Forests of Quercus petraea, sometimes of Quercus robur, or of either or both oaks and Fagus sylvatica, accompanied by a cortège of sub-Atlantic and submeridional acidophile species (e.g. Pinus sylvestris, Carpinus betulus), developed in central and southern-central Europe, outside of the main Atlantic-influenced range of the Quercion. Quercus-dominated acidophilous forests of the western Hercynian ranges and their periphery, developed in more Atlantic conditions as substitution forests of the Luzulo-Fagion beech forests have been associated with them because of a shared contingent of sub-Atlantic accompanying species and similarities in overall appearance.