A systematic study of Berkheya and allies (Compositae)

Abstract

Berkheya Ehrh. is a genus of daisies in the tribe Arctotideae, subtribe Gorteriinae with over 80 species, most of which occur in southern Africa. This genus has centres of diversity associated with the montane regions of South Africa, including the Drakensberg Alpine Centre and Mpumalanga escarpment regions. Previous molecular and morphological studies indicate that Berkheya is paraphyletic. I present phylogenies based on nrDNA (ITS; Internal Transcribed Spacer) and cpDNA (psbA-trnH) sequence data analysed with Bayesian Inference and Parsimony. A phylogeny of combined cp- and nrDNA is also presented. These phylogenies are used to assess generic limits and to investigate the biogeographic patterns of Berkheya and its allies. The ITS phylogeny shows five well supported clades of Berkheya, two of which (Clades I and 2) are monophyletic summer rainfall region clades. Clades 3 to 5 are all paraphyletic winter rainfall clades with Cullumia occurring in the third clade, Cuspida occurring in the fourth and Didelta in Clade 5. Both psbA-trnH and combined phylogenies show concordance with the clade distribution shown in the ITS phylogeny. The ITS phylogeny was used to analyse correspondence with Roessler's (1959) Series. It was found that the phylogeny showed considerable agreement with Roessler's series, but B. bipinnatifida and B. spinosa of Series Speciosae may require some revision as well as taxa of monotypic series Cruciatae and Angustae. It is suggested that the latter series be merged with Cullumia species to form a single series. Some consideration should be taken to include Didelta species into Series Fruticosae as Didelta occurs in subclade 5b of Clade 5 with other Series Fruticosae taxa. Achene morphology was examined from species from each of the five clades to investigate the relationships of Berkheya and its allies, as well as to determine if there were any consistent achene features for each clade. The structure of surface cells on the fruit, the presence, absence and morphology of twin hairs as well as the structure of the pappus scales were found to be most useful in reflecting phylogenetic relationships within the clades. When compared with the clades of the ITS phylogeny, achene morphology showed consistent characters between taxa occurring in the same clades. As the most comprehensive study involving Berkheya, this phylogenenetic investigation was able to confirm that Berkheya is a paraphyletic genus with Didelta, Cullumia and Cuspida needing to be subsumed into Berkheya. An alternative classification is that taxa of Clade 5 could possibly be erected as an expanded Didelta, separate and sister to Berkheya.