Diagnosing destabilization risk in global land carbon sinks.

Global net land carbon uptake or net biome production (NBP) has increased during recent decades1. Whether its temporal variability and autocorrelation have changed during this period, however, remains elusive, even though an increase in both could indicate an increased potential for a destabilized carbon sink2,3. Here, we investigate the trends and controls of net terrestrial carbon uptake and its temporal variability and autocorrelation from 1981 to 2018 using two atmospheric-inversion models, the amplitude of the seasonal cycle of atmospheric CO2 concentration derived from nine monitoring stations distributed across the Pacific Ocean and dynamic global vegetation models. We find that annual NBP and its interdecadal variability increased globally whereas temporal autocorrelation decreased. We observe a separation of regions characterized by increasingly variable NBP, associated with warm regions and increasingly variable temperatures, lower and weaker positive trends in NBP and regions where NBP became stronger and less variable. Plant species richness presented a concave-down parabolic spatial relationship with NBP and its variability at the global scale whereas nitrogen deposition generally increased NBP. Increasing temperature and its increasing variability appear as the most important drivers of declining and increasingly variable NBP. Our results show increasing variability of NBP regionally that can be mostly attributed to climate change and that may point to destabilization of the coupled carbon-climate system.

The three major axes of terrestrial ecosystem function.

The leaf economics spectrum1,2 and the global spectrum of plant forms and functions3 revealed fundamental axes of variation in plant traits, which represent different ecological strategies that are shaped by the evolutionary development of plant species2. Ecosystem functions depend on environmental conditions and the traits of species that comprise the ecological communities4. However, the axes of variation of ecosystem functions are largely unknown, which limits our understanding of how ecosystems respond as a whole to anthropogenic drivers, climate and environmental variability4,5. Here we derive a set of ecosystem functions6 from a dataset of surface gas exchange measurements across major terrestrial biomes. We find that most of the variability within ecosystem functions (71.8%) is captured by three key axes. The first axis reflects maximum ecosystem productivity and is mostly explained by vegetation structure. The second axis reflects ecosystem water-use strategies and is jointly explained by variation in vegetation height and climate. The third axis, which represents ecosystem carbon-use efficiency, features a gradient related to aridity, and is explained primarily by variation in vegetation structure. We show that two state-of-the-art land surface models reproduce the first and most important axis of ecosystem functions. However, the models tend to simulate more strongly correlated functions than those observed, which limits their ability to accurately predict the full range of responses to environmental changes in carbon, water and energy cycling in terrestrial ecosystems7,8.

The essential role of biodiversity in the key axes of ecosystem function.

Biodiversity is essential for maintaining the terrestrial ecosystem multifunctionality (EMF). Recent studies have revealed that the variations in terrestrial ecosystem functions are captured by three key axes: the maximum productivity, water use efficiency, and carbon use efficiency of the ecosystem. However, the role of biodiversity in supporting these three key axes has not yet been explored. In this study, we combined the (i) data collected from more than 840 vegetation plots across a large climatic gradient in China using standard protocols, (ii) data on plant traits and phylogenetic information for more than 2,500 plant species, and (iii) soil nutrient data measured in each plot. These data were used to systematically assess the contribution of environmental factors, species richness, functional and phylogenetic diversity, and community-weighted mean (CWM) and ecosystem traits (i.e., traits intensity normalized per unit land area) to EMF via hierarchical partitioning and Bayesian structural equation modeling. Multiple biodiversity attributes accounted for 70% of the influence of all the variables on EMF, and ecosystems with high functional diversity had high resource use efficiency. Our study is the first to systematically explore the role of different biodiversity attributes, including species richness, phylogenetic and functional diversity, and CWM and ecosystem traits, in the key axes of ecosystem functions. Our findings underscore that biodiversity conservation is critical for sustaining EMF and ultimately ensuring human well-being.

From atoms to ecosystems: elementome diversity meets ecosystem functioning.

The elemental composition of plants (the elementome) is a reliable indicator of their functional traits and the ecological strategies that they follow, and thus represents a good predictor of how ecosystems work. Biodiversity and, especially, functional diversity are also widely recognized as important drivers of ecosystem functioning, mainly because of niche partitioning amongst different species. Here, I review evidence indicating that plant elementomes relate to their ecological niches and how plant elemental concentrations may shift in response to abiotic and biotic drivers. I propose the use of ecosystem elementome diversity as a universal metric to compare ecosystems and investigate diversity-ecosystem functioning relationships. Future research using this promising novel approach will bring together elementomes, diversity, and ecosystem functioning.

Global maps and factors driving forest foliar elemental composition: the importance of evolutionary history.

Consistent information on the current elemental composition of vegetation at global scale and the variables that determine it is lacking. To fill this gap, we gathered a total of 30 912 georeferenced records on woody plants foliar concentrations of nitrogen (N), phosphorus (P) and potassium (K) from published databases, and produced global maps of foliar N, P and K concentrations for woody plants using neural networks at a resolution of 1 km2 . We used data for climate, atmospheric deposition, soil and morphoclimatic groups to train the neural networks. Foliar N, P and K do not follow clear global latitudinal patterns but are consistent with the hypothesis of soil substrate age. We additionally built generalized linear mixed models to investigate the evolutionary history effect together with the effects of environmental effects. In this comparison, evolutionary history effects explained most of the variability in all cases (mostly > 60%). These results emphasize the determinant role of evolutionary history in foliar elemental composition, which should be incorporated in upcoming dynamic global vegetation models.

MASTREE+: Time-series of plant reproductive effort from six continents.

Significant gaps remain in understanding the response of plant reproduction to environmental change. This is partly because measuring reproduction in long-lived plants requires direct observation over many years and such datasets have rarely been made publicly available. Here we introduce MASTREE+, a data set that collates reproductive time-series data from across the globe and makes these data freely available to the community. MASTREE+ includes 73,828 georeferenced observations of annual reproduction (e.g. seed and fruit counts) in perennial plant populations worldwide. These observations consist of 5971 population-level time-series from 974 species in 66 countries. The mean and median time-series length is 12.4 and 10 years respectively, and the data set includes 1122 series that extend over at least two decades (≥20 years of observations). For a subset of well-studied species, MASTREE+ includes extensive replication of time-series across geographical and climatic gradients. Here we describe the open-access data set, available as a.csv file, and we introduce an associated web-based app for data exploration. MASTREE+ will provide the basis for improved understanding of the response of long-lived plant reproduction to environmental change. Additionally, MASTREE+ will enable investigation of the ecology and evolution of reproductive strategies in perennial plants, and the role of plant reproduction as a driver of ecosystem dynamics.

Aún existen importantes vacíos en la comprensión de la respuesta reproductiva de las plantas al cambio medioambiental, en parte, porque su monitoreo en especies de plantas longevas requiere una observación directa durante muchos años, y estos conjuntos de datos rara vez han estado disponibles. Aquí presentamos a MASTREE +, una base de datos que recopila series de tiempo de la reproducción de las plantas de todo el planeta, poniendo a disposición estos datos de libre acceso para la comunidad científica. MASTREE + incluye 73.828 puntos de observación de la reproducción anual georreferenciados (ej. conteos de semillas y frutos) en poblaciones de plantas perennes en todo el mundo. Estas observaciones consisten en 5971 series temporales a nivel de población provenientes de 974 especies en 66 países. La mediana de la duración de las series de tiempo es de 10 años (media = 12.4 años) y el conjunto de datos incluye 1.122 series de al menos dos décadas (≥20 años de observaciones). Para un subconjunto de especies bien estudiadas, MASTREE +incluye un amplio conjunto de series temporales replicadas en gradientes geográficos y climáticos. Describimos el conjunto de datos de acceso abierto disponible como un archivo.csv y presentamos una aplicación web asociada para la exploración de datos. MASTREE+ proporcionará la base para mejorar la comprensión sobre la respuesta reproductiva de plantas longevas al cambio medioambiental. Además, MASTREE+ facilitará los avances en la investigación de la ecología y la evolución de las estrategias reproductivas en plantas perennes y el papel de la reproducción vegetal como determinante de la dinámica de ecosistemas.

Bryophyte C:N:P stoichiometry, biogeochemical niches and elementome plasticity driven by environment and coexistence.

Ecological stoichiometry and studies of biogeochemical niches have mainly focused on plankton and vascular plants, but the phenotypically closest modern relatives of early plants, bryophytes, have been largely neglected. We analysed C:N:P stoichiometries and elemental compositions (K, Na, Mg, Ca, S, Fe) of 35 widely distributed bryophyte species inhabiting springs. We estimated bryophyte C:N:P ratios and their biogeochemical niches, investigated how elementomes respond to the environment and determined whether they tend to diverge more for coexisting than non-coexisting individuals and species. The median C:N:P was 145:8:1, intermediate between Redfield's ratio for marine plankton and those for vascular plants. Biogeochemical niches were differentiated amongst species and were phylogenetically conserved. Differences in individual and species-specific elementomes increased with coexistence between species. Our results provide an evolutionary bridge between the ecological stoichiometries of algae and vascular plants and suggest that differences in elementomes could be used to understand community assemblages and functional diversity.

Increasing climatic sensitivity of global grassland vegetation biomass and species diversity correlates with water availability.

Grasslands are key repositories of biodiversity and carbon storage and are heavily impacted by effects of global warming and changes in precipitation regimes. Patterns of grassland dynamics associated with variability in future climate conditions across spatiotemporal scales are yet to be adequately quantified. Here, we performed a global meta-analysis of year and growing season sensitivities of vegetation aboveground biomass (AGB), aboveground net primary productivity (ANPP), and species richness (SR) and diversity (Shannon index, H) to experimental climate warming and precipitation shifts. All four variables were sensitive to climate change. Their sensitivities to shifts in precipitation were correlated with local background water availability, such as mean annual precipitation (MAP) and aridity, and AGB and ANPP sensitivities were greater in dry habitats than in nonwater-limited habitats. There was no effect of duration of experiment (short vs long term) on sensitivities. Temporal trends in ANPP and SR sensitivity depended on local water availability; ANPP sensitivity to warming increased over time and SR sensitivity to irrigation decreased over time. Our results provide a global overview of the sensitivities of grassland function and diversity to climate change that will improve the understanding of ecological responses across spatiotemporal scales and inform policies for conservation in dry climates.

The effect of global change on soil phosphatase activity.

Soil phosphatase enzymes are produced by plant roots and microorganisms and play a key role in the cycling of phosphorus (P), an often-limiting element in terrestrial ecosystems. The production of these enzymes in soil is the most important biological strategy for acquiring phosphate ions from organic molecules. Previous works showed how soil potential phosphatase activity is mainly driven by climatic conditions and soil nitrogen (N) and carbon. Nonetheless, future trends of the activity of these enzymes under global change remain little known. We investigated the influence of some of the main drivers of change on soil phosphatase activity using a meta-analysis of results from 97 published studies. Our database included a compilation of N and P fertilization experiments, manipulation experiments with increased atmospheric CO2 concentration, warming, and drought, and studies comparing invaded and non-invaded ecosystems. Our results indicate that N fertilization leads to higher phosphatase activity, whereas P fertilization has the opposite effect. The rise of atmospheric CO2 levels or the arrival of invasive species also exhibits positive response ratios on the activity of soil phosphatases. However, the occurrence of recurrent drought episodes decreases the activity of soil phosphatases. Our analysis did not reveal statistically significant effects of warming on soil phosphatase activity. In general, soil enzymatic changes in the reviewed experiments depended on the initial nutrient and water status of the ecosystems. The observed patterns evidence that future soil phosphatase activity will not only depend on present-day soil conditions but also on potential compensations or amplifications among the different drivers of global change. The responses of soil phosphatases to the global change drivers reported in this study and the consideration of cost-benefit approaches based on the connection of the P and N cycle will be useful for a better estimation of phosphatase production in carbon (C)-N-P models.

Is forest fecundity resistant to drought? Results from an 18-yr rainfall-reduction experiment.

Recruitment is a primary determinant of the long-term dynamics of plant populations in changing environments. However, little information is known about the effects of anthropogenic environmental changes on reproductive ecology of trees. We evaluated the impact of experimentally induced 18 yr of drought on reproduction of three contrasting forest trees: Quercus ilex, Phillyrea latifolia and Arbutus unedo. Rainfall reduction did not decrease tree fecundity. Drought, however, affected the allocation of resources in Q. ilex and A. unedo but not the more drought tolerant P. latifolia. Larger crop production by Q. ilex and A. unedo was associated with a stronger decrease in growth in the rainfall-reduction plots compared with the control plots, suggesting that these species were able to maintain their fecundity by shifting their allocation of resources away from growth. Our results indicated resistance to change in tree fecundity in Mediterranean-type forest subjected to an average 15% decrease in the amount of soil moisture, suggesting that these ecosystems may adapt to a progressive increase in arid conditions. However, the species-specific reductions in growth may indirectly affect future fecundity and ultimately shift community composition, even without immediate direct effects of drought on tree fecundity.

The role of climate, foliar stoichiometry and plant diversity on ecosystem carbon balance.

Global change is affecting terrestrial carbon (C) balances. The effect of climate on ecosystem C balance has been largely explored, but the roles of other concurrently changing factors, such as diversity and nutrient availability, remain elusive. We used eddy-covariance C-flux measurements from 62 ecosystems from which we compiled information on climate, ecosystem type, stand age, species abundance and foliar concentrations of N and P of the main species, to assess their importance in the ecosystem C balance. Climate and productivity were the main determinants of ecosystem C balance and its stability. In P-rich sites, increasing N was related to increased gross primary production and respiration and vice versa, but reduced net C uptake. Our analyses did not provide evidence for a strong relation between ecosystem diversity and their productivity and stability. Nonetheless, these results suggest that nutrient imbalances and, potentially, diversity loss may alter future global C balance.

Investigating the relationship between climate, stand age, and temporal trends in masting behavior of European forest trees.

Masting-temporally variable seed production with high spatial synchrony-is a pervasive strategy in wind-pollinated trees that is hypothesized to be vulnerable to climate change due to its correlation with variability in abiotic conditions. Recent work suggests that aging may also have strong effects on seed production patterns of trees, but this potential confounding factor has not been considered in previous times series analysis of climate change effects. Using a 54 year dataset for seven dominant species in 17 forests across Poland, we used the proportion of seed-producing trees (PST) to contrast the predictions of the climate change and aging hypotheses in Abies alba, Fagus sylvatica, Larix decidua, Picea abies, Pinus sylvestris, Quercus petraea, and Quercus robur. Our results show that in all species, PST increased over time and that this change correlated most strongly with stand age, while the standardized precipitation-evapotranspiration index, a measure of drought, contributed to temporal trends in PST of F. sylvatica and Q. robur. Temporal variability of PST also increased over time in all species except P. sylvestris, while trends in temporal autocorrelation and among-stand synchrony reflect species-specific masting strategies. Our results suggest a pivotal role of plant ontogeny in driving not only the extent but also variability and synchrony of reproduction in temperate forest trees. In a time of increasing forest regrowth in Europe, we therefore call for increased attention to demographic effects such as aging on plant reproductive behavior, particularly in studies examining global change effects using long-term time series data.

Towards comparable assessment of the soil nutrient status across scales-Review and development of nutrient metrics.

Nutrient availability influences virtually every aspect of an ecosystem, and is a critical modifier of ecosystem responses to global change. Although this crucial role of nutrient availability in regulating ecosystem structure and functioning has been widely acknowledged, nutrients are still often neglected in observational and experimental synthesis studies due to difficulties in comparing the nutrient status across sites. In the current study, we explain different nutrient-related concepts and discuss the potential of soil-, plant- and remote sensing-based metrics to compare the nutrient status across space. Based on our review and additional analyses on a dataset of European, managed temperate and boreal forests (ICP [International Co-operative Programme on Assessment and Monitoring of Air Pollution Effects on Forests] Forests dataset), we conclude that the use of plant- and remote sensing-based metrics that rely on tissue stoichiometry is limited due to their strong dependence on species identity. The potential use of other plant-based metrics such as Ellenberg indicator values and plant-functional traits is also discussed. We conclude from our analyses and review that soil-based metrics have the highest potential for successful intersite comparison of the nutrient status. As an example, we used and adjusted a soil-based metric, previously developed for conifer forests across Sweden, against the same ICP Forests data. We suggest that this adjusted and further adaptable metric, which included the organic carbon concentration in the upper 20 cm of the soil (including the organic fermentation-humus [FH] layer), the C:N ratio and pHCaCl2 of the FH layer, can be used as a complementary tool along with other indicators of nutrient availability, to compare the background nutrient status across temperate and boreal forests dominated by spruce, pine or beech. Future collection and provision of harmonized soil data from observational and experimental sites is crucial for further testing and adjusting the metric.

Environmental Veto Synchronizes Mast Seeding in Four Contrasting Tree Species.

Synchronized and variable reproduction by perennial plants, called mast seeding, is a major reproductive strategy of trees. The need to accumulate sufficient resources after depletion following fruiting (resource budget), the efficiency of mass flowering for outcross pollination (pollen coupling), or the external factors preventing reproduction (environmental veto) could all synchronize masting. We used seed production data for four species (Quercus ilex, Quercus humilis, Sorbus aucuparia, and Pinus albicaulis) to parametrize resource budget models of masting. Based on species life-history characteristics, we hypothesized that pollen coupling should synchronize reproduction in S. aucuparia and P. albicaulis, while in Q. ilex and Q. humilis, environmental veto should be a major factor. Pollen coupling was stronger in S. aucuparia and P. albicaulis than in oaks, while veto was more frequent in the latter. Yet in all species, costs of reproduction were too small to impose a replenishment period. A synchronous environmental veto, in the presence of environmental stochasticity, was sufficient to produce observed variability and synchrony in reproduction. In the past, vetoes like frost events that prevent reproduction have been perceived as negative for plants. In fact, they could be selectively favored as a way to create mast seeding.

Nutrient scarcity strengthens soil fauna control over leaf litter decomposition in tropical rainforests.

Soil fauna is a key control of the decomposition rate of leaf litter, yet its interactions with litter quality and the soil environment remain elusive. We conducted a litter decomposition experiment across different topographic levels within the landscape replicated in two rainforest sites providing natural gradients in soil fertility to test the hypothesis that low nutrient availability in litter and soil increases the strength of fauna control over litter decomposition. We crossed these data with a large dataset of 44 variables characterizing the biotic and abiotic microenvironment of each sampling point and found that microbe-driven carbon (C) and nitrogen (N) losses from leaf litter were 10.1 and 17.9% lower, respectively, in the nutrient-poorest site, but this among-site difference was equalized when meso- and macrofauna had access to the litterbags. Further, on average, soil fauna enhanced the rate of litter decomposition by 22.6%, and this contribution consistently increased as nutrient availability in the microenvironment declined. Our results indicate that nutrient scarcity increases the importance of soil fauna on C and N cycling in tropical rainforests. Further, soil fauna is able to equalize differences in microbial decomposition potential, thus buffering to a remarkable extent nutrient shortages at an ecosystem level.

Nutrient-rich plants emit a less intense blend of volatile isoprenoids.

The emission of isoprenoids (e.g. isoprene and monoterpenes) by plants plays an important defensive role against biotic and abiotic stresses. Little is known, however, about the functional traits linked to species-specific variability in the types and rates of isoprenoids emitted and about possible co-evolution of functional traits with isoprenoid emission type (isoprene emitter, monoterpene emitter or both). We combined data for isoprene and monoterpene emission rates per unit dry mass with key functional traits (foliar nitrogen (N) and phosphorus (P) concentrations, and leaf mass per area) and climate for 113 plant species, covering the boreal, wet temperate, Mediterranean and tropical biomes. Foliar N was positively correlated with isoprene emission, and foliar P was negatively correlated with both isoprene and monoterpene emission rate. Nonemitting plants generally had the highest nutrient concentrations, and those storing monoterpenes had the lowest concentrations. Our phylogenetic analyses found that the type of isoprenoid emission followed an adaptive, rather than a random model of evolution. Evolution of isoprenoids may be linked to nutrient availability. Foliar N and P are good predictors of the type of isoprenoid emission and the rate at which monoterpenes, and to a lesser extent isoprene, are emitted.

The Moran effect and environmental vetoes: phenological synchrony and drought drive seed production in a Mediterranean oak.

Masting is the highly variable production of synchronized seed crops, and is a common reproductive strategy in plants. Weather has long been recognized as centrally involved in driving seed production in masting plants. However, the theory behind mechanisms connecting weather and seeding variation has only recently been developed, and still lacks empirical evaluation. We used 12-year long seed production data for 255 holm oaks (Quercus ilex), as well as airborne pollen and meteorological data, and tested whether masting is driven by environmental constraints: phenological synchrony and associated pollination efficiency, and drought-related acorn abscission. We found that warm springs resulted in short pollen seasons, and length of the pollen seasons was negatively related to acorn production, supporting the phenological synchrony hypothesis. Furthermore, the relationship between phenological synchrony and acorn production was modulated by spring drought, and effects of environmental vetoes on seed production were dependent on last year's environmental constraint, implying passive resource storage. Both vetoes affected among-tree synchrony in seed production. Finally, precipitation preceding acorn maturation was positively related to seed production, mitigating apparent resource depletion following high crop production in the previous year. These results provide new insights into mechanisms beyond widely reported weather and seed production correlations.

The role of nutrients, productivity and climate in determining tree fruit production in European forests.

Fruit production (NPPf ), the amount of photosynthates allocated to reproduction (%GPPf ) and their controls for spatial and species-specific variability (e.g. nutrient availability, climate) have been poorly studied in forest ecosystems. We characterized fruit production and its temporal behaviour for several tree species and resolved the effects of gross primary production (GPP), climate and foliar nutrient concentrations. We used data for litterfall and foliar nutrient concentration from 126 European forests and related them to climatic data. GPP was estimated for each forest using a regression model. Mean NPPf ranged from c. 10 to 40 g C m-2  yr-1 and accounted for 0.5-3% of GPP. Forests with higher GPPs produced larger fruit crops. Foliar zinc (Zn) and phosphorus (P) concentrations were associated positively with NPPf , whereas foliar Zn and potassium (K) were negatively related to its temporal variability. Maximum NPPf and interannual variability of NPPf were higher in Fagaceae than in Pinaceae species. NPPf and %GPPf were similar amongst the studied species despite the different reproductive temporal behaviour of Fagaceae and Pinaceae species. We report that foliar concentrations of P and Zn are associated with %GPPf , NPPf and its temporal behaviour.

Fungal composition associated with host tree identity mediates nutrient addition effects on wood microbial respiration.

Fungi are key decomposers of deadwood, but the impact of anthropogenic changes in nutrients and temperature on fungal community and its consequences for wood microbial respiration are not well understood. Here, we examined how nitrogen and phosphorus additions (field experiment) and warming (laboratory experiment) together influence fungal composition and microbial respiration from decomposing wood of angiosperms and gymnosperms in a subtropical forest. Nutrient additions significantly increased wood microbial respiration via fungal composition, but effects varied with nutrient types and taxonomic groups. Specifically, phosphorus addition significantly increased wood microbial respiration (65%) through decreased acid phosphatase activity and increased abundance of fast-decaying fungi (e.g., white rot), while nitrogen addition marginally increased it (30%). Phosphorus addition caused a greater increase in microbial respiration in gymnosperms than in angiosperms (83.3% vs. 46.9%), which was associated with an increase in Basidiomycota:Ascomycota operational taxonomic unit abundance in gymnosperms but a decrease in angiosperms. The temperature dependencies of microbial respiration were remarkably constant across nutrient levels, consistent with metabolic scaling theory hypotheses. This is because there was no significant interaction between temperature and wood phosphorus availability or fungal composition, or the interaction among the three factors. Our results highlight the key role of tree identity in regulating nutrient response of wood microbial respiration through controlling fungal composition. Given that the range of angiosperm species may expand under climate warming and forest management, our data suggest that expansion will decrease nutrient effects on forest carbon cycling in forests previously dominated by gymnosperm species.

Chlorophyll fluorescence variation in two Mediterranean forest species over a 21-year drought treatment period.

The Mediterranean is among the regions predicted to be most affected by climate change due to rising temperatures and increasing frequency as well as intensity of extreme weather events, such as drought. These shifts in climatic conditions might lead to changes in species community composition by enabling the increase of drought-tolerant species at the expense of less tolerant ones. This hypothesis was tested in the current study using chlorophyll fluorescence data from a 21-year precipitation exclusion experiment in a Mediterranean forest for two co-dominant Quercus ilex L. and Phillyrea latifolia L. species with contrasting low and high levels of drought tolerance, respectively. The maximum potential quantum efficiency of photosystem II (PSII) (Fv/Fm), photochemical efficiency of PSII (yield) and non-photochemical quenching (NPQ) varied seasonally. The Fv/Fm and NPQ levels were positively correlated with air temperature and standardized precipitation-evapotranspiration index (SPEI), whereas yield, which was greater under drought treatment, was negatively associated with vapor pressure deficit and SPEI. The Fv/Fm values showed similar increase in the two species over the 21-year study period regardless of treatment and in parallel to progressive warming. By contrast, yield values were higher in Q. ilex than in P. latifolia, while NPQ values were higher in P. latifolia than in Q. ilex. Notably, high yield values were also observed in the drought-treated plots. Throughout the study, plants in the drought-treated plots exhibited decreased basal area, leaf biomass and aerial cover due to high stem mortality. In addition, a continuous increase in temperature was detected in summer and autumn, which might explain the observed increase in Fv/Fm values over the study period. Higher yield and lower NPQ detected in Q. ilex could be attributed to less competition for resources in the drought-treated plots and acclimation of Q. ilex plants over the study period. Our results indicate that reduction in stem density could improve forest resilience to climate change-induced drought conditions.