleshy-fruited taxa (subf. Myrtoideae) are not monophyletic. Vicariance of a widespread warm temperate Southern Hemisphere distribution is likely in explaining the South American-Australasian disjunction, with subsequent dispersal events between the two and to Africa and the Mediterranean basin.