Molecular phylogenetics of the Espeletia complex (Asteraceae): evidence from nrDNA ITS sequences on the closest relatives of an Andean adaptive radiation.

The subtribe Espeletiinae (Asteraceae, Heliantheae) comprises morphologically and ecologically diverse plants endemic to the tropical montane paramos of the Andes of Venezuela, Colombia, and Ecuador. Though the ecophysiology and ecology of this adaptive radiation have been well studied, relationships among taxa in the subtribe and between the subtribe and other taxa in the Heliantheae are poorly known. In this study, sequences from the internal transcribed spacer (ITS) region of nuclear ribosomal DNA are used to test previous hypotheses about the phylogenetic position of the Espeletiinae within the Heliantheae and to determine which taxa are the subtribe's closest relatives. Gene phylogenies based on maximum parsimony analyses reveal that the Espeletiinae clade is nested well within the subtribe Melampodiinae and thus should be considered a monophyletic complex of species, not a separate subtribe. The most parsimonious gene trees suggest that the genus Ichthyothere may be the sister taxon to the Espeletia complex and that the genus Smallanthus and a species of Rumfordia are likely among the complex's other closest living relatives. These data offer preliminary insights into the origins of this adaptive radiation and the broader phylogenetic context in which it occurred.