Phylogenetic relationships and evolutionary trends in Tillandsia subgenus Diaphoranthema and xerophytic species of subgenus Phytarrhiza (Bromeliaceae: Tillandsioideae)

Tillandsia subgenera Diaphoranthema and Phytarrhiza s.s. are closely related epiphytic and epilithic plants adapted to xeric habitats in the Americas. Several studies suggested that subgenus Diaphoranthema and the xerophytic species of subgenus Phytarrhiza should not be segregated into different subgenera. In recent phylogenetic analyses of Tillandsioideae, subgenus Phytarrhiza s.l. is revealed as highly polyphyletic, showing a xerophytic lineage closely related to subgenus Diaphoranthema. Moreover, the evolution of neotenic traits, a selfing breeding system and polyembryony have yet to be fully investigated in a phylogenetic context. Infrageneric classifications and phylogenetic relationships were inferred using the plastid matK-trnK and nuclear ETS markers and morphology. Phylogenetic analyses of individual and combined molecular data sets were performed under parsimony and Bayesian inference. Our results confirm subgenus Diaphoranthema as monophyletic if three xeric species of subgenus Phytarrhiza are included. In subgenus Diaphoranthema, aggregates Loliacea and Rectangula were recovered as monophyletic, whereas Myosura was paraphyletic and Caliginosa, Capillaris and Recurvata were polyphyletic. Alternative groupings and morphological trends are discussed. Analysis of morphological characters associated with heterochrony indicates that neoteny may not be the only paedomorphic process in the evolution of this group. Furthermore, paedomorphosis would occur in some vegetative structures, inflorescences and flowers, whereas some fruit and seed traits could be modified by peramorphosis.