K. Uzunova et al., ANATOMICAL CHANGES AND EVOLUTIONARY TRENDS IN THE FOLIAR EPIDERMIS OFEXTANT AND FOSSIL EURO-MEDITERRANEAN OAKS (FAGACEAE), Plant systematics and evolution, 204(3-4), 1997, pp. 141-159
The differentiation of foliar epidermis structures, cuticles, stomata,
and indumentum of most of the extant Euro-Mediterranean taxa of Querc
us have been studied by light and scanning electron microscopy, and co
mpared with other genera and fossil material. Results allow the recogn
ition of the subgg. Sclerophyllodrys, Cerris and Quercus, and of vario
us sections and even individual species. The cyclocytic stomata type,
regarded as plesiomorphic, is found in the geologically oldest represe
ntatives of the family (Dryophyllum, resp. Eotrigonobalanus) and in th
e relatively old, evergreen to semi-evergreen species of Quercus subgg
. Sclerophyllodrys and Cerris. These taxa also have relatively thick a
nd smooth cuticles. Anomocytic stomata, thinner cuticles and the prese
nce of crystalline wax flakes are regarded as apomorphic features and
characterize the deciduous taxa of subg. Quercus. These changes are in
terpreted as ecological adaptations to the changing climates from the
Tertiary to the present. An analysis of the various character combinat
ions found in macrofossils and the often poor species separation in Qu
ercus suggest that hybridization has played a major role in the evolut
ion of the Euro-Mediterranean oaks.