The oaks of western Eurasia: Traditional classifications and evidence from two nuclear markers

Phylogenetic relationships among species of Quercus (oaks) from western Eurasia including the western part of the Himalayas are examined for the first time. Based on ITS and 5S-IGS data three major infrageneric groups are recognized for western Eurasia: the cerroid, iliciod, and roburoid oaks. While individuals of the cerroid and ilicoid groups cluster according to their species, particularly in the 5S-IGS analyses, individuals of species of roburoid oaks do not cluster with exception of Quercus pontica. The Cypriot endemic Quercus alnifolia belongs to the ilicoid oaks, in contrast to traditional views placing it within the cerroid oaks. Based on all ITS data available, the groups identified for western Eurasia can be integrated into a global in frageneric framework for Quercus. The Ilex group is resurrected as a well-defined group that comprises taxa traditionally placed into six subsections of Q. sects. Cerris and Lepidobalanus (white oaks) sensu Camus. Phylogenetic reconstructions suggest two major lineages within Quercus, each consisting of three infrageneric groups. Within the first lineage, the Quercus group (roburoid oaks in western Eurasia) and the Lobatae group evolved by "budding" as is reflected by incomplete lineage sorting, high variability within groups, and low differentiation among groups. The groups of the second lineage, including the Cyclobalanopsis, Cerris (cerroid oaks in western Eurasia), and Ilex (ilicoid oaks in western Eurasia) groups, evolved in a more tree-like fashion.