Pollen characteristics used in determination and systematics of Quercus (Fagaceae): new data and verification of previous concepts

Quercus (oaks) is a large and important genus of woody angiosperms occurring in a wide range of environments and often occupying a dominant position in temperate forests of the Northern Hemisphere or being a minor component in some subtropical and tropical forests. A reliable determination of fossil dispersed oak pollen requires combined light, scanning electron (SEM) and transmission electron microscopy (TEM) studies and relevant comparative data on extant oak species. We provide SEM and TEM data on 22 extant Quercus spp., representing all sections of the genus. The main objectives of our study were to clarify the nature of the so-called geniculus, a characteristic equatorial bulge in the exine of the colpus found in many oak species, and to test previous hypotheses about the structural basic units of the exine in Quercus. Our ontogenetic study and oxidative experiment involving several Quercus spp. and one Trigonobalanus sp. resulted in a more accurate interpretation of the nature of the geniculus and exine elements, their different developmental states and, therefore, their more reliable application in the systematics and evolution of the genus Quercus and Fagaceae in general.