Biogeography and systematics of Carex subgenus Uncinia (Cyperaceae): A unique radiation for the genus Carex in the Southern Hemisphere

Carex subg. Uncinia (Cyperaceae) constitutes one of six currently recognized Carex subgenera. This subgenus is mainly distributed on the American continent and in the Pacific region, and it is the only subgenus almost entirely absent from the Old World and primarily diversified in the Southern Hemisphere. It includes some of the few Carex species with clear epizoochoric traits: the representatives of C. sect. Uncinia possess utricles with an exserted and hooked rachilla that allows the diaspores to attach to feather or hair. We performed phylogenetic (ITS, ETS-1f, matK), biogeographic, and ancestral state reconstruction analyses to elucidate the systematic structure, origin and dispersal routes, and major morphological evolutionary patterns of the different lineages within the subgenus. Our phylogenetic reconstructions revealed that the subgenus comprises seven different clades that mostly match previously recognized sections. One of the clades, however, represents a new section described herein as C. sect. Wheelerianae. Unispicate lineages evolved repeatedly from ancestors bearing multispicate inflorescences, while the presence of a rachilla, often pictured as a plesiomorphy in Carex, seems to have developed four independent times in the evolution of C. subg. Uncinia. The origin of the subgenus dates back to the beginning of the Miocene, probably in North America from where it colonized the Southern Hemisphere. It first dispersed to South America during the Early Miocene. Later, in the Middle Miocene, representatives of C. sect. Uncinia would reach the Pacific Southwest region (New Zealand, Australasia) from South America in at least two independent dispersal events. The vast majority of the biogeographic events seem to be explained by long-distance dispersal. The remarkable dispersal ability of C. sect. Uncinia enabled by the hooked rachilla has allowed it to reach remote archipelagos in the Pacific and Subantarctic regions, probably bird-mediated.