Sequence of plant responses to droughts of different timescales: lessons from holm oak (Quercus ilex) forests

The functional traits of plants in regions of the world with a Mediterranean climate have been shaped to tolerate periods of water deficit. These species are adapted to summer droughts but may not be able to cope with future increases in drought intensity, duration, and/or frequency. Here, we review the mechanisms and traits of drought resistance and recovery of the well-studied holm oak (Quercus ilex), which we propose as a model species for Mediterranean-type ecosystems. Our aim was to understand the differences and links between the responses of Q. ilex to summer droughts, extreme droughts, and long-term drought experiments. A main goal was to provide an integral picture of drought responses across organisational and temporal scales for identifying the most relevant processes that are likely to contribute to determining the future of Mediterranean vegetation. Evidence from long-term drought experiments showed that acclimation processes from the molecular (e.g. epigenetic changes) to the ecosystem level (e.g. reductions in stand density) mitigate the effects of drought. Changes in leaf morphology and hydraulics, leaf-to-shoot allometry, and root functioning are among the key mechanisms for overcoming increasing drought. The duration of drought determines its severity in terms of canopy loss and stem mortality. Although Q. ilex can vigorously resprout after such episodes, its resilience may be subsequently reduced. In the future, higher frequency of return of extreme droughts will challenge thus the capacity of these forests to recover. The insights provided by this review of the complex interplay of processes that determine the response of trees to droughts of different duration, intensity, and frequency will also help us to understand the likely responses of other resprouting angiosperms in seasonally dry ecosystems that share similar functional traits with Q. ilex.