Increasing carbon discrimination rates and depth of water uptake favor the growth of Mediterranean evergreen trees in the ecotone with temperate deciduous forests

Tree populations at the low-altitudinal or -latitudinal limits of species' distributional ranges are predicted to retreat toward higher altitudes and latitudes to track the ongoing changes in climate. Studies have focused on the climatic sensitivity of the retreating species, whereas little is known about the potential replacements. Competition between tree species in forest ecotones will likely be strongly influenced by the ecophysiological responses to heat and drought. We used tree-ring widths and δ¹³ C and δ¹⁸ O chronologies to compare the growth rates and long-term ecophysiological responses to climate in the temperate-Mediterranean ecotone formed by the deciduous Fagus sylvatica and the evergreen Quercus ilex at the low altitudinal and southern latitudinal limit of F. sylvatica (NE Iberian Peninsula). F. sylvatica growth rates were similar to those of other southern populations and were surprisingly not higher than those of Q. ilex, which were an order of magnitude higher than those in nearby drier sites. Higher Q. ilex growth rates were associated with high temperatures, which have increased carbon discrimination rates in the last 25 years. In contrast, stomatal regulation in F. sylvatica was proportional to the increase in atmospheric CO₂ . Tree-ring δ¹⁸ O for both species were mostly correlated with δ¹⁸ O in the source water. In contrast to many previous studies, relative humidity was not negatively correlated with tree-ring δ¹⁸ O but had a positive effect on Q. ilex tree-ring δ¹⁸ O. Furthermore, tree-ring δ¹⁸ O decreased in Q. ilex over time. The sensitivity of Q. ilex to climate likely reflects the uptake of deep water that allowed it to benefit from the effect of CO₂ fertilization, in contrast to the water-limited F. sylvatica. Consequently, Q. ilex is a strong competitor at sites currently dominated by F. sylvatica and could be favored by increasingly warmer conditions.