In and out of the Qinghai-Tibet Plateau: divergence time estimation and historical biogeography of the large arctic-alpine genus Saxifraga L.

Aim Geologically dynamic areas often harbour remarkable levels of biodiversity. Among other factors, mountain building is assumed to be a precondition for species radiation, and yet, the potential role of immigration as a source of biodiversity prior to radiation is often neglected. Here, we studied the biogeographical history of the large genus Saxifraga to unravel the role played by the Qinghai-Tibet Plateau (QTP) for the diversification of this genus and to understand factors that have led to the establishment of high biodiversity in and around this region. Location QTP and surrounding mountain ranges and worldwide distribution range of Saxifraga. Methods Using a total of 420 taxa (321 ingroup taxa) comprising more than 60% of extant Saxifraga species, we studied the evolutionary history of Saxifraga by performing phylogenetic analyses (maximum likelihood and Bayesian inference on nuclear ITS and plastid trnL-trnF, matK sequences), divergence time estimation (using uncorrelated log-normal clock models and four fossil constraints in beast) and ancestral range estimation (using BioGeoBEARS). Results Saxifraga originated in North America around 74 (64-83) Ma, dispersed to South America and northern Asia during its early diversification and colonized Europe and the QTP region by the Late Eocene. The QTP region was colonized several times independently, followed in some lineages by rapid radiations, temporally coinciding with recent uplifts of the Hengduan Mountains at the southeastern fringe of the QTP. Subsequently, several lineages dispersed out of Tibet. Main conclusions Immigration, recent rapid radiation and lineage persistence were all important processes for the establishment of a rich species stock of Saxifraga in the QTP region. Because floristic exchanges between the neighbouring areas and the QTP region were bi-directional, the spatio-temporal evolution of Saxifraga contrasts with the 'out of QTP' pattern, which has often been assumed for northern temperate plants.