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This insight article comments on: Ziegler C, Cochard, H, Stahl C, Bastien Gérard LF, Goret J, Heuret P, Levionnois S, Maillard P, Bonal D, Coste S. 2024. Residual water losses mediate the trade-off between growth and drought survival across saplings of 12 tropical rainforest tree species with contrasting hydraulic strategies. Journal of Experimental Botany 75, 4128–4147. [ABSTRACT FROM AUTHOR]

Polylepis trees occur throughout the Andean mountain region, and it is the tree genus that grows at the highest elevation worldwide. In the humid Andes where moisture is rarely limiting, Polylepis trees must adapt to extreme environmental conditions, especially rapid fluctuations in temperature, ultraviolet radiation and vapor pressure deficit (VPD). However, Polylepis ' water-use patterns remain largely unknown despite the importance of understanding their response to microclimate variation to determine their capacity to maintain resilience under future environmental change. We conducted a study in a Polylepis reticulata Kunth forest in the Ecuadorian Andes to evaluate its tree water-use dynamics and to identify the main environmental drivers of transpiration. Tree sap flow was monitored simultaneously with soil volumetric water content (VWC) and microclimate during 2 years for trees growing in forest edge and interior locations. We found that sap flow was primarily controlled by VPD and that VWC exerted a secondary role in driving sap flow dynamics. The highest values for sap flow rates were found when VPD > 0.15 kPa and VCW < 0.73 cm3 cm−3, but these threshold conditions only occurred during brief periods of time and were only found in 11% of our measurements. Moreover, these brief windows of more favorable conditions occurred more frequently in forest edge compared with forest interior locations, resulting in edge trees maintaining 46% higher sap flow compared with interior trees. Our results also suggest that P. reticulata has a low stomatal control of transpiration, as the sap flow did not decline with increasing VPD. This research provides valuable information about the potential impacts of projected future increases in VPD due to climate change on P. reticulata water-use dynamics, which include higher sap flow rates leading to greater transpirational water loss due to this species' poor stomatal control. [ABSTRACT FROM AUTHOR]

Regional-level applications of dynamic vegetation models are challenging because they need to accommodate the variation in plant functional diversity, which requires moving away from broadly defined functional types. Different approaches have been adopted in the last years to incorporate a trait-based perspective into modeling exercises. A common parametrization strategy involves using trait data to represent functional variation between individuals while discarding taxonomic identity. However, this strategy ignores the phylogenetic signal of trait variation and cannot be employed when predictions for specific taxa are needed, such as in applications to inform forest management planning. An alternative strategy involves adapting the taxonomic resolution of model entities to that of the data source employed for large-scale initialization and estimating functional parameters from available plant trait databases, adopting diverse solutions for missing data and non-observable parameters. Here we report the advantages and limitations of this second strategy according to our experience in the development of MEDFATE (version 2.9.3), a novel cohort-based and trait-enabled model of forest dynamics, for its application over a region in the western Mediterranean Basin. First, 217 taxonomic entities were defined according to woody species codes of the Spanish National Forest Inventory. While forest inventory records were used to obtain some empirical parameter estimates, a large proportion of physiological, morphological, and anatomical parameters were matched to measured plant traits, with estimates extracted from multiple databases and averaged at the required taxonomic level. Estimates for non-observable key parameters were obtained using meta-modeling and calibration exercises. Missing values were addressed using imputation procedures based on trait covariation, taxonomic averages or both. The model properly simulated observed historical changes in basal area, with a performance similar to an empirical model trained for the same region. While strong efforts are still required to parameterize trait-enabled models for multiple taxa, and to incorporate intra-specific trait variability, estimation procedures such as those presented here can be progressively refined, transferred to other regions or models and iterated following data source changes by employing automated workflows. We advocate for the adoption of trait-enabled and population-structured models for regional-level projections of forest function and dynamics. [ABSTRACT FROM AUTHOR]

Forest models are instrumental for understanding and projecting the impact of climate change on forests. A considerable number of forest models have been developed in the last decades. However, few systematic and comprehensive model comparisons have been performed in Europe that combine an evaluation of modelled carbon and water fluxes and forest structure. We evaluate 13 widely used, state‐of‐the‐art, stand‐scale forest models against field measurements of forest structure and eddy‐covariance data of carbon and water fluxes over multiple decades across an environmental gradient at nine typical European forest stands. We test the models' performance in three dimensions: accuracy of local predictions (agreement of modelled and observed annual data), realism of environmental responses (agreement of modelled and observed responses of daily gross primary productivity to temperature, radiation and vapour pressure deficit) and general applicability (proportion of European tree species covered). We find that multiple models are available that excel according to our three dimensions of model performance. For the accuracy of local predictions, variables related to forest structure have lower random and systematic errors than annual carbon and water flux variables. Moreover, the multi‐model ensemble mean provided overall more realistic daily productivity responses to environmental drivers across all sites than any single individual model. The general applicability of the models is high, as almost all models are currently able to cover Europe's common tree species. We show that forest models complement each other in their response to environmental drivers and that there are several cases in which individual models outperform the model ensemble. Our framework provides a first step to capturing essential differences between forest models that go beyond the most commonly used accuracy of predictions. Overall, this study provides a point of reference for future model work aimed at predicting climate impacts and supporting climate mitigation and adaptation measures in forests. [ABSTRACT FROM AUTHOR]

Regional-level applications of dynamic vegetation models are challenging because they need to accommodate the variation in plant functional diversity, which requires moving away from broadly-defined functional types. Different approaches have been adopted in the last years to incorporate a trait-based perspective into modeling exercises. A common parametrization strategy involves using trait data to represent functional variation between individuals while discard taxonomic identity, but this strategy ignores the phylogenetic signal of trait variation and cannot be employed when predictions for specific taxa are needed, as in applications to inform forest management planning. An alternative strategy involves adapting the taxonomic resolution of model entities to that of the data source employed for large-scale initialization and estimating functional parameters from available plant trait databases while adopting alternative solutions for missing data and non-observable parameters. Here we report the advantages and limitations of this second strategy according to our experience in the development of MEDFATE (v. 2.8.1), a novel cohort-based and trait-enabled model of forest dynamics, for its application over a region in the Western Mediterranean Basin. First, 217 taxonomic entities were defined according to woody species codes of the Spanish National Forest Inventory. While forest inventory data were used to obtain some empirical parameter estimates, a large proportion of physiological, morphological, and anatomical parameters were mapped to measured plant traits, with estimates extracted from multiple databases and averaged at the required taxonomic level. Estimates for non-observable key parameters were obtained using meta-modeling and calibration exercises. Missing values were filled using imputation procedures based on trait coordination, taxonomic averages or both. The model properly simulated observed historical basal area changes, with a performance similar to an empirical model trained for the same region. While strong efforts are still required to parameterize trait-enabled models for multiple taxa, estimation procedures can be progressively refined, transferred to other regions or models and iterated following data source changes by employing automated workflows. We advocate for the adoption of trait-enabled population-structured models for regional-level projections of forest function and dynamics. [ABSTRACT FROM AUTHOR]

Riparian zones play a fundamental role in regulating the amount of carbon (C) and nitrogen (N) that is exported from catchments. However, C and N removal via soil gaseous pathways can influence local budgets of greenhouse gas (GHG) emissions and contribute to climate change. Over a year, we quantified soil effluxes of carbon dioxide (CO2) and nitrous oxide (N2O) from a Mediterranean riparian forest in order to understand the role of these ecosystems on catchment GHG emissions. In addition, we evaluated the main soil microbial processes that produce GHG (mineralization, nitrification, and denitrification) and how changes in soil properties can modify the GHG production over time and space. Riparian soils emitted larger amounts of CO2 (1.2- 10 gCm-2 d-1) than N2O (0.001-0.2 mgNm-2 d-1) to the atmosphere attributed to high respiration and low denitrification rates. Both CO2 and N2O emissions showed a marked (but antagonistic) spatial gradient as a result of variations in soil water content across the riparian zone. Deep groundwater tables fueled large soil CO2 effluxes near the hillslope, while N2O emissions were higher in the wet zones adjacent to the stream channel. However, both CO2 and N2O emissions peaked after spring rewetting events, when optimal conditions of soil water content, temperature, and N availability favor microbial respiration, nitrification, and denitrification. Overall, our results highlight the role of water availability on riparian soil biogeochemistry and GHG emissions and suggest that climate change alterations in hydrologic regimes can affect the microbial processes that produce GHG as well as the contribution of these systems to regional and global biogeochemical cycles. [ABSTRACT FROM AUTHOR]

Mediterranean riparian zones act as vegetation shelters for several deciduous tree species at the edge of their bioclimatic distribution, e.g. alder ( Alnus glutinosa ), black poplar ( Populus nigra ) or ash ( Fraxinus excelsior ). Current global warming and human induced disturbances may worsen their growing conditions. Under such circumstances, black locust ( Robinia pseudoacacia ) is outcompeting autochthonous tree species. Here, we provide evidences of black locust better growth and water use performance than alder and ash. We compare the temporal and spatial patterns of transpiration and the stem basal area increments of alder, black poplar, common ash and black locust, all of them co-occurring in a mixed riparian Mediterranean forest. Black locust presented the lowest transpiration values per basal area unit (4.0 mm·m −2 ·growing season −1 ). Although tree transpiration was mainly driven by energy availability instead of water, ash transpiration was constrained by water availability at soil water contents below 0.08 cm 3 ·cm −3 . Black locust was the only tree species growing all over the water availability gradient present in the study site, and it did not present any significant difference in sap flow values across this gradient. Furthermore, black locust and black poplar were the species with higher growth-based water use efficiency (5.4 g·cm −1 ·m −3 and 3.6 g·cm −1 ·m −3 , respectively); ash and alder were the less efficient ones (2.8 g·cm −1 ·m −3 and 1.9 g·cm −1 ·m −3 respectively). The good performance of black locust is relevant to understand its great successful invasion of Mediterranean riparian forests, particularly after human-induced disturbances, as forest management. [ABSTRACT FROM AUTHOR]

Riparian zones play a fundamental role in regulating the amount of carbon (C) and nitrogen (N) that is exported from catchments. However, C and N removal via soil gaseous pathways can influence local budgets of greenhouse gases (GHG) emissions and contribute to climate change. Over a year, we quantified soil effluxes of carbon dioxide (CO2) and nitrous oxide (N2O) from a Mediterranean riparian forest in order to understand the role of these ecosystems on catchment GHG emissions. In addition, we evaluated the main soil microbial processes that produce GHG (mineralization, nitrification, and denitrification) and how changes in soil properties can modify the GHG production over time and space. Mediterranean riparian soils emitted large amounts of CO2 to the atmosphere (1.2-10 g C m-2 d-1), but were powerless sources of N2O (0.001-0.2 mg N m-2 d-1) due to low denitrification rates. Both CO2 and N2O emissions showed a marked (but antagonistic) spatial gradient as a result of variations in soil moisture across the riparian zone. Deep groundwater tables fueled large soil CO2 effluxes near the hillslope, while N2O emissions were higher in the wet zones adjacent to the stream channel. However, both CO2 and N2O emissions peaked after spring rewetting events, when optimal conditions of soil moisture, temperature, and N availability favor microbial respiration, nitrification, and denitrification. Overall, our results highlight the role of riparian soils as hotspots of GHG emissions, and suggest that future alterations in hydrologic regimes can affect the microbial processes that produce GHG as well as the contribution of these systems to climate change. [ABSTRACT FROM AUTHOR]

Drought has been shown to reduce soil respiration (SR) in previous studies. Meanwhile, studies of the effect of forest management on SR yielded contrasting results. However, little is known about the combined effect of drought and forest management on SR. To investigate if the drought stress on SR can be mitigated by thinning, we implemented plots of selective thinning and 15% reduced rainfall in a mixed forest consisting of the evergreen Quercus ilex and deciduous Quercus cerrioides; we measured SR seasonally from 2004 to 2007. Our results showed a clear soil moisture threshold of 9%; above this value, SR was strongly dependent on soil temperature, with Q10 of 3·0-3.8. Below this threshold, the relationship between SR and soil temperature weakened. We observed contrasting responses of SR of target oak species to drought and thinning. Reduced rainfall had a strong negative impact on SR of Q. cerrioides, whereas the effect on SR for Q. ilex was marginal or even positive. Meanwhile, selective thinning increased SR of Q. cerrioides, but reduced that of Q. ilex. Overall, our results showed that the negative effect of drought on SR can be offset through selective thinning, but the effect is attenuated with time. [ABSTRACT FROM AUTHOR]

In the last decades, the global interest in the role of forests as carbon sinks has grown, and thus, studies aimed at estimating tree biomass have progressively increased. However, few surveys have focused on young coppices, although they are abundant worldwide in areas regenerating after disturbance (e.g. wildfire, clearcutting). In the Mediterranean Basin, young coppices are very frequent, and most of them are formed by evergreen and deciduous oaks ( Quercus spp.). In this survey, we have studied the biomass allocation patterns of two oaks coexisting in coppices, the evergreen Quercus ilex and the deciduous Quercus cerrioides, comparing them in the light of their different leaf habit, which may influence their physiological performance in the context of climate change. We have also obtained allometric equations for each species and its components, which we have used to calculate the carbon stock in the sampled area, as an insight into the potential of young oak coppices to sequester carbon. The results indicate a higher biomass investment of Q. ilex in the stump and a higher leafiness and allocation to roots in Q. cerrioides. In the light of these differences, the evergreen Q. ilex could be defined as a 'resource-saving' species in comparison with the more 'resource-demanding' Q. cerrioides. The allometric equations were able to predict from 78 to 99 % of the variation in biomass using diameter as predictor variable for all the tree components aside of the stump. Overall carbon stock estimation in the young coppice of our study area was 43.2 Mg ha, of which 62 % is stored belowground. These results highlight the importance of biomass allocation in the belowground compartment in Mediterranean young oak coppices as a temporal carbon sink. Moreover, they provide evidence that these forests may store a relevant amount of carbon, often ignored in forest inventories. [ABSTRACT FROM AUTHOR]

Fire is a major agent involved in landscape transformation and an indirect cause of changes in species composition. Responses to fire may vary greatly depending on life histories and functional traits of species. We have examined the taxonomic and functional responses to fire of eight taxonomic animal groups displaying a gradient of dietary and mobility patterns: Gastropoda, Heteroptera, Formicidae, Coleoptera, Araneae, Orthoptera, Reptilia and Aves. The fieldwork was conducted in a Mediterranean protected area on 3 sites (one unburnt and two burnt with different postfire management practices) with five replicates per site. We collected information from 4606 specimens from 274 animal species. Similarity in species composition and abundance between areas was measured by the Bray-Curtis index and ANOSIM, and comparisons between animal and plant responses by Mantel tests. We analyze whether groups with the highest percentage of omnivorous species, these species being more generalist in their dietary habits, show weak responses to fire (i.e. more similarity between burnt and unburnt areas), and independent responses to changes in vegetation. We also explore how mobility, i.e. dispersal ability, influences responses to fire. Our results demonstrate that differences in species composition and abundance between burnt and unburnt areas differed among groups. We found a tendency towards presenting lower differences between areas for groups with higher percentages of omnivorous species. Moreover, taxa with a higher percentage of omnivorous species had significantly more independent responses of changes in vegetation. High- (e.g. Aves) and low-mobility (e.g. Gastropoda) groups had the strongest responses to fire (higher R scores of the ANOSIM); however, we failed to find a significant general pattern with all the groups according to their mobility. Our results partially support the idea that functional traits underlie the response of organisms to environmental changes caused by fire. [ABSTRACT FROM AUTHOR]

Mediterranean forest growth is constrained by drought and high temperatures during summer. Effects of climate change on these forests depend on how changes in water availability and temperature will take place. A process-based forest growth model, growth of trees is limited by water in the Mediterranean (GOTILWA+), was applied in the Mediterranean region on Quercus ilex, Pinus halepensis, P. pinaster, P. sylvestris and Fagus sylvatica forests. The effects of climate change on growth were analysed, as well as the effect of thinning cycle length, combined with the assumption of different soil depths. Thinning cycle lengths was included because it can affect the response of stands to climatic conditions, and soil depth because this is positively related to soil water-holding capacity and consequently may change the effects of drought. The simulation period covered 140 years (1961–2100). Model results show that leaf area index (LAI) may increase, favoured by the increase of atmospheric CO2, particularly at sites where rainfall is relatively high and climatic conditions not too warm. The predicted increase in temperature significantly influenced mean leaf life span (MLLS). MLLS of F. sylvatica would increase with climate change, implying a longer growing period. Conversely, MLLS of evergreen species would be reduced, accelerating leaf turnover. In general, our results showed a higher production promoted by projected climate change in response to the increasing atmospheric CO2 concentration and rainfall in the region. Temperature increase would have different consequences for production. In F. sylvatica, the longer growing period would promote higher production, particularly when water is not limiting. On the other hand, Q. ilex and Pinus species would expend more carbon in maintaining and producing leaves to replace those lost in increased turnover rate. As expected, access of roots to deeper soil results in an increased final wood yield (FWY) due to an improved water balance that promotes higher transpiration, photosynthesis and growth. In general, the shorter the harvest cycle, the larger the FWY, because of less tree mortality between harvesting events. According to our results, temperature and rainfall may constrain growth during certain periods but if rainfall increases in the future, a positive effect on growth is likely. [Copyright &y& Elsevier]

Summer drought is a characteristic trait of Mediterranean climates and can limit primary production. The holm oak (Quercus ilex L.) forest at the Prades Experimental Complex of Catchments (PECC, NE Spain) was managed by clearcutting until the 1950s. As a result of this practice, the forest has a high density (13000 stems ha−1) that is causing growth stagnation and intense competition for water. A thinning experiment was conducted to release resources, reduce stem competition and improve forest productivity. Minirhizotrons were installed in two plots in each of the thinned and control stands and fine root dynamics quantified to a soil depth of 60 cm over a 2-year period (June 1994–May 1996).Soil water content was higher and soil water penetrated deeper in the soil (30–50 cm) in the thinned plots than in the control plots. In the control plots, soil temperature did not vary substantially with depth (mean = 11.65 °C), whereas soil temperature was higher in shallow horizons compared with deep horizons in the thinned plots. Mean soil temperature was one degree higher in the thinned plots than in the control plots (12.9 versus 11.9 °C). There were always more roots in the top 20 cm of soil in thinned plots than in control plots. In the thinned plots, more than 50% of root density (root no. cm−2) was located in the top 20 cm of soil, whereas root density never reached 50% in the 0–20 cm soil layer of the control plots. Roots longevity varied from 35 to 471 days, and there were significant differences between soil depths. Root longevity was greater in the control plots than in the thinned plots. [ABSTRACT FROM AUTHOR]

• Black locust growth is currently limited by low temperatures in Mediterranean region. • We inverse calibrated GOTILWA + vegetation model from observed sap flow data. • Increasing severity of climate change enhances black locust growth. • Growth enhancement is driven by an increased daily productivity and an extended growing season. Mediterranean riparian forests are comparably humid environments that provide shelter for several broadleaved deciduous tree species at their southernmost distribution margin. The stability of these communities, however, is threatened by climate change as well as invasive tree species, such as black locust (Robinia pseudoacacia L.). So far, black locust's European distribution appears to be mostly limited by low temperatures, but global warming might enhance its growth in colder areas. Moreover, R. pseudoacacia can better access water from the phreatic level than some native non-phreatophytic tree species such as European ash (Fraxinus excelsior L.). In this study, we compare the performance of European ash, a native deciduous tree species at its southernmost distribution border, with the invasive black locust, under a range of climate change projections, in a stand located at N.E. Spain. We first use Bayesian inference to calibrate the GOTILWA + vegetation model against sap flow data for both tree species. We then project each tree species' performance under several climate change scenarios. Our results indicate that increasing temperatures will enlarge black locust's vegetative period, leading to substantially increased annual productivity if the phreatic water table keeps reachable. For European ash, we project a slight increase in productivity, but with higher uncertainty. Our findings suggest that black locust will profit more from global warming than the native European ash, which is concerning because of the already detrimental impact of black locust for the local ecosystems. We conclude that climate change has the potential to stimulate black locust growth on Mediterranean riparian forests. Forest management should therefore include mechanisms to avoid black locust establishment, such as avoid clear-cutting and maintaining closed riparian forest canopies. [ABSTRACT FROM AUTHOR]

Isoprene and monoterpenes are highly reactive organic compounds, emitted by most plant species, which play an important role in air chemistry and air pollution. Different leaf-scale isoprenoid emission models are available. These models are scaled to the canopy through coupling them to terrestrial biogeochemical models and thus used to generate regional emissions inventories. Although the leaf scale models have been shown to perform similarly, large unexplained differences exist in regional emissions inventories. This may be explained in part by the complete lack of inter-comparisons of emission model estimates when scaled from the leaf to the canopy. In this paper we address this problem by coupling four different isoprene emission models (Guenther et al. model, Niinemets et al. model, BIM2 and the Martin et al. model) to two terrestrial biogeochemical model platforms (MoBiLE, GOTILWA+) that describe canopy structure differently. Simulations of isoprene emissions for the Puechabon Mediterranean holm oak stand are analysed, with both canopy photosynthesis models constrained using FLUXNET measurements. The results demonstrate that even with constrained canopy level photosynthesis, large model platform dependent within canopy differences can exist in both modelled photosynthesis and emissions. This results in large differences in modelled isoprenoid emissions, due to the relatively higher sensitivity of emissions to canopy microclimate, in particular temperature. This is the first time emission results from two biogeochemical platforms have been compared, and demonstrates that different canopy descriptions can lead to larger differences in modelled emissions than that attributable to the difference between the emission models themselves. This is an important aspect that has not been acknowledged by the emission modelling community. [Copyright &y& Elsevier]

Abstract: Climate change will increase the frequency and the intensity of droughts in the Mediterranean region, likely reducing growth and increasing mortality of holm oaks (Quercus ilex), one of the most abundant species of Mediterranean forests. In water-limited systems such as those of the Mediterranean, carbon allocation patterns strongly favour belowground accumulation, especially in large subterranean structures called lignotubers. The resilience of these forests depends largely on the replenishment rate of these carbon reserves after disturbances. An experimental thinning, with two intensities (removal of 40% and 80% of basal area), was performed in 1992 in a holm oak forest at the Prades Experimental Complex of Catchments (NE Spain). In 2002, a second thinning was carried out in subplots within the former experimental 0.5 ha plots. Samples from the lignotubers of holm oak trees were analyzed for starch, and both mobile and immobile chemical components, in order to assess the resilience of holm oaks to repeated disturbances. Our results show that after 10 years, starch stocks in the lignotubers have only recovered to half their former values. Removing 40% of the basal area instead of 80% is suggested to be the better managing option for this kind of forests. [Copyright &y& Elsevier]

Recent studies projecting future climate change impacts on forests mainly consider either the effects of climate change on productivity or on disturbances. However, productivity and disturbances are intrinsically linked because 1) disturbances directly affect forest productivity (e.g. via a reduction in leaf area, growing stock or resource-use efficiency), and 2) disturbance susceptibility is often coupled to a certain development phase of the forest with productivity determining the time a forest is in this specific phase of susceptibility. The objective of this paper is to provide an overview of forest productivity changes in different forest regions in Europe under climate change, and partition these changes into effects induced by climate change alone and by climate change and disturbances. We present projections of climate change impacts on forest productivity from state-of-the-art forest models that dynamically simulate forest productivity and the effects of the main European disturbance agents (fire, storm, insects), driven by the same climate scenario in seven forest case studies along a large climatic gradient throughout Europe. Our study shows that, in most cases, including disturbances in the simulations exaggerate ongoing productivity declines or cancel out productivity gains in response to climate change. In fewer cases, disturbances also increase productivity or buffer climate-change induced productivity losses, e.g. because low severity fires can alleviate resource competition and increase fertilization. Even though our results cannot simply be extrapolated to other types of forests and disturbances, we argue that it is necessary to interpret climate change-induced productivity and disturbance changes jointly to capture the full range of climate change impacts on forests and to plan adaptation measures.