Crassula, insights into an old, arid-adapted group of southern African leaf-succulents.

The Crassulaceae is an important family in the Greater Cape Floristic Region of southern Africa and is the seventh largest family in the arid Succulent Karoo Biome. After the Aizoaceae it is the largest group of leaf-succulents in southern Africa. This is the first investigation of a broad selection (68%) of the ±170 species of Crassula. We used data from three chloroplast and two nuclear gene-regions, which yielded many informative characters and provided good resolution among the species. We show that only five of the 20 sections in Crassula are monophyletic. However, the clades recovered show close correlation with the two subgenera that were once recognized. Crassula contains more than 25 succulent annual species which are not closely related to each other but form early-diverging branches in each of the three major clades. One of these major clades contains far more perennial species than the others and is the greatest diversification within Crassula. This diversification mostly arose within the last 10 million years (my) and spread across much of southern Africa. Members of the smaller two major clades are often soft- and flat-leaved perennials (many with basic chromosome number x = 8, with high levels of polyploidy). Those in the largest diversification (where a basic chromosome number of x = 7 predominates) show other arid-adaptations (more highly succulent leaves with a dense covering of hairs or papillae or a smooth xeromorphic epidermis). Their flowers are also more variable in shape and bee-, moth- and butterfly-pollinated species are known among them. We establish that Crassula arose in the Greater Cape Floristic Region of southern Africa. While much of its diversity has evolved in the last 10 my, Crassula nevertheless contains species that are much older and itself arose ±46 my ago. Since all its species are succulent it is possible that they are part of an early arid-adapted flora that contributed to the Succulent Karoo Biome in the western part of southern Africa. Consequently this Biome may not be assembled only from 'young lineages' as is usually thought to be the case.

Recent radiation of Brachystelma and Ceropegia (Apocynaceae) across the Old World against a background of climatic change.

The genera Brachystelma Sims and Ceropegia L. of the Ceropegieae (Apocynaceae-Asclepiadoideae) consist of ±320 species of geophytes and slender climbers with a tendency to stem-succulence in Ceropegia. They occur in and around the semi-arid, mainly tropical parts of the Old World. For 146 species (around half of the total) from most of the geographic range of the genera, we analysed data from two nuclear and five plastid regions. The evolution of Ceropegia is very complex, with at least 13 mostly well-supported lineages, one of which is sister to the ±350 species of stapeliads. Species of Brachystelma have evolved at least four times, with most of them nested within two separate major lineages. So, neither Brachystelma nor Ceropegia is monophyletic. We recover a broad trend, in two separate major lineages, from slender climbers to small, geophytic herbs. Several clades are recovered in which all species possess an underground tuber. Small, erect, non-climbing, geophytic species of Ceropegia with a tuber are nested among species of Brachystelma. Consequently, the distinctive tubular flowers used to define Ceropegia do not reflect relationships. This re-iterates the great floral plasticity in the Ceropegieae, already established for the stapeliads. Both major lineages exhibit a trend from tubular flowers with faint, often fruity odours, pollinated by very small Dipteran flies, to flatter flowers often with a bad odour, pollinated by larger flies. Most of the diversity in Brachystelma and Ceropegia is recent and arose within the last 3my against a background of increased aridification or extreme climatic variability during the Pliocene. In the ingroup, diversity is highest in Southern Africa, followed by Tropical East Africa and other arid parts of Africa, the Arabian Peninsula and India. Many disjunctions are revealed and these are best explained by recent, long distance dispersal. In Africa, the diversity arises from the presence of many different lineages over wide areas but there is also evidence of closely related species growing together with different pollinators.

Evolution of the stapeliads (Apocynaceae-Asclepiadoideae) - repeated major radiation across Africa in an Old World group.

The stapeliads of the Ceropegieae (Apocynaceae-Asclepiadoideae), are approximately 340 species of stem-succulents placed in around 30 genera, found in semi-arid parts of the Old World. Here we sampled 192 species (i.e. nearly two thirds of the total) from across the full geographic range of the group and analysed data from the two nuclear regions (nuclear ribosomal ITS and ncpGS) and five plastid regions (psbA-trnH intergenic spacer, rps16 intron, trnL-trnF intergenic spacer, trnS-trnG intergenic region and the non-coding rpl32-trnL region). We find that the stapeliads radiated first in the northern hemisphere from Africa to southern Europe and Myanmar. This radiation subtends a grade of minor clades in the south-western corner of the African continent. These were followed by a single clade containing major radiation back across Africa from South Africa to tropical Arabia (but no further east than Dhofar, Oman), which includes also a single early spread into Madagascar. We establish the monophyly of many of the genera, such as Echidnopsis Hook.f., Hoodia Hook., Huernia R. Br., Piaranthus R. Br., Rhytidocaulon P.R.O. Bally and Tridentea Haw., but find that Duvalia Haw., Orbea Haw., Stapelia L. and Tromotriche Haw. are polyphyletic. We show that in certain vegetative features, there is broad cohesion across clades. Florally, on the other hand, the stapeliads exhibit considerable plasticity and we are able to show that very differently shaped flowers as well as large and small flowers evolved repeatedly among closely related species.