Diagnostics, taxonomy, nomenclature and distribution of perennial Sesuvium (Aizoaceae) in Africa.

The taxonomy of perennial Sesuvium species in Africa has been poorly investigated until now. Previously five perennial species of Sesuvium were recognised in Africa (S. congense, S. crithmoides, S. mesembryanthemoides, S. portulacastrum, and S. sesuvioides). Based on the differing number of stamens, S. ayresii is accepted here as being distinct from S. portulacastrum. Field observations in Angola also led the authors to conclude that S. crystallinum and S. mesembryanthemoides are conspecific with S. crithmoides. A new subspecies, Sesuvium portulacastrum subsp. persoonii, is described from West Africa (Cape Verde, Gambia, Guinea-Bissau, Mauritania, Senegal). The molecular phylogeny indicates the position of S. portulacastrum subsp. persoonii within the "American lineage" as a part of the Sesuvium portulacastrum complex which needs further studies. A diagnostic key and taxonomic notes are provided for the six perennial species of Sesuvium found in Africa and recognised by the authors (S. ayresii, S. congense, S. crithmoides, S. portulacastrum subsp. portulacastrum, S. portulacastrum subsp. persoonii, S. verrucosum and the facultatively short-lived S. sesuvioides). The distribution of S. crithmoides, previously considered to be endemic to Angola, is now confirmed for the seashores of Republic of Congo and DR Congo. The American species S. verrucosum is reported for the first time for Africa (the Macaronesian islands: Cape Verde and the Canaries). It is locally naturalised in Gran Canaria, being a potentially invasive species. These findings as well as new records of S. verrucosum from Asia and the Pacific Islands confirm its proneness to transcontinental introduction. Lectotypes of S. brevifolium, S. crithmoides, S. crystallinum and S. mesembryanthemoides are selected. The seed micromorphology and anatomy of the perennial African species is studied. Compared to the seeds of some annual African Sesuvium investigated earlier, those of perennial species are smooth or slightly alveolate. The aril is one-layered and parenchymatous in all species and usually tightly covers the seed. The aril detachments from the seed coat that form a white stripe near the cotyledon area easily distinguish S. verrucosum from other species under study.

Out of southern Africa: Origin, biogeography and age of the Aizooideae (Aizoaceae).

The Aizooideae is an early-diverging lineage within the Aizoaceae. It is most diverse in southern Africa, but also has endemic species in Australasia, Eurasia and South America. We derived a phylogenetic hypothesis from Bayesian and Maximum Likelihood analyses of plastid DNA-sequences. We find that one of the seven genera, the fynbos-endemic Acrosanthes, does not belong to the Aizooideae, but is an ancient sister-lineage to the subfamilies Mesembryanthemoideae & Ruschioideae. Galenia and Plinthus are embedded inside Aizoon and Aizoanthemum is polyphyletic. The Namibian endemic Tetragonia schenckii is sister to Tribulocarpus of the Sesuvioideae. For the Aizooideae, we explored their possible age by means of relaxed Bayesian dating and used Bayesian Binary MCMC reconstruction of ancestral areas to investigate their area of origin. Early diversification occurred in southern Africa in the Eocene-Oligocene, with a split into a mainly African lineage and an Eurasian-Australasian-African-South American lineage. These subsequently radiated in the early Miocene. For Tetragonia, colonisation of Australasia via long-distance dispersal from Eurasia gave rise to the Australasian lineage from which there were subsequent dispersals to South America and Southern Africa. Despite the relatively old age of the Aizooideae, more than half the species have radiated since the Pleiocene, coinciding with the large and rapid diversification of the Ruschioideae. The lineage made up of Tetragonia schenckii &Tribulocarpus split from the remainder of the Sesuvioideae already in the mid Oligocene and its disjunct distribution between Namibia and north-east Africa may be the result of a previously wider distribution within an early Arid African flora. Our reconstruction of ancestral character-states indicates that the expanding keels giving rise to hygrochastic fruits originated only once, i.e. after the split of the Sesuvioideae from the remainder of the Aizoaceae and that they were subsequently lost many times. Variously winged and spiky fruits, adapted to dispersal by wind and animals, have evolved independently in the Aizooideae and the Sesuvioideae. There is then a greater diversity of dispersal systems in the earlier lineages than in the Mesembryanthemoideae and Ruschioideae, where dispersal is mainly achieved by rain.

A phylogenetic hypothesis for the recently diversified Ruschieae (Aizoaceae) in southern Africa.

The Ruschieae is a large tribe of about 1600 species of succulent perennials. They form a major component of the arid parts of the Greater Cape Floristic Region, both in numbers of species and in their density of coverage. So far phylogenetic relationships within the tribe have been unresolved, largely through the paucity of variable molecular characters and this is ascribed to the tribe's recent and rapid radiation. Our phylogeny is based on 10 chloroplast gene regions and represents a nearly complete sampling of the 100 currently recognised genera of the Ruschieae. These chloroplast regions yielded relatively few phylogenetically informative characters, consequently providing only limited resolution in and poor support for many parts of the phylogeny. Nevertheless, for the first time, we provide well-supported evidence that taxa with mostly mesomorphic, often ephemeral leaves and weakly persistent fruits form a basal grade of lineages in the Ruschieae. These lineages subtend a large polytomy of taxa with almost exclusively xeromorphic, persistent leaves and strongly persisting fruits. Among the basal grade of lineages, those occurring within the winter-rainfall region typically shed their leaves or form (at least partly) a protective, dry sheath around the apical bud during the dry summer months, as a means of escaping the summer drought. This contrasts with taxa of the basal grade from outside the winter-rainfall region, in which the leaves persist. Our results show that, in both strongly and weakly persistent fruits, specialised characteristics of the fruit evolved repeatedly and so these structures are highly homoplasious. Perhaps as a consequence of repeated changes towards increased persistence and specialisation of leaves and fruits, several clades show little morphological cohesion. However, as in other groups in the Cape Flora, most clades in the Ruschieae represent regional groupings. Our analysis of sequences of the nuclear gene 'chloroplast-expressed glutamine synthetase' (ncpGS) revealed extensive paralogy within the Ruschieae, but found an intact reading frame in all its members. More data on the cytology of the Ruschieae is needed to evaluate whether the paralogy observed is due to gene duplication or polyploidy.