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Understanding ecosystem responses to compound disturbance regimes and the influence of specific sequences of events in determining ecosystem shifts remains a challenge. We use a modelling framework for Mediterranean-type ecosystems to assess the effects of fire–drought interactions on long-term vegetation dynamics and to identify disturbance-driven changes in trait composition (tree seeder vs. tree resprouter prevalence) and ecosystem state (forest vs. non-forest). Changes in tree seeder and the tree resprouter dominance show nonlinear, threshold-type trends over gradients of increasing compound disturbance frequency. Vegetation composition thresholds mostly occur in a narrow range of the compound fire–drought disturbance space. Additionally, trait compositional switches and the likelihood of sudden changes in ecosystem state are promoted by fire-drought interactions. Distinct sequences of disturbance events cause vegetation transitions, disrupting ecosystem resilience, even under moderate recurrence of individual disturbances. An extreme drought year followed by one or two large fire events promotes shifts from resprouter- to seeder dominance. Contrastingly, a large crown fire followed by an extreme drought promotes changes from seeder to resprouter dominance. This disturbance sequence is also a mechanism strong enough to trigger sudden shifts in ecosystem state (from forest to non-forest). Synthesis. Thresholds of change in vegetation composition occur over a narrow range of the modelled gradients of compound fire-drought recurrence, and the loss of ecosystem resilience is contingent on particular sequences of disturbance events. Overall, our findings highlight that disturbance interactions define the relative location of tipping points in ecosystem state, and that effects and feedbacks of compound disturbance regimes increase the long-term likelihood of sudden ecosystem shifts and, therefore, uncertainty in predicting vegetation state.

Mediterranean forests and fire regimes are closely intertwined. Global change is likely to alter both forest dynamics and wildfire activity, ultimately threatening the provision of ecosystem services and posing greater risks to society. In this paper we evaluate future wildfire behavior by coupling climate projections with simulation models of forest dynamics and wildfire hazard. To do so, we explore different forest management scenarios reflecting different narratives related to EU forestry (promotion of carbon stocks, reduction of water vulnerability, biomass production and business-as-usual) under the RCP 4.5 and RCP 8.5 climate pathways in the period 2020-2100. We used as a study model pure submediterranean Pinus nigra forests of central Catalonia (NE Spain). Forest dynamics were simulated from the 3rd National Forest Inventory (143 stands) using SORTIE-nd software based on climate projections under RCPs 4.5 and 8.5. The climate products were also used to estimate fuel moisture conditions (both live and dead) and wind speed. Fuel parameters and fire behavior were then simulated, selecting crown fire initiation potential and rate of spread as key indicators. The results revealed consistent trade-offs between forest dynamics, climate and wildfire. Despite the clear influence exerted by climate, forest management modulates fire behavior, resulting in different trends depending on the climatic pathway. In general, the maintenance of current practices would result in the highest rates of crown fire activity, while management for water vulnerability reduction is postulated as the best alternative to surmount the increasingly hazardous conditions envisaged in RCP 8.5., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2022 The Authors. Published by Elsevier Ltd.. All rights reserved.)

Minimum water potential (Ψ min ) is a key variable for characterizing dehydration tolerance and hydraulic safety margins (HSMs) in plants. Ψ min is usually estimated as the absolute minimum tissue Ψ experienced by a species, but this is problematic because sample extremes are affected by sample size and the underlying probability distribution. We compare alternative approaches to estimate Ψ min and assess the corresponding uncertainties and biases; propose statistically robust estimation methods based on extreme value theory (EVT); and assess the implications of our results for the characterization of hydraulic risk. Our results show that current estimates of Ψ min and HSMs are biased, as they are strongly affected by sample size. Because sampling effort is generally higher for species living in dry environments, the differences in current Ψ min estimates between these species and those living under milder conditions are partly artefactual. When this bias is corrected using EVT methods, resulting HSMs tend to increase substantially with resistance to embolism across species. Although data availability and representativeness remain the main challenges for proper determination of Ψ min , a closer look at Ψ distributions and the use of statistically robust methods to estimate Ψ min opens new ground for characterizing plant hydraulic risks., (© 2021 The Authors. New Phytologist © 2021 New Phytologist Foundation.)

Efforts to develop mechanistic tree growth models are hindered by the uncertainty of whether and when tree growth responses to environmental factors are driven by carbon assimilation or by biophysical limitations of wood formation. In this study, we used multiannual weekly wood-formation monitoring of two conifer species (Larix decidua and Picea abies) along a 900 m elevational gradient in the Swiss Alps to assess the biophysical effect of temperature and water potential on wood formation. To this end, we developed a model that simulates the effect of water potential on turgor-driven cambial division, modulated by the effect of temperature on enzymatic activity. The model reproduced the observed phenology of tracheid production, as well as intra- and interannual tracheid production dynamics of both species along the elevational gradient, although interannual model performance was lower. We found that temperature alone explains the onset of tracheid production, yet water potential appears necessary to predict the ending and the total amount of tracheids produced annually. We conclude that intra-annual cambial activity is strongly constrained by both temperature and water potential at all elevations, independently of carbon assimilation. At the interannual scale, biophysical constraints likely interact with other factors., (© 2020 The Authors. New Phytologist © 2020 New Phytologist Trust.)

The extent to which water availability can be used to predict the enlargement and final dimensions of xylem conduits remains an open issue. We reconstructed the time course of tracheid enlargement in Pinus sylvestris trees in central Spain by repeated measurements of tracheid diameter on microcores sampled weekly during a 2 yr period. We analyzed the role of water availability in these dynamics empirically through time-series correlation analysis and mechanistically by building a model that simulates daily tracheid enlargement rate and duration based on Lockhart's equation and water potential as the sole input. Tracheid enlargement followed a sigmoid-like time course, which varied intra- and interannually. Our empirical analysis showed that final tracheid diameter was strongly related to water availability during tracheid enlargement. The mechanistic model was calibrated and successfully validated (R 2  = 0.92) against the observed tracheid enlargement time course. The model was also able to reproduce the seasonal variations of tracheid enlargement rate, duration and final diameter (R 2  = 0.84-0.99). Our results support the hypothesis that tracheid enlargement and final dimensions can be modeled based on the direct effect of water potential on turgor-driven cell expansion. We argue that such a mechanism is consistent with other reported patterns of tracheid dimension variation., (© 2019 The Authors New Phytologist © 2019 New Phytologist Trust.)

Understanding ecosystem vulnerability is essential in risk management to anticipate disasters. While valuable efforts have been made to characterize vulnerability components (exposure, sensitivity, and response capacity) at particular ecosystem stages, there is still a lack of context-specific studies accounting for the temporal dimension of vulnerability. In this study, we developed a procedure to identify the main natural dynamics of monospecific and mixed forests and to assess the variations of sensitivity and response capacity to fire along successional dynamics. In the procedure, we generated forest chronosequences by summarizing the dynamics between consecutive surveys of permanent plots into a set of longer successional trajectories represented in a multidimensional space. Then, we calculated several variables of sensitivity and response capacity to fire of forest stages associated with each trajectory and we assessed their variation along succession. The procedure was applied to Mediterranean forests in Spain dominated by a pine species poorly adapted to severe crown fires. We found that forest vulnerability components varied differently among successional trajectories, which depended on the composition and structure of their initial stages and the environmental context in which they occurred. Autosuccessional dynamics of pine forests showed relatively low sensitivity to fire along trajectories. However, their response capacity was related to the changes in shrub cover. In contrast, diversifying dynamics showed an increasing sensitivity to fire, but also a higher response capacity the greater the functional diversity along succession. These results highlight the need for considering the temporal dimension of vulnerability in risk management and the importance of assessing sensitivity and response capacity as independent components of vulnerability that can be modified through management at critical forest stages., (Copyright © 2019 Elsevier Ltd. All rights reserved.)

Fire regimes are shifting or are expected to do so under global change. Current fire suppression is not able to control all wildfires, and its capability to do so might be compromised under harsher climate conditions. Alternative fire management strategies may allow to counteract predicted fire trends, but we lack quantitative tools to evaluate their potential effectiveness at the landscape scale. Here, we sought to quantify changes in fire regimes induced after the implementation of different fire management strategies. We developed and applied a new version of the model MEDFIRE in Catalonia (Mediterranean region of ~32,000 km 2 in NE Spain). We first projected burnt area from 2016 to 2100 resulting from climate change under the Representative Concentration Pathway 8.5 scenario of HadGEM-CC model and under current fire suppression levels. We then evaluated the impacts of four fire management strategies: 'Let it burn', fixed effort of prescribed burning with two different spatial allocations, and adaptive prescribed burning dynamically adjusting efforts according to recent past fires. Results predicted the emergence of novel climates associated with similar barometric configurations to current conditions but with higher temperatures (i.e. hot wind events). These novel climates led to an increase in burnt area, which was partially counteracted by negative fire-vegetation feedbacks. All prescribed burning scenarios decreased the amount of high-intensity fires and extreme fire events. The 'Let it burn' strategy, although less costly, was not able to reduce the extent of high-intensity fires. The adaptive prescribed burning scenario resulted in the most cost-efficient strategy. Our results provide quantitative evidence of fire management effectiveness, and bring to light key insights that could guide the design of fire policies fit for future novel climate conditions. We propose adaptive landscape management focused on the reduction of fire negative impacts rather than on the elimination of this disturbance from the system., (Copyright © 2019 Elsevier B.V. All rights reserved.)