Leaf variations in Elaeagnus angustifolia related to environmental heterogeneity.

Elaeagnus angustifolia (Russian olive) is a Eurasian tree that has become naturalized and has invaded zones along watercourses in many arid and semiarid regions of the world. These habitats are characterized by vertical environmental gradients, thus trees must develop some plasticity to adapt to the wide range of site conditions. This study was undertaken to test the hypothesis that variations in leaf anatomy and morphology of E. angustifolia reflect their adaptability to the differences in the microclimate that occur within the canopy of single trees. Foliar architecture, blade and petiole epidermal and internal anatomy were examined in leaves at different canopy positions and related to environmental conditions. Upper sun-leaves are exposed to higher solar irradiance and lower air humidity and are smaller, more slender and thicker than the lower, half-exposed and shade-leaves. Color varies between the leaves at different levels, from silvery grey-green in the upper strata, to dark green in the lower one. Bicolor is more evident in half-exposed and shaded leaves. When compared with the lower half-exposed and shade-leaves, the upper leaves of E. angustifolia have a greater areole density, a higher mesophyll proportion and stomatal density. Trichomes are multicellular, pedestalled, stellate-branched or peltate and their form and density can be associated with leaf color and appearance. The slender petioles of the upper leaves have proportionally more epidermis, collenchyma and phloem and less parenchyma and xylem than those of lower leaves, when observed in transverse sections. Foliar morphological and anatomical variability in E. angustifolia may be considered an adaptive advantage that enables leaves to develop and function in habitats marked by strong variations of solar radiation, air temperature and humidity.