LAUSR

Tree-ring intra-annual stable isotopes (δ13C and δ18O) are powerful tools for revealing plant ecophysiological responses to climatic extremes. We analyzed interannual and fine-scale intra-annual variability of tree-ring δ13C and δ18O in Pinus massoniana from southeastern China to explore environmental drivers and potential trade-offs between the main physiological controls. We show that wet season relative humidity (May-October RH) drove interannual variability of δ18O and intra-annual variability of tree-ring δ18O. It also drove intra-annual variability of tree-ring δ13C, whereas interannual variability was mainly controlled by February-May temperature and September-October RH. Furthermore, intra-annual tree-ring δ18O variability was larger during wet years compared to dry years, whereas δ13C variability was lower during wet years compared to dry years. As a result of these differences in intra-annual variability amplitude, process-based models (we used the Roden model for δ18O and the Farquhar model for δ13C) captured the intra-annual δ18O pattern better in wet years compared to dry years, whereas intra-annual δ13C pattern was better simulated in dry years compared to wet years. This result suggests a potential asymmetric bias in process-based models in capturing the interplay of the different mechanistic processes (i.e., isotopic source and leaf-level enrichment) operating in dry versus wet years. We therefore propose an intra-annual conceptual model considering a dynamic trade-off between the isotopic source and leaf-level enrichment in different tree-ring parts to understand how climate and ecophysiological processes drive intra-annual tree-ring stable isotopic variability under humid climate conditions.