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Droughts negatively impact forests by reducing growth and increasing\uddefoliation leading to forest dieback as the climate becomes warmer and drier. However,\udthe timing and severity of droughts determine how differently or intensively water shortage\udaffects primary (shoot and leaf formation) and secondary growth (stem radial growth based\udon tree-ring widths). We compare the impact of two severe droughts (2005, 2012),\udshowing different climatic characteristics on the growth responses of three Mediterranean\udholm oak stands in northeastern Spain. We also quantify climate trends and drought\udseverity. Then, we use remote sensing data to infer how those droughts impacted forest\udproductivity. Both droughts were characterized by warm and dry spring conditions leading\udto reduced budburst, low shoot production, asynchrony in primary growth and decreased\udproductivity and scarce radial growth, particularly in 2005. However, defoliation peaked in\ud2012 when radial growth showed minimum values and early spring and late summer\udtemperatures reached maximum values. We discuss how uncoupled and resilient are the\udresponses of primary and secondary growth to drought. Finally, these findings are used to\udgain insight into the drought-related drivers of defoliation in Spanish holm oak forests.We acknowledge support by the CSIC Open Access Publication Initiative through its Unit of Information Resources for Research (URICI).Peer reviewe

Timing of Drought Triggers Distinct Growth Responses in Holm Oak: Implications to Predict Warming-Induced Forest Defoliation and Growth Decline