Increased forest vulnerability is being reflected as more widespread and severe drought-induced decline episodes. In particular, the Mediterranean area is revealing a high susceptibility to phenomena of loss in tree vitality across species. Within tree species, oaks (Quercus spp.) are experiencing extensive decline in many countries. However, in the wake of the so-called "oak decline phenomenon", the attention on these species has generally been limited. In this paper, we review the current available literature on oakdecline cases reported within the Mediterranean Basin, with particular remark for those occurred in Italy and Spain. More specifically our main aims were to: (i) provide an update on the patterns and mechanisms of decline by focusing on tree-ring and wood-anatomical variables; (ii) provide some hints for improving the resistance and resilience of oak stands experiencing decline. Our review reveals that drought is reported as the main driver triggering oak decline within the Mediterranean Basin, although other causes (i.e., increasing temperature, pathogens attack or excessive stand density) could exacerbate decline. In most reported cases, drought induced a substantial reduction of growth and changes in some wood anatomical properties. Indeed, growth decline prior death is also indicated as an early-warning signal of impending death. In ring-porous oak species, declining trees were often characterized by a very low production of latewood and a decrease in lumen area of the widest earlywood vessels, suggesting a potential reduction of hydraulic conductivity. Moreover, hydraulic dysfunction is reported as the main cause of decline. Finally, regarding management actions that should be considered for improving the resilience of declining stands and preserve the species-specific stand composition, it could be useful to shorten the rotation period of coppice stands or promoting their gradual conversion towards high forests, and favoring more drought-resistant species should also be considered. In addition, regeneration prior to regeneration cuts should be improved by anticipating seed dispersal or by planting oak seedlings obtained from local germoplasm.

Drought stress causes forest dieback that is often explained by two interrelated mechanisms, namely hydraulic failure and carbon starvation. However, it is still unclear which functional and structural alterations, related to these mechanisms, predispose to dieback. Here we apply a multi-proxy approach for the characterization of tree structure (radial growth, wood anatomy) and functioning (δ13C, δ18O and non-structural carbohydrates (NSCs)) in tree rings before and after drought-induced dieback. We aim to discriminate which is the main mechanism and to assess which variables can act as early-warning proxies of drought-triggered damage. The study was tailored in southern Italy in two forests (i.e., San Paolo (SP) and Oriolo (OR)) where declining and non-declining trees of a ring-porous tree species (Quercus frainetto Ten.) showing anisohydric behavior coexist. Both stands showed growth decline in response to warm and dry spring conditions, although the onset of dieback was shifted between them (2002 in SP and 2009 in OR). Declining trees displayed a sharp growth drop after this onset with reductions of 49% and 44% at SP and OR sites, respectively. Further, contrary to what we expected, declining trees showed a lower intrinsic water-use efficiency compared with non-declining trees after the dieback onset (with reductions of 9.7% and 5.6% at sites SP and OR, respectively), due to enhanced water loss through transpiration, as indicated by the lower δ18O values. This was more noticeable at the most drought-affected SP stand. Sapwood NSCs did not differ between declining and non-declining trees, indicating no carbon starvation in affected trees. Thus, the characterized structural and functional alterations partially support the hydraulic failure mechanism of dieback. Finally, we show that growth data are reliable early-warning proxies of drought-triggered dieback.

The objective of this study is to globally assess the effects of atmospheric nitrogen deposition and climate, associated with rising levels of atmospheric CO2 , on the variability of carbon isotope discrimination (Δ(13) C), and intrinsic water-use efficiency (iWUE) of angiosperm and conifer tree species. Eighty-nine long-term isotope tree-ring chronologies, representing 23 conifer and 13 angiosperm species for 53 sites worldwide, were extracted from the literature, and used to obtain long-term time series of Δ(13) C and iWUE. Δ(13) C and iWUE were related to the increasing concentration of atmospheric CO2 over the industrial period (1850-2000) and to the variation of simulated atmospheric nitrogen deposition and climatic variables over the period 1950-2000. We applied generalized additive models and linear mixed-effects models to predict the effects of climatic variables and nitrogen deposition on Δ(13) C and iWUE. Results showed a declining Δ(13) C trend in the angiosperm and conifer species over the industrial period and a 16.1% increase of iWUE between 1850 and 2000, with no evidence that the increased rate was reduced at higher ambient CO2 values. The temporal variation in Δ(13) C supported the hypothesis of an active plant mechanism that maintains a constant ratio between intercellular and ambient CO2 concentrations. We defined linear mixed-effects models that were effective to describe the variation of Δ(13) C and iWUE as a function of a set of environmental predictors, alternatively including annual rate (Nrate ) and long-term cumulative (Ncum ) nitrogen deposition. No single climatic or atmospheric variable had a clearly predominant effect, however, Δ(13) C and iWUE showed complex dependent interactions between different covariates. A significant association of Nrate with iWUE and Δ(13) C was observed in conifers and in the angiosperms, and Ncum was the only independent term with a significant positive association with iWUE, although a multi-factorial control was evident in conifers.

Hydraulic theory suggests that tall trees are at greater risk of drought-triggered death caused by hydraulic failure than small trees. In addition the drop in growth, observed in several tree species prior to death, is often interpreted as an early-warning signal of impending death. We test these hypotheses by comparing size, growth, and wood-anatomy patterns of living and now-dead trees in two Italian oak forests showing recent mortality episodes. The mortality probability of trees is modeled as a function of recent growth and tree size. Drift-diffusion-jump (DDJ) metrics are used to detect early-warning signals. We found that the tallest trees of the anisohydric Italian oak better survived drought contrary to what was predicted by the theory. Dead trees were characterized by a lower height and radial-growth trend than living trees in both study sites. The growth reduction of now-dead trees started about 10 years prior to their death and after two severe spring droughts during the early 2000s. This critical transition in growth was detected by DDJ metrics in the most affected site. Dead trees were also more sensitive to drought stress in this site indicating different susceptibility to water shortage between trees. Dead trees did not form earlywood vessels with smaller lumen diameter than surviving trees but tended to form wider latewood vessels with a higher percentage of vessel area. Since living and dead trees showed similar competition we did not expect that moderate thinning and a reduction in tree density would increase the short-term survival probability of trees.

Forest decline induced by climate change is a global phenomenon that affects many tree species, mainly in drought-prone areas as the Mediterranean region. In southern Italy, several oak species have shown decline symptoms and elevated mortality since the 2000s due to drought stress. However, it remains to be answered whether decline occurred alone or whether a pathogen was also involved. To this aim, we compared two coexisting oak species in a forest located in southern Italy which are assumed to be less (Quercus cerris) and more tolerant to drought (Quercus pubescens). We sampled fifteen couples of neighboring declining (D) and non-declining (ND) trees of both species. Wood cores were taken from all trees to perform dendrochronological analyses to detect the decline onset and link it to potential climatic drivers. Carbon isotope ratios (d13C) were analyzed in wood of the two vigor classes to compare their water-use efficiency. Phytophthora presence was also assessed in soil samples from ten D-ND couples of trees per species. The oak species most affected by drought-induced decline in terms of leaf shedding and mortality was Q. cerris, i.e., the least tolerant to drought. In both species, the D trees showed a reduced growth rate compared with ND trees from 2000 onward when drought and warming intensified. Q. pubescens showed higher growth sensitivity to precipitation, temperature and drought than Q. cerris. This sensitivity to climate was magnified in D trees whose growth decreased in response to warm and dry conditions during the prior winter and the late summer. The Q. pubescens D trees were more efficient in their water use than ND trees before the growth divergence between D and ND trees amplified. In the studied area, Phytophthora quercina was isolated from 40% of the sampled trees, and tended to be more frequent amongst ND than amongst D trees. Our data suggests that droughts and warm summer conditions triggered oak decline. The high prevalence of P. quercina in the studied area warrants further study as a potential predisposing factor.