Losses of leaf area owing to herbivory and early senescence in three tree species along a winter temperature gradient.

In temperate climates, evergreen leaves have to survive throughout low temperature winter periods. Freezing and chilling injuries can lead to accelerated senescence of part of the leaf surface, which contributes to a reduction of the lifespan of the photosynthetic machinery and of leaf lifetime carbon gain. Low temperatures are also associated with changes in foliar chemistry and morphology that affect consumption by herbivores. Therefore, the severity of foliar area losses caused by accelerated senescence and herbivory can change along winter temperature gradients. The aim of this study is to analyse such responses in the leaves of three evergreen species (Quercus ilex, Q. suber and Pinus pinaster) along a climatic gradient. The leaves of all three species presented increased leaf mass per area (LMA) and higher concentrations of structural carbohydrates in cooler areas. Only the two oak species showed visible symptoms of damage caused by herbivory, this being less intense at the coldest sites. The leaves of all three species presented chlorotic and necrotic spots that increased in size with leaf age. The foliar surface affected by chlorosis and necrosis was larger at the sites with the coldest winters. Therefore, the effects of the winter cold on the lifespan of the photosynthetic machinery were contradictory: losses of leaf area due to accelerated senescence increased, but there was a decrease in losses caused by herbivory. The final consequences for carbon assimilation strongly depend on the exact timing of the appearance of the damage resulting from low temperature and grazing by herbivores.