Cineraria L. (Senecioneae, Asteraceae) - its taxonomy, phylogeny, phytogeography and conservation

Abstract

This study aimed to investigate the phylogeny of Cineraria L. to elucidate its position in the tribe Senecioneae (Asteraceae), to delimit its generic and species boundaries, explore species relationships (infrageneric structure) and produce a monograph of the genus. It also aimed to examine its distribution, phytogeographic affinities, levels of rarity and endemism in Cineraria, to identify some of the factors contributing to rarity in Cineraria and to highlight implications for conservation. Phylogenetic analyses were performed using morphological and molecular (DNA sequence) characters to elucidate relationships within the genus and between Cineraria and selected related genera in the subtribe Senecioninae. The phylogenetic species concept was applied - suites of diagnostic characters were used to characterise species. The phenetic approach, using Cluster Analysis and Principal Coordinates Analysis, was applied to investigate variation in two highly variable species, C. deltoidea Sond. and C. lobata L’Hér. Species distributions were mapped and the number of species per degree square was plotted for southern Africa to identify centres of diversity and endemism. Rare species were identified and categorised according to Rabinowitz’s criteria of geographic range, habitat specificity and local population size. Cineraria now has a more homogenous generic concept, characterised as herbs or subshrubs with palmately veined leaves, radiate, calyculate capitula, penicillate style apices and obovate, compressed cypselae with two distinct margins or wings and a substantial carpopodium. Eleven species have been removed from the genus and two new genera, Bolandia Cron and Oresbia Cron & B.Nord. have been established to accommodate three of the species. Two species have been reassigned to and another reinstated in Senecio L. The affinities of five species remain unresolved. Cineraria now comprises 35 species with four new subspecies and two new varieties recognised. Eight species have been placed in synonymy and five new species have been described during this revision. The status of C. deltoidea as a single, highly variable species, widespread throughout the eastern mountains of Africa, has been confirmed. Cineraria lobata has been shown to be a very variable species and the geographic and morphological variation has been formally (and informally) recognised. No infrageneric classification has been applied to Cineraria as a robust phylogeny of all the species has yet to be hypothesised. A southern African origin for Cineraria in the Western and/or Eastern Cape is postulated, based on the current distribution of the sister genus Bolandia and Cineraria mollis DC. iv Cineraria appears to have undergone rapid speciation fairly recently, as indicated by the lack of variation in the molecular data analysed, with reticulate evolution playing an important role in its evolutionary history (as seen by the lack of congruence between the chloroplast and nuclear DNA sequences). The centre of diversity of Cineraria is the KwaZulu-Natal Midlands, part of the Maputaland-Pondoland Centre of endemism in southern Africa. Cineraria has an afromontane affinity, and fifteen species endemic to specific mountains or regions of endemism and five near-endemics have been identified in Cineraria. Eleven species have been shown to be rare (in the sense of low abundance, restricted range and high habitat specificity), however only five are considered to be threatened as indicated by IUCN Red Data Criteria. However, at least seven species are Data Deficient and require further investigation. Causes of rarity in Cineraria are linked to narrow habitat specificity, particularly soil or rock type and/or altitudinal range.