Reduced growth sensitivity to climate in bark-beetle infested Aleppo pines: Connecting climatic and biotic drivers of forest dieback

Drought and bark-beetle infestation are major and often interconnected drivers of forest dieback and tree death. These two stressors may interact and accelerate forest mortality, since warmer and drier conditions boost beetle attacks and reduce tree growth. However, the way in which drought and bark-beetle infestation interact and affect declining or dying trees is still poorly understood. To disentangle the long-term interaction between the two stressors, we quantified radial growth (basal area increment), resin production, mortality and intrinsic water-use efficiency (iWUE) inferred from wood-carbon isotope discrimination. We compared trees infested and recently killed by two abundant bark beetle species (Orthotomicus erosus and Tomicus piniperda) with non-infested living trees in a drought-prone Aleppo pine plantation. Growth and iWUE showed similar values in infested and non-infested trees. Since bark-beetle-infested trees did not grow less than non-infested trees, our results did not support the hypothesis of higher costs of resin production at the expense of stem-wood formation. Radial growth was enhanced by cool and wet winter conditions prior to the growing season. However, infested trees showed lower growth responsiveness than did non-infested trees with respect to this climatic driver of growth. Infested trees also showed a lower resin-duct production two years prior to death than non-infested trees. The growth responsiveness to climate should be characterized in bark-beetle-infested trees, since a weak correlation between climate and growth can be regarded as a predisposing factor of infestation-induced tree death. Such reduced responsiveness to climate stress could be linked to the tree vulnerability to beetle attacks in drought-prone forests.