RESUMEN
Increasingly frequent severe drought events are pushing Mediterranean forests to unprecedented responses. Lack of management leads to dense forests that are highly susceptible to drought stress, potentially resulting in extensive dieback and increased vulnerability to other disturbances. Forest treatments like thinning and slash burning reduce competition for resources and have the potential to enhance tree growth and vigor and minimize tree vulnerability to drought. Here, we used tree rings to study the growth and physiological response of black pine (Pinus nigra) to drought in northeastern Spain under different treatments, including two thinning intensities (light and heavy, with 10% and 40% basal area reduction, respectively) followed by two understory treatments (clearing alone and in combination with slash burning), resulting in a research design of four treatments plus an untreated control with three replicates. Specifically, we studied basal area increment (BAI), resilience indices, and intrinsic water use efficiency (iWUE) using carbon and oxygen isotope composition (δ13C and δ18O in tree-ring cellulose) before and after treatments. Our results showed that BAI and resistance to drought increased in the heavy-thin (burned and unburned) and light-thin burned units. Resilience increased in the burned units regardless of the thinning intensity, while recovery was not affected by treatment. Slash burning additionally increased BAI in the light-thin and resistance and resilience in the heavy-thin units compared with clearing alone. The stable isotope analysis revealed a minor effect of treatments on δ13C and δ18O. No change in iWUE among treatments was presumably linked to a proportional increase in both net CO2 assimilation and stomatal conductance, which particularly increased in the heavy-thin (burned and unburned) and light-thin burned units, indicating that these trees were the least affected by drought. This study shows that management approaches aimed at reducing wildfire hazard can also increase the vigor of dominant trees under drought stress. By reducing competition both from the overstory and the understory, thinning followed by clearing alone or in combination with slash burning promotes tree growth and vigor and increases its resistance and resilience to drought.
RESUMEN
Prescribed burning is used to reduce fuel hazard but underburning can damage standing trees. The effect of burning on needle terpene storage, a proxy for secondary metabolism, in fire-damaged pines is poorly understood despite the protection terpenes confer against biotic and abiotic stressors. We investigated variation in needle terpene storage after burning in three Mediterranean pine species featuring different adaptations to fire regimes. In two pure-stands of Pinus halepensis Mill. and two mixed-stands of Pinus sylvestris L. and Pinus nigra ssp. salzmanni (Dunal) Franco, we compared 24 h and 1 year post-burning concentrations with pre-burning concentrations in 20 trees per species, and evaluated the relative contribution of tree fire severity and physiological condition (δ13C and N concentration) on temporal terpene dynamics (for mono- sesqui- and diterpenes). Twenty-four hours post-burning, monoterpene concentrations were slightly higher in P. halepensis than at pre-burning, while values were similar in P. sylvestris. Differently, in the more fire-resistant P. nigra monoterpene concentrations were lower at 24 h, compared with pre-burning. One year post-burning, concentrations were always lower compared with pre- or 24 h post-burning, regardless of the terpene group. Mono- and sesquiterpene variations were negatively related to pre-burning δ13C, while diterpene variations were associated with fire-induced changes in needle δ13C and N concentration. At both post-burning times, mono- and diterpene concentrations increased significantly with crown scorch volume in all species. Differences in post-burning terpene contents as a function of the pine species' sensitivity to fire suggest that terpenic metabolites could have adaptive importance in fire-prone ecosystems in terms of flammability or defence against biotic agents post-burning. One year post-burning, our results suggest that in a context of fire-induced resource availability, pines likely prioritize primary rather than secondary metabolism. Overall, this study contributes to the assessment of the direct and indirect effects of fire on pine terpene storage, providing valuable information about their vulnerability to biotic and abiotic stressors throughout time.