Phylogeny, morphology, and biogeography of Chuquiraga, an Andean-Patagonian genus of Asteraceae-Barnadesioideae

Chuquiraga is a genus of 23 species of evergreen shrubs endemic to South America. It is distributed principally along the Andes from Colombia to Chile and Argentina, and it is especially diversified in the Central Andes and in the deserts and semideserts of southern South America. The genus exhibits a wide array of leaf-morphology types and two different head and floral types apparently related to hummingbird and insect pollination. In this study, phylogenetic relationships among Chuquiraga species were resolved by parsimony cladistic analysis using morphological characters. The resulting cladogram was used to interpret morphological, ecological, and biogeographical patterns in a historical context. Biotic and abiotic environmental factors hypothesized to have exerted selective pressure on morphological traits of its species were optimized onto the phylogeny to suggest how and when these factors may have affected the evolution and diversification of the genus. Results suggest an origin of the genus in southern South America, with two major evolutionary radiations, one more northern in the Central and Northern Andes, and the other in the Southern Andes and the North Chilean, Patagonia and Monte Deserts. Pollination by hummingbirds seems to have been an important factor in the origin of the northern clade, affecting floral morphology. Herbivory by vertebrates and increased aridity seem to have been important selective forces in the evolution and diversification of the southern clade, especially affecting leaf morphology. These changes were probably associated with the major elevation of the Andes in late Tertiary and with the hyperaridization and climatic fluctuations of Pleistocene and Holocene times.