We manipulated nutrient availability and competing vegetation to investigate the phenotypic plasticity of leaf morphology and nutrient content of saplings and seedlings of the evergreen holm oak Quercus ilex subsp. ballota in response to nutrient pulses. Different factorial experimental designs were established with N and/or P pulse fertilization as factors in a field experiment in Catalonia (northeast Spain) and with P fertilization, soil type, and competitive situation as factors in a pot experiment. In the field, P fertilization decreased leaf mass per area (LMA) 9–13% in the 3 y following the fertilization application. No significant effect was observed from variations in N supply in any leaf cohort. In the pot experiment, P fertilization increased the leaf size, pre-dawn leaf water content, and root growth and decreased the LMA. P fertilization strongly increased P leaf concentrations, as in the field experiment. Neighbour competition induced increases in LMA in the second leaf cohort and in leaf density and size in both leaf cohorts; it also reduced root growth and tended to increase P leaf concentrations. Soil type had a significant effect on individual leaf area, leaf mass, and LMA, the latter being higher in siliceous soils than in calcareous and artificial soils. The pattern of change in leaf thickness across the different soil types was the opposite of that for leaf P concentrations, P contents per unit of leaf area, and leaf density. These results point to a morphological leaf plasticity in response to P availability, competitive situation, and soil type. The relations between the changes in leaf morphology, leaf nutrient content, root growth, and pre-dawn leaf water content changed depending on the factor that induced them. However, leaf morphological traits changed in combination with other plant traits such as root growth and nutrient contents in order to balance resource availability and production capacity under different environmental conditions, such as changing degrees of neighbour competition or water availability. This plasticity of sclerophyllous leaf traits in response to nutrient pulses makes Quercus ilex subsp. ballota well suited to adapt to the unpredictable changes and stress situations likely to result from current and immediate future climate and other changes in the Mediterranean region.
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Vol. 13 • No. 2