Molecular phylogenetics and morphology reveal the Plettkea lineage including several members of Arenaria and Pycnophyllopsis to be a clade of 21 South American species nested within Stellaria (Caryophyllaceae, Alsineae)

Abstract

Caryophyllaceae with a cushion-like life form occur with a large number of species at the higher altitudes of the Andes (3500-5000 m) and have evolved convergently in several different lineages. Based on molecular phylogenetic analysis it is shown that members of the former genera Plettkea and Pycnophyllopsis, but also certain species previously classified as Arenaria constitute a subclade nested within the monophyletic genus Stellaria. Both plastid (trnK-matK-psbA + trnL-F) and nuclear (nrITS) trees converged on such a highly supported 'Plettkea' clade. Morphologically, the members of the 'Plettkea' subclade of Stellaria are further characterized by reduced to completely absent petals and seeds with a more or less conspicuous tuberculate testa. This clade is described as S. sect. Plettkea (Mattf.) Montesinos & Borsch. Species-level relationships within S. sect. Plettkea are also congruently inferred by plastid and nuclear genomic compartments, with three further sublineages recognized: Altogether, our detailed taxonomic revision showed that the 'Plettkea' clade in fact constitutes an Andean radiation of 21 species within Stellaria, four of which are described as new to science. Earlier treatments indicated just a few species with a putative placement. The results of this investigation underscore the importance of fieldwork and integrated molecular-morphological approaches to assess the species diversity in Andean plant groups. In addition to the phylogenetic analysis, we provide a taxonomic backbone including all names and types, descriptions and information on distribution and ecology and a key for identification. Regarding the next relatives of the S. sect. Plettkea clade, our plastid trees depict the 'Nitentes' clade of Stellaria as sister, whereas nrITS instead suggests a sister group relationship of the 'Nitentes' with the speciose 'Larbreae' clade. Our inferred relationships of major clades further deviate from published molecular trees by indicating an early branching position of the 'Petiolares' clade.