SOURCES OF THE ARCTIC FLORA: ORIGINS OF ARCTIC SPECIES IN RANUNCULUS AND RELATED GENERA.

The arctic biome is a relatively young ecosystem with ~2300 species of vascular plants. We studied the genus Ranunculus as an example of the origin and evolution of the arctic flora. For this purpose we used molecular phylogenetic and clock analyses based on evaluation of nuclear ITS and chloroplast matK-trnK DNA sequences in 194 taxa of Ranunculus and closely related genera. Taxa occurring in the Arctic arose form seven phylogenetic lineages of Ranunculus and also in the genera Coptidium and Halerpestes. Two clades of Ranunculus are species-rich in the Arctic, i.e., Ranunculus sect. Ranunculus and R. sect. Auricomus (both from R. subg. Ranunculus), but this is due to a number of arctic "microtaxa" morphologically barely separate from R. acris in the former clade and the widely agamospermic species complex of R. auricomus in the latter. Lineages with species adapted to wetlands or aquatic habitats are significant groups represented in the arctic flora (R. subg. Ranunculus sectt. Flammula and Hecatonia/Xanthobatrachium, R. subg. Batrachium, genus Coptidium) but show no clear signs of radiation in the Arctic or the northern boreal zone, except for sectt. Hecatonia/Xanthobatrachium, with R. hyperboreus and R. sceleratus subsp. reptabundus. Astonishingly few of the otherwise numerous lineages of Ranunculus with distributions in the higher mountain systems of Eurasia and North America have acted as "founding sources" for the arctic flora. The only clear example is that of the arctic-alpine R. glacialis and the Beringian R. chamissonis from the lineage of subg. R. sectt. Aconitifolii/Crymodes, although there might be others in sect. Auricomus not recovered in the current molecular data. Lineages that gave rise to arctic taxa diverged from each other from the early Miocene (R. glacialis/R. chamissonis, Coptidium, lineages in Halerpestes) and continued at an even rate throughout the Tertiary. There are no signs that the intense climate changes of the late Pliocene and the Quaternary substantially accelerated or impeded diversification in Ranunculus. Only the crown group split of R. acris and its relatives is clearly of Quaternary age. A detailed comparison concerning morphology, karyology, and life form excludes fundamental differences between taxa of Ranunculus in the Arctic and their respective closest relatives in regions south of it. Ecological traits, e.g., preferences for dry or moist soils or growth in open and sheltered conditions, also do not differ between arctic and nonarctic̣ taxa. Migration into the Arctic thus started from different phylogenetic lineages and at different times, without development of obvious special traits in the adaptation to arctic environments. This recurrent pattern in Ranunculus differs from that seen in other arctic genera, e.g., Artemisia, in which special traits of adaptation to arctic environments are found. In Ranunculus, the origin of the open arctic biome primarily favored range expansions of taxa/species already adapted to wet habitats in cold areas and depending on rapid dispersal.