Iridaceae 'Out of Australasia'? phylogeny, biogeography, and divergence time based on plastid DNA sequences

The current infrafamilial taxonomy of the Iridaceae recognizes four subfamilies; Isophysidoideae (1: 1); Nivenioideae (6: ca. 92), Iridoideae (29: 890), and Crocoideae (29: 1032). Phylogenetic analyses of sequences of five plastid DNA regions, rbcL, rps4, trnL-F, matK, and rps16, confirm most aspects of this classification and the evolutionary patterns that they imply, importantly the sisiter relationship of Isophysidoideae to the remainder of the family and the monophyly of Iridoideae. Subfamily Nivenioideae is, however, paraphyletic; Crocoideae is consistently found nested within it, sister to the core Nivenioideae, the woody Klattia, Nivenia, and Witsenia. This clade is sister to Aristea, which in turn is sister to the Madagascan Geosiris, and then to the Australasian Patersonia. We treat Aristea, Geosiris, and Patersonia as separate subfamilies, Aristeoideae and the new Geosiridaceae and Patersonioideae, rendering Nivenioideae and Crocoideae monophyletic. The alternative, uniting a widely circumscribed Nivenioideae and Crocoideae, seems undesirable because Nivenioideae have none of the numerous synapomorphies of Crocoideae, and that subfamily includes more than half the total species of Iridaceae. Main synapomorphies of Crocoideae are: pollen operculate; exine perforate; ovule campylotropous; root xylem vessels with simple perforations; rootstock a corm; inflorescence usually a spike; plants deciduous. Four more derived features of Crocoideae are shared only with core Nivenioideae: flowers long-lived; perianth tube well developed; flowers sessile; and septal nectaries present. The genera of the latter subfamily are evergreen shrubs, have monocot-type secondary growth, tangentially flattened seeds, and the inflorescence unit is a binate rhipidium. The latter feature unites core Nivenioideae with Aristea, Geosiris, and Patersonia, which have fugaceous flowers and, with few exceptions, a blue perianth. Molecular-based phylogenetic trees using sequences from five plastid DNA regions now show discrete generic clusters within Crocoideae and Iridoideae, the foundation for the tribal classification. The five tribe classification of Iridoideae, initially based on morphological characters and subsequently supported by a four plastid DNA region sequence analysis, continues to receive support using additional DNA sequences. Application of molecular clock techniques to our phylogeny indicates that the Iridaceae differentiated in the late Cretaceous and diverged from the next most closely related family, Doryanthaceae circa 82 mya, thus during the Campanian. The Tasmanian Isophysis is the only extant member of the clade sister to the remainder of the Iridaceae, from which it may have diverged 66 mya, in the Maastrichtian. The generic phylogeny shows the proximal clades of the family are all Australasian, which corroborates past hypotheses that the Iridaceae originated in Antarctica-Australasia, although its subsequent radiation occurred elsewhere, notably in southern Africa and temperate and highland South America at the end of the Eocene or later.