Drought legacy in mature spruce alleviates physiological stress during recurrent drought

Forest ecosystems are facing severe and prolonged droughts with delayed recovery, known as "drought legacy". This study presents positive legacy effects following a long-term, experimental drought and subsequent recovery in a mature mixed Norway spruce and European beech forest. Approximately 50 mature trees were exposed to five consecutive years of summer drought by completely excluding growing season precipitation from May 2014 to June 2019. Experimental drought recovery started in July 2019, after which the trees received natural precipitation. Taking advantage of the natural summer drought of 2022, following the unique long-term experimental drought, we investigated how drought legacy affects tree physiological responses to recurrent drought. The long-term experimental drought resulted in a 60% reduction in spruce leaf area, which was still reduced by 30% 4 years after the drought release. This slow recovery and associated reduced water use resulted in higher soil water availability under spruce during the 2022 drought, leading to significantly reduced physiological drought stress: about two times higher predawn leaf water potential, leaf gas exchange and sap flow density in legacy spruce compared to previous controls. Furthermore, neighbouring beech, displaying no leaf area reduction during the experimental drought, also had higher predawn leaf water potential and leaf gas exchange during the 2022 drought compared to previous controls, likely benefitting from the reduced water use of spruce. The slow recovery of spruce leaf area as a pronounced drought legacy effect proved advantageous for trees in alleviating physiological stress and overcoming future drought events.