Tree drought-mortality risk depends more on intrinsic species resistance than on stand species diversity.

Increasing tree diversity is considered a key management option to adapt forests to climate change. However, the effect of species diversity on a forest's ability to cope with extreme drought remains elusive. In this study, we assessed drought tolerance (xylem vulnerability to cavitation) and water stress (water potential), and combined them into a metric of drought-mortality risk (hydraulic safety margin) during extreme 2021 or 2022 summer droughts in five European tree diversity experiments encompassing different biomes. Overall, we found that drought-mortality risk was primarily driven by species identity (56.7% of the total variability), while tree diversity had a much lower effect (8% of the total variability). This result remained valid at the local scale (i.e within experiment) and across the studied European biomes. Tree diversity effect on drought-mortality risk was mediated by changes in water stress intensity, not by changes in xylem vulnerability to cavitation. Significant diversity effects were observed in all experiments, but those effects often varied from positive to negative across mixtures for a given species. Indeed, we found that the composition of the mixtures (i.e., the identities of the species mixed), but not the species richness of the mixture per se, is a driver of tree drought-mortality risk. This calls for a better understanding of the underlying mechanisms before tree diversity can be considered an operational adaption tool to extreme drought. Forest diversification should be considered jointly with management strategies focussed on favouring drought-tolerant species.

Hydraulic plasticity and water use regulation act to maintain the hydraulic safety margins of Mediterranean trees in rainfall exclusion experiments.

Hydraulic failure due to xylem embolism has been identified as one of the main mechanisms involved in drought-induced forest decline. Trees vulnerability to hydraulic failure depends on their hydraulic safety margin (HSM). While it has been shown that HSM globally converges between tree species and biomes, there is still limited knowledge regarding how HSM can adjust locally to varying drought conditions within species. In this study, we relied on three long-term partial rainfall exclusion experiments to investigate the plasticity of hydraulic traits and HSM for three Mediterranean tree species (Quercus ilex L., Quercus pubescens Willd., and Pinus halepensis Mill.). For all species, a homeostasis of HSM in response to rainfall reduction was found, achieved through different mechanisms. For Q. ilex, the convergence in HSM is attributed to the adjustment of both the turgor loss point (Ψtlp) and the water potential at which 50% of xylem conductivity is lost due to embolism (P50). In contrast, the maintenance of HSM for P. halepensis and Q. pubescens is related to its isohydric behavior for the first and leaf area adjustment for the latter. It remains to be seen whether this HSM homeostasis can be generalized and if it will be sufficient to withstand extreme droughts expected in the Mediterranean region.

Isohydricity and hydraulic isolation explain reduced hydraulic failure risk in an experimental tree species mixture.

Species mixture is promoted as a crucial management option to adapt forests to climate change. However, there is little consensus on how tree diversity affects tree water stress, and the underlying mechanisms remain elusive. By using a greenhouse experiment and a soil-plant-atmosphere hydraulic model, we explored whether and why mixing the isohydric Aleppo pine (Pinus halepensis, drought avoidant) and the anisohydric holm oak (Quercus ilex, drought tolerant) affects tree water stress during extreme drought. Our experiment showed that the intimate mixture strongly alleviated Q. ilex water stress while it marginally impacted P. halepensis water stress. Three mechanistic explanations for this pattern are supported by our modeling analysis. First, the difference in stomatal regulation between species allowed Q. ilex trees to benefit from additional soil water in mixture, thereby maintaining higher water potentials and sustaining gas exchange. By contrast, P. halepensis exhibited earlier water stress and stomatal regulation. Second, P. halepensis trees showed stable water potential during drought, although soil water potential strongly decreased, even when grown in a mixture. Model simulations suggested that hydraulic isolation of the root from the soil associated with decreased leaf cuticular conductance was a plausible explanation for this pattern. Third, the higher predawn water potentials for a given soil water potential observed for Q. ilex in mixture can-according to model simulations-be explained by increased soil-to-root conductance, resulting from higher fine root length. This study brings insights into the mechanisms involved in improved drought resistance of mixed species forests.

Plant hydraulics at the heart of plant, crops and ecosystem functions in the face of climate change.

Plant hydraulics is crucial for assessing the plants' capacity to extract and transport water from the soil up to their aerial organs. Along with their capacity to exchange water between plant compartments and regulate evaporation, hydraulic properties determine plant water relations, water status and susceptibility to pathogen attacks. Consequently, any variation in the hydraulic characteristics of plants is likely to significantly impact various mechanisms and processes related to plant growth, survival and production, as well as the risk of biotic attacks and forest fire behaviour. However, the integration of hydraulic traits into disciplines such as plant pathology, entomology, fire ecology or agriculture can be significantly improved. This review examines how plant hydraulics can provide new insights into our understanding of these processes, including modelling processes of vegetation dynamics, illuminating numerous perspectives for assessing the consequences of climate change on forest and agronomic systems, and addressing unanswered questions across multiple areas of knowledge.

Drivers and implications of the extreme 2022 wildfire season in Southwest Europe.

Wildfire is a common phenomenon in Mediterranean countries but the 2022 fire season has been extreme in southwest Europe (Portugal, Spain and France). Here we provide a preliminary but comprehensive analysis of 2022's wildfire season in southwest Europe. Burned area has exceeded the 2001-2021 median by a factor of 52 in some regions and large wildfires (>500 ha) started to occur in June-July, earlier than the traditional fire season. These anomalies were associated with record-breaking values of fuel dryness, atmospheric water demand and pyrometeorological conditions. Live fuel moisture content was below the historical minima for almost 50 % of the season in some regions. A few large wildfires were responsible for 82 % of the burned area and, in turn, 47 % of the area burned occurred in protected areas. Shrublands, transitional woodlands and conifer forests (but not eucalypt plantations) were the land cover types most affected by extreme fires. As climate change intensifies, we can expect such fire seasons to become the new normal in large parts of the continent, potentially leading to major negative impacts on rural economies. These results highlight the need for landscape level fuel management also in protected areas, to avoid fire-induced biodiversity losses and landscape scale degradation. Our results have important policy implications and indicate that fire prevention should be explicitly addressed within continental forest legislation and strategies.

Plant hydraulic modelling of leaf and canopy fuel moisture content reveals increasing vulnerability of a Mediterranean forest to wildfires under extreme drought.

Fuel moisture content (FMC) is a crucial driver of forest fires in many regions world-wide. Yet, the dynamics of FMC in forest canopies as well as their physiological and environmental determinants remain poorly understood, especially under extreme drought. We embedded a FMC module in the trait-based, plant-hydraulic SurEau-Ecos model to provide innovative process-based predictions of leaf live fuel moisture content (LFMC) and canopy fuel moisture content (CFMC) based on leaf water potential ( ψ Leaf ). SurEau-Ecos-FMC relies on pressure-volume (p-v) curves to simulate LFMC and vulnerability curves to cavitation to simulate foliage mortality. SurEau-Ecos-FMC accurately reproduced ψ Leaf and LFMC dynamics as well as the occurrence of foliage mortality in a Mediterranean Quercus ilex forest. Several traits related to water use (leaf area index, available soil water, and transpiration regulation), vulnerability to cavitation, and p-v curves (full turgor osmotic potential) had the greatest influence on LFMC and CFMC dynamics. As the climate gets drier, our results showed that drought-induced foliage mortality is expected to increase, thereby significantly decreasing CFMC. Our results represent an important advance in our capacity to understand and predict the sensitivity of forests to wildfires.

Prediction of regional wildfire activity in the probabilistic Bayesian framework of Firelihood.

Modeling wildfire activity is crucial for informing science-based risk management and understanding the spatiotemporal dynamics of fire-prone ecosystems worldwide. Models help disentangle the relative influences of different factors, understand wildfire predictability, and provide insights into specific events. Here, we develop Firelihood, a two-component, Bayesian, hierarchically structured, probabilistic model of daily fire activity, which is modeled as the outcome of a marked point process: individual fires are the points (occurrence component), and fire sizes are the marks (size component). The space-time Poisson model for occurrence is adjusted to gridded fire counts using the integrated nested Laplace approximation (INLA) combined with the stochastic partial differential equation (SPDE) approach. The size model is based on piecewise-estimated Pareto and generalized Pareto distributions, adjusted with INLA. The Fire Weather Index (FWI) and forest area are the main explanatory variables. Temporal and spatial residuals are included to improve the consistency of the relationship between weather and fire occurrence. The posterior distribution of the Bayesian model provided 1,000 replications of fire activity that were compared with observations at various temporal and spatial scales in Mediterranean France. The number of fires larger than 1 ha across the region was coarsely reproduced at the daily scale, and was more accurately predicted on a weekly basis or longer. The regional weekly total number of larger fires (10-100 ha) was predicted as well, but the accuracy degraded with size, as the model uncertainty increased with event rareness. Local predictions of fire numbers or burned areas also required a longer aggregation period to maintain model accuracy. The estimation of fires larger than 1 ha was also consistent with observations during the extreme fire season of the 2003 unprecedented heat wave, but the model systematically underrepresented large fires and burned areas, which suggests that the FWI does not consistently rate the actual danger of large fire occurrence during heat waves. Firelihood enabled a novel analysis of the stochasticity underlying fire hazard, and offers a variety of applications, including fire hazard predictions for management and projections in the context of climate change.

Increased likelihood of heat-induced large wildfires in the Mediterranean Basin.

Wildfire activity is expected to increase across the Mediterranean Basin because of climate change. However, the effects of future climate change on the combinations of atmospheric conditions that promote wildfire activity remain largely unknown. Using a fire-weather based classification of wildfires, we show that future climate scenarios point to an increase in the frequency of two heat-induced fire-weather types that have been related to the largest wildfires in recent years. Heat-induced fire-weather types are characterized by compound dry and warm conditions occurring during summer heatwaves, either under moderate (heatwave type) or intense (hot drought type) drought. The frequency of heat-induced fire-weather is projected to increase by 14% by the end of the century (2071-2100) under the RCP4.5 scenario, and by 30% under the RCP8.5, suggesting that the frequency and extent of large wildfires will increase throughout the Mediterranean Basin.

Recent climate hiatus revealed dual control by temperature and drought on the stem growth of Mediterranean Quercus ilex.

A better understanding of stem growth phenology and its climate drivers would improve projections of the impact of climate change on forest productivity. Under a Mediterranean climate, tree growth is primarily limited by soil water availability during summer, but cold temperatures in winter also prevent tree growth in evergreen forests. In the widespread Mediterranean evergreen tree species Quercus ilex, the duration of stem growth has been shown to predict annual stem increment, and to be limited by winter temperatures on the one hand, and by the summer drought onset on the other hand. We tested how these climatic controls of Q. ilex growth varied with recent climate change by correlating a 40-year tree ring record and a 30-year annual diameter inventory against winter temperature, spring precipitation, and simulated growth duration. Our results showed that growth duration was the best predictor of annual tree growth. We predicted that recent climate changes have resulted in earlier growth onset (-10 days) due to winter warming and earlier growth cessation (-26 days) due to earlier drought onset. These climatic trends partly offset one another, as we observed no significant trend of change in tree growth between 1968 and 2008. A moving-window correlation analysis revealed that in the past, Q. ilex growth was only correlated with water availability, but that since the 2000s, growth suddenly became correlated with winter temperature in addition to spring drought. This change in the climate-growth correlations matches the start of the recent atmospheric warming pause also known as the 'climate hiatus'. The duration of growth of Q. ilex is thus shortened because winter warming has stopped compensating for increasing drought in the last decade. Decoupled trends in precipitation and temperature, a neglected aspect of climate change, might reduce forest productivity through phenological constraints and have more consequences than climate warming alone.

Photosynthetic sensitivity to drought varies among populations of Quercus ilex along a rainfall gradient.

Drought frequency and intensity are expected to increase in the Mediterranean as a consequence of global climate change. To understand how photosynthetic capacity responds to long-term water stress, we measured seasonal patterns of stomatal (SL), mesophyll (MCL) and biochemical limitations (BL) to net photosynthesis (Amax) in three Quercus ilex (L.) populations from sites differing in annual rainfall. In the absence of water stress, stomatal conductance (gs), maximum carboxylation capacity (Vcmax), photosynthetic electron transport rate (Jmax) and Amax were similar among populations. However, as leaf predawn water potential (Ψl,pd) declined, the population from the wettest site showed steeper declines in gs, Vcmax, Jmax and Amax than those from the drier sites. Consequently, SL, MCL and BL increased most steeply in response to decreasing Ψl,pd in the population from the wettest site. The higher sensitivity of Amax to drought was primarily the result of stronger stomatal regulation of water loss. Among-population differences were not observed when gs was used instead of Ψl,pd as a drought stress indicator. Given that higher growth rates, stature and leaf area index were observed at the wettest site, we speculate that hydraulic architecture may explain the greater drought sensitivity of this population. Collectively, these results highlight the importance of considering among-population differences in photosynthetic responses to seasonal drought in large scale process-based models of forest ecosystem function.