Origin of the Woody Macaronesian Sempervivoideae and the Phylogenetic Position of the East African Species of Aeonium

Evolutionary relationships among the genera of Macaronesian Sempervivoideae, Aeonium, Aichryson, Greenovia, and Monanthes, were studied using sequence variation of the chloroplast DNA trnL (UAA) - trnF (GAA) spacer and the nuclear ribosomal Internal Transcribed Spacer 2 (ITS2). Phylogenetic analysis indicates that the Moroccan Sedum sect. Monanthoidea is the sister taxon of the Macaronesian Sempervivoideae. In combination with the terminal position in the Macaronesian Sempervivoideae of the East African A. leucoblepharum, which has formerly been assumed to be the sister taxon of the other species of Macaronesian Sempervivoideae (i.e. Aichryson and Monanthes), a recent remigration to Africa is suggested. Statistical support for the terminal position of A. leucoblepharum using only spacer sequences is without homoplasy but not high since only single mutations in both the chloroplast and nuclear sequence characterize the clade containing A. leucoblepharum. A. leucoblepharum and the Canarian species with a similar growth-form share 50% of the RAPDs. Within a clade comprising woody species with yellow flowers and a herbaceous rosette, the highest genetic divergence, as determined with RAPDs, is found between A. simsii and the woody Macaronesian and African species. The extremely close genetic ties among the woody and branched (sub)shrubs indicate that, when compared to the other species of the genus, the woody, African Aeonium species are not the sister group of the Macaronesian Sempervivoideae and substantiate the view that an ancestor of A. leucoblepharum recently migrated from the Canary Islands to East Africa and Arabia through long distance dispersal, rather than being a relict of an African Aeonium flora from the Tertiary.