An updated infrageneric classification of the oaks: review of previous taxonomic schemes and synthesis of evolutionary patterns
In this paper, we review major classification schemes proposed for oaks by John Claudius Loudon, Anders Sandoe Orsted, William Trelease, Otto Karl Anton Schwarz, Aimee Antoinette Camus, Yuri Leonardovich Menitsky, and Kevin C. Nixon. Classifications of oaks (Fig. 1) have thus far been based entirely on morphological characters. They differed profoundly from each other because each taxonomist gave a different weight to distinguishing characters; often characters that are homoplastic in oaks. With the advent of molecular phylogenetics our view has considerably changed. One of the most profound changes has been the realisation that the traditional split between the East Asian subtropical to tropical subgenus Cyclobalanopsis and the subgenus Quercus that includes all other oaks is artificial. The traditional concept has been replaced by that of two major clades, each comprising three infrageneric groups: a Palearctic-Indomalayan clade including Group Ilex (Ilex oaks), Group Cerris (Cerris oaks) and Group Cyclobalanopsis (cycle-cup oaks), and a predominantly Nearctic clade including Group Protobalanus (intermediate or golden cup oaks), Group Lobatae (red oaks) and Group Quercus (white oaks, with most species in America and some 30 species in Eurasia). The main morphological feature characterising these phylogenetic lineages is pollen morphology, a character overlooked in traditional classifications. This realisation, along with the now available (molecular-)phylogenetic framework, opens new avenues for biogeographic, ecological and evolutionary studies and a re-appraisal of the fossil record. We provide an overview about recent advances in these fields and outline how the results of these studies contribute to the establishment of a unifying systematic scheme of oaks. Ultimately, we propose an updated classification of Quercus recognising two subgenera with eight sections. This classification considers morphological traits, molecular-phylogenetic relationships, and the evolutionary history of one of the most important temperate woody plant genera.\n\nO_FIG O_LINKSMALLFIG WIDTH=156 HEIGHT=200 SRC=\"FIGDIR/small/168146_fig1.gif\" ALT=\"Figure 1\">\nView larger version (49K):\norg.highwire.dtl.DTLVardef@d5a05org.highwire.dtl.DTLVardef@91295forg.highwire.dtl.DTLVardef@3338fcorg.highwire.dtl.DTLVardef@e24a58_HPS_FORMAT_FIGEXP M_FIG O_FLOATNOFigure 1:C_FLOATNO Classification schemes for Quercus from Loudon to Nixon (see Appendix). Colour coding denotes the actual systematic affiliation of species included in each taxon: of the Old World or mid-latitude clade section Cyclobalanopsis (cycle-cup oaks, yellow), section Cerris (Cerris oaks; orange), and section Ilex (Ilex oaks; green); and of the New World or high-latitude clade section Quercus (white oaks s.str.; blue), sections Virentes (cyan) and Ponticae (dark blue), section Protobalanus (intermediate oaks; purple), and section Lobatae (red oaks; red). Colour gradients are proportional, i.e. reflect the proportion of species with different systematic affiliation included in each taxon. Note: Menitsky (1984) and Trelease (1924) only treated the Eurasian and American oaks, respectively, and provided classifications in (nearly) full agreement with current phylogenies.\n\nC_FIG