Phylogenetics of Puya (Bromeliaceae): Placement, major lineages, and evolution of Chilean species.

Puya (Bromeliaceae), a large genus of terrestrial bromeliads found throughout a range of elevations in the Andes and central Chile, is of great systematic, evolutionary, and biogeographical interest. This first molecular phylogenetic study of Puya and related bromeliads employs matK, trnS-trnG, rps16, and PHYC sequences. Chloroplast DNA, nuclear DNA, and combined DNA data all place Puya closest to subfamily Bromelioideae. Nuclear and combined data support Puya as monophyletic, and the two subgenera are nonmonophyletic. All data indicate that the Chilean species of Puya are early diverging within the genus, consistent with Chilean genera as the first-diverging members of subfamily Bromelioideae. Central Chile is identified as a key region for understanding the biogeographical history of Bromeliaceae, as is true with other South American plant groups. A complicated history involving early chloroplast capture and later secondary hybridization and/or introgression is seen in Chilean lineages. These events help explain the occurrence of sterile inflorescence tips, floral color and shape, and leaf indument. The ecological radiation of Puya appears coincident with the final, recent rise of the Andes and subsequent high-elevation habitat diversification. Additionally, geographical distribution, rather than moisture or elevational adaptations, correlates to species relationships. Evolution of CAM photosynthesis has occurred multiple times.