Effects of Drought on Xylem Anatomy and Water-Use Efficiency of Two Co-Occurring Pine Species

Exploring how drought influences growth, performance, and survival in different species is crucial to understanding the impacts of climate change on forest ecosystems. Here, we investigate the responses of two co-occurring pines (Pinus nigra and Pinus sylvestris) to interannual drought in east-central Spain by dendrochronological and wood anatomical features integrated with isotopic ratios of carbon (δ13C) and oxygen (δ18O) in tree rings. Our results showed that drought induces both species to allocate less carbon to build tracheid cell-walls but increases tracheid lumen diameters, particularly in the transition wood between early and latewood, potentially maximizing hydraulic conductivity but reducing resistance to embolism at a critical phase during the growing season. The thicker cell-wall-to-lumen ratio in P. nigra could imply that its xylem may be more resistant to bending stress and drought-induced cavitation than P. sylvestris. In contrast, the higher intrinsic water-use efficiency (iWUE) in P. sylvestris suggests that it relies more on a water-saving strategy. Our results suggest that narrower cell-walls and reduced growth under drought are not necessarily linked to increased iWUE. At our site P. nigra showed a higher growth plasticity, grew faster and was more competitive than P. sylvestris. In the long term, these sustained differences in iWUE and anatomical characters could affect forest species performance and composition, particularly under increased drought stress.