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Microclimate research gained renewed interest over the last decade and its importance for many ecological processes is increasingly being recognized. Consequently, the call for high-resolution microclimatic temperature grids across broad spatial extents is becoming more pressing to improve ecological models. Here, we provide a new set of open-access bioclimatic variables for microclimate temperatures of European forests at 25 × 25 m2 resolution.
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Microclima , Árboles , Temperatura , Bosques , EcosistemaRESUMEN
The stability of ecological communities is critical for the stable provisioning of ecosystem services, such as food and forage production, carbon sequestration, and soil fertility. Greater biodiversity is expected to enhance stability across years by decreasing synchrony among species, but the drivers of stability in nature remain poorly resolved. Our analysis of time series from 79 datasets across the world showed that stability was associated more strongly with the degree of synchrony among dominant species than with species richness. The relatively weak influence of species richness is consistent with theory predicting that the effect of richness on stability weakens when synchrony is higher than expected under random fluctuations, which was the case in most communities. Land management, nutrient addition, and climate change treatments had relatively weak and varying effects on stability, modifying how species richness, synchrony, and stability interact. Our results demonstrate the prevalence of biotic drivers on ecosystem stability, with the potential for environmental drivers to alter the intricate relationship among richness, synchrony, and stability.
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Plantas/clasificación , Secuestro de Carbono , Cambio Climático , Ecosistema , Desarrollo de la Planta , Plantas/metabolismo , Suelo/químicaRESUMEN
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.
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Pradera , Agua , Biomasa , Cambio Climático , Ecosistema , LluviaRESUMEN
Ecological research heavily relies on coarse-gridded climate data based on standardized temperature measurements recorded at 2 m height in open landscapes. However, many organisms experience environmental conditions that differ substantially from those captured by these macroclimatic (i.e. free air) temperature grids. In forests, the tree canopy functions as a thermal insulator and buffers sub-canopy microclimatic conditions, thereby affecting biological and ecological processes. To improve the assessment of climatic conditions and climate-change-related impacts on forest-floor biodiversity and functioning, high-resolution temperature grids reflecting forest microclimates are thus urgently needed. Combining more than 1200 time series of in situ near-surface forest temperature with topographical, biological and macroclimatic variables in a machine learning model, we predicted the mean monthly offset between sub-canopy temperature at 15 cm above the surface and free-air temperature over the period 2000-2020 at a spatial resolution of 25 m across Europe. This offset was used to evaluate the difference between microclimate and macroclimate across space and seasons and finally enabled us to calculate mean annual and monthly temperatures for European forest understories. We found that sub-canopy air temperatures differ substantially from free-air temperatures, being on average 2.1°C (standard deviation ± 1.6°C) lower in summer and 2.0°C higher (±0.7°C) in winter across Europe. Additionally, our high-resolution maps expose considerable microclimatic variation within landscapes, not captured by the gridded macroclimatic products. The provided forest sub-canopy temperature maps will enable future research to model below-canopy biological processes and patterns, as well as species distributions more accurately.
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Bosques , Microclima , Cambio Climático , Temperatura , ÁrbolesRESUMEN
Aim: The aim was to evaluate the effects of climate warming on biodiversity across spatial scales (i.e., α-, ß- and γ-diversity) and the effects of patch openness and experimental context on diversity responses. Location: Global. Time period: 1995-2017. Major taxa studied: Fungi, invertebrates, phytoplankton, plants, seaweed, soil microbes and zooplankton. Methods: We compiled data from warming experiments and conducted a meta-analysis to evaluate the effects of warming on different components of diversity (such as species richness and equivalent numbers) at different spatial scales (α-, ß- and γ-diversity, partitioning ß-diversity into species turnover and nestedness components). We also investigated how these effects were modulated by system openness, defined as the possibility of replicates being colonized by new species, and experimental context (duration, mean temperature change and ecosystem type). Results: Experimental warming did not affect local species richness (α-diversity) but decreased effective numbers of species by affecting species dominance. Warming increased species spatial turnover (ß-diversity), although no significant changes were detected at the regional scale (γ-diversity). Site openness and experimental context did not significantly affect our results, despite significant heterogeneity in the effect sizes of α- and ß-diversity. Main conclusions: Our meta-analysis shows that the effects of warming on biodiversity are scale dependent. The local and regional inventory diversity remain unaltered, whereas species composition across temperature gradients and the patterns of species dominance change with temperature, creating novel communities that might be harder to predict.
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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.
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Sequías , Quercus , Ecosistema , Fertilidad , Bosques , ÁrbolesRESUMEN
Soil enzymes are central in the response of terrestrial ecosystems to climate change, and their study can be crucial for the models' implementation. We investigated for 1 year the effects of warming and seasonality on the potential activities of five soil extracellular enzymes and their relationships with soil moisture, phosphorus (P) concentration, and other soil parameters in a P-limited Mediterranean semiarid shrubland. The site was continuously subjected to warming since 1999, and we compared data from this study to analogous data from 2004. Warming uniformly increased all enzymes activities, but only when a sufficient amount of soil water was available. Seasonality unevenly altered enzyme activities, thus affecting enzymatic stoichiometry. P deficiency affected enzymatic stoichiometry, favoring the activities of the phosphatases. The effect of warming was stronger in 2014 than 2004, excluding the hypothesis of acclimation of rhizospheric responses to higher temperatures and suggesting that further increases in extracellular enzymatic activities are to be expected if sufficient water is available. Climatic warming will likely generally stimulate soil enzymatic activities and accelerate nutrient mineralization and similar ecological processes such as the production and degradation of biomass and changes in community composition, but which will be limited by water availability, especially in Mediterranean soils in summer. Winters in such ecosystems will benefit from a general increase in activity and production, but biological activity could even decrease in summer, potentially leading to a negative overall balance of nutrient mineralization. This study suggests that a general increase in activity due to warming could lead to faster mineralization of soil organic matter and water consumption in colder climates, until one of these factors in turn becomes limiting. Such trade-offs between water and temperature in relation with enzyme activity should be considered in biogeochemical models.
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Ecosistema , Suelo , Cambio Climático , Estaciones del Año , Microbiología del SueloRESUMEN
Changes in rainfall amounts and patterns have been observed and are expected to continue in the near future with potentially significant ecological and societal consequences. Modelling vegetation responses to changes in rainfall is thus crucial to project water and carbon cycles in the future. In this study, we present the results of a new model-data intercomparison project, where we tested the ability of 10 terrestrial biosphere models to reproduce the observed sensitivity of ecosystem productivity to rainfall changes at 10 sites across the globe, in nine of which, rainfall exclusion and/or irrigation experiments had been performed. The key results are as follows: (a) Inter-model variation is generally large and model agreement varies with timescales. In severely water-limited sites, models only agree on the interannual variability of evapotranspiration and to a smaller extent on gross primary productivity. In more mesic sites, model agreement for both water and carbon fluxes is typically higher on fine (daily-monthly) timescales and reduces on longer (seasonal-annual) scales. (b) Models on average overestimate the relationship between ecosystem productivity and mean rainfall amounts across sites (in space) and have a low capacity in reproducing the temporal (interannual) sensitivity of vegetation productivity to annual rainfall at a given site, even though observation uncertainty is comparable to inter-model variability. (c) Most models reproduced the sign of the observed patterns in productivity changes in rainfall manipulation experiments but had a low capacity in reproducing the observed magnitude of productivity changes. Models better reproduced the observed productivity responses due to rainfall exclusion than addition. (d) All models attribute ecosystem productivity changes to the intensity of vegetation stress and peak leaf area, whereas the impact of the change in growing season length is negligible. The relative contribution of the peak leaf area and vegetation stress intensity was highly variable among models.
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Ciclo del Carbono , Ecosistema , Hojas de la Planta , Estaciones del Año , AguaRESUMEN
Productivity of tropical lowland moist forests is often limited by availability and functional allocation of phosphorus (P) that drives competition among tree species and becomes a key factor in determining forestall community diversity. We used non-target 31P-NMR metabolic profiling to study the foliar P-metabolism of trees of a French Guiana rainforest. The objective was to test the hypotheses that P-use is species-specific, and that species diversity relates to species P-use and concentrations of P-containing compounds, including inorganic phosphates, orthophosphate monoesters and diesters, phosphonates and organic polyphosphates. We found that tree species explained the 59% of variance in 31P-NMR metabolite profiling of leaves. A principal component analysis showed that tree species were separated along PC 1 and PC 2 of detected P-containing compounds, which represented a continuum going from high concentrations of metabolites related to non-active P and P-storage, low total P concentrations and high N:P ratios, to high concentrations of P-containing metabolites related to energy and anabolic metabolism, high total P concentrations and low N:P ratios. These results highlight the species-specific use of P and the existence of species-specific P-use niches that are driven by the distinct species-specific position in a continuum in the P-allocation from P-storage compounds to P-containing molecules related to energy and anabolic metabolism.
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Metaboloma , Metabolómica , Fósforo/metabolismo , Bosque Lluvioso , Árboles/metabolismo , Guyana Francesa , Hojas de la Planta/metabolismo , Especificidad de la EspecieRESUMEN
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.
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Bosque Lluvioso , Animales , Carbono , Nitrógeno , Hojas de la Planta , Suelo/químicaRESUMEN
Global warming and reduced precipitation may trigger large-scale species losses and vegetation shifts in ecosystems around the world. However, currently lacking are practical ways to quantify the sensitivity of species and community composition to these often-confounded climatic forces. Here we conducted long-term (16 yr) nocturnal-warming (+0.6°C) and reduced precipitation (-20% soil moisture) experiments in a Mediterranean shrubland. Climatic niche groups (CNGs) - species ranked or classified by similar temperature or precipitation distributions - informatively described community responses under experimental manipulations. Under warming, CNGs revealed that only those species distributed in cooler regions decreased. Correspondingly, under reduced precipitation, a U-shaped treatment effect observed in the total community was the result of an abrupt decrease in wet-distributed species, followed by a delayed increase in dry-distributed species. Notably, while partially correlated, CNG explanations of community response were stronger for their respective climate parameter, suggesting some species possess specific adaptations to either warming or drought that may lead to independent selection to the two climatic variables. Our findings indicate that when climatic distributions are combined with experiments, the resulting incorporation of local plant evolutionary strategies and their changing dynamics over time leads to predictable and informative shifts in community structure under independent climate change scenarios.
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Clima , Sequías , Calentamiento Global , Ecosistema , Estaciones del Año , Especificidad de la Especie , TemperaturaRESUMEN
Global warming and recurring drought are expected to accelerate water limitation for plant communities in semiarid Mediterranean ecosystems and produce directional shifts in structure and composition that are not easily detected, and supporting evidence is scarce. We conducted a long-term (17 years) nocturnal-warming (+0.6°C) and drought (-40% rainfall) experiments in an early-successional Mediterranean shrubland to study the changes in community structure and composition, contrasting functional groups and dominant species, and the superimposed effects of natural extreme drought. Species richness decreased in both the warming and drought treatments. Responses to the moderate warming were associated with decreases in herb abundance, and responses to the drought were associated with decreases in both herb and shrub abundances. The drought also significantly decreased community diversity and evenness. Changes in abundance differed between herbs (decreases) and shrubs (increases or no changes). Both warming and drought, especially drought, increased the relative species richness and abundance of shrubs, favoring the establishment of shrubs. Both warming and drought produced significant shifts in plant community composition. Experimental warming shifted the community composition from Erica multiflora toward Rosmarinus officinalis, and drought consistently shifted the composition toward Globularia alypum. The responses in biodiversity (e.g., community biodiversity, changes of functional groups and compositional shifts) were also strongly correlated with atmospheric drought (SPEI) in winter-spring and/or summer, indicating sensitivity to water limitation in this early-successional Mediterranean ecosystem, especially to natural extreme droughts. Our results suggest that the shifts in species assembles and community diversity and composition are accelerated by the long-term nocturnal-warming and drought, combined with natural severe droughts, and that the magnitude of the impacts of climate change is also correlated with the successional status of ecosystem. The results thus highlight the necessity for assessing the impacts on ecosystemic functioning and services and developing effective measures for conserving biodiversity.
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Cambio Climático , Sequías , Ecosistema , Biodiversidad , Ericaceae , Estaciones del AñoRESUMEN
Well-defined productivity-precipitation relationships of ecosystems are needed as benchmarks for the validation of land models used for future projections. The productivity-precipitation relationship may be studied in two ways: the spatial approach relates differences in productivity to those in precipitation among sites along a precipitation gradient (the spatial fit, with a steeper slope); the temporal approach relates interannual productivity changes to variation in precipitation within sites (the temporal fits, with flatter slopes). Precipitation-reduction experiments in natural ecosystems represent a complement to the fits, because they can reduce precipitation below the natural range and are thus well suited to study potential effects of climate drying. Here, we analyse the effects of dry treatments in eleven multiyear precipitation-manipulation experiments, focusing on changes in the temporal fit. We expected that structural changes in the dry treatments would occur in some experiments, thereby reducing the intercept of the temporal fit and displacing the productivity-precipitation relationship downward the spatial fit. The majority of experiments (72%) showed that dry treatments did not alter the temporal fit. This implies that current temporal fits are to be preferred over the spatial fit to benchmark land-model projections of productivity under future climate within the precipitation ranges covered by the experiments. Moreover, in two experiments, the intercept of the temporal fit unexpectedly increased due to mechanisms that reduced either water loss or nutrient loss. The expected decrease of the intercept was observed in only one experiment, and only when distinguishing between the late and the early phases of the experiment. This implies that we currently do not know at which precipitation-reduction level or at which experimental duration structural changes will start to alter ecosystem productivity. Our study highlights the need for experiments with multiple, including more extreme, dry treatments, to identify the precipitation boundaries within which the current temporal fits remain valid.
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Clima , Ecología/métodos , Ecosistema , Lluvia , AguaRESUMEN
Climate change is predicted to increase the aridity in the Mediterranean Basin and severely affect forest productivity and composition. The responses of forests to different timescales of drought, however, are still poorly understood because extreme and persistent moderate droughts can produce nonlinear responses in plants. We conducted a rainfall-manipulation experiment in a Mediterranean forest dominated by Quercus ilex, Phillyrea latifolia, and Arbutus unedo in the Prades Mountains in southern Catalonia from 1999 to 2014. The experimental drought significantly decreased forest aboveground-biomass increment (ABI), tended to increase the litterfall, and decreased aboveground net primary production throughout the 15 years of the study. The responses to the experimental drought were highly species-specific. A. unedo suffered a significant reduction in ABI, Q. ilex experienced a decrease during the early experiment (1999-2003) and in the extreme droughts of 2005-2006 and 2011-2012, and P. latifolia was unaffected by the treatment. The drought treatment significantly increased branch litterfall, especially in the extremely dry year of 2011, and also increased overall leaf litterfall. The drought treatment reduced the fruit production of Q. ilex, which affected seedling recruitment. The ABIs of all species were highly correlated with SPEI in early spring, whereas the branch litterfalls were better correlated with summer SPEIs and the leaf and fruit litterfalls were better correlated with autumn SPEIs. These species-specific responses indicated that the dominant species (Q. ilex) could be partially replaced by the drought-resistant species (P. latifolia). However, the results of this long-term study also suggest that the effect of drought treatment has been dampened over time, probably due to a combination of demographic compensation, morphological and physiological acclimation, and epigenetic changes. However, the structure of community (e.g., species composition, dominance, and stand density) may be reordered when a certain drought threshold is reached.
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Cambio Climático , Sequías , Ericaceae/fisiología , Oleaceae/fisiología , Quercus/fisiología , Aclimatación , Biomasa , Ericaceae/crecimiento & desarrollo , Bosques , Nueces/crecimiento & desarrollo , Nueces/fisiología , Oleaceae/crecimiento & desarrollo , Hojas de la Planta/crecimiento & desarrollo , Hojas de la Planta/fisiología , Brotes de la Planta/crecimiento & desarrollo , Brotes de la Planta/fisiología , Quercus/crecimiento & desarrollo , España , Estrés FisiológicoRESUMEN
Vegetation in water-limited ecosystems relies strongly on access to deep water reserves to withstand dry periods. Most of these ecosystems have shallow soils over deep groundwater reserves. Understanding the functioning and functional plasticity of species-specific root systems and the patterns of or differences in the use of water sources under more frequent or intense droughts is therefore necessary to properly predict the responses of seasonally dry ecosystems to future climate. We used stable isotopes to investigate the seasonal patterns of water uptake by a sclerophyll forest on sloped terrain with shallow soils. We assessed the effect of a long-term experimental drought (12 years) and the added impact of an extreme natural drought that produced widespread tree mortality and crown defoliation. The dominant species, Quercus ilex, Arbutus unedo and Phillyrea latifolia, all have dimorphic root systems enabling them to access different water sources in space and time. The plants extracted water mainly from the soil in the cold and wet seasons but increased their use of groundwater during the summer drought. Interestingly, the plants subjected to the long-term experimental drought shifted water uptake toward deeper (10-35 cm) soil layers during the wet season and reduced groundwater uptake in summer, indicating plasticity in the functional distribution of fine roots that dampened the effect of our experimental drought over the long term. An extreme drought in 2011, however, further reduced the contribution of deep soil layers and groundwater to transpiration, which resulted in greater crown defoliation in the drought-affected plants. This study suggests that extreme droughts aggravate moderate but persistent drier conditions (simulated by our manipulation) and may lead to the depletion of water from groundwater reservoirs and weathered bedrock, threatening the preservation of these Mediterranean ecosystems in their current structures and compositions.
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Sequías , Árboles/fisiología , Agua/metabolismo , Hidrología , Longevidad , Estaciones del Año , España , Especificidad de la Especie , Árboles/crecimiento & desarrolloRESUMEN
At the molecular level, folivory activity on plants has mainly been related to the foliar concentrations of nitrogen (N) and/or particular metabolites. We studied the responses of different nutrients and the whole metabolome of Quercus ilex to seasonal changes and to moderate field experimental conditions of drought, and how this drought may affect folivory activity, using stoichiometric and metabolomic techniques. Foliar potassium (K) concentrations increased in summer and consequently led to higher foliar K : phosphorus (P) and lower carbon (C) : K and N : K ratios. Foliar N : P ratios were not lowest in spring as expected by the growth rate hypothesis. Trees exposed to moderate drought presented higher concentrations of total sugars and phenolics and these trees also experienced more severe folivory attack. The foliar increases in K, sugars and antioxidant concentrations in summer, the driest Mediterranean season, indicated enhanced osmoprotection under natural drought conditions. Trees under moderate drought also presented higher concentrations of sugars and phenolics; a plant response to avoid water loss. These shifts in metabolism produced an indirect relationship between increased drought and folivory activity.
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Sequías , Herbivoria/fisiología , Metaboloma , Hojas de la Planta/metabolismo , Quercus/fisiología , Árboles/fisiología , Aminoácidos/metabolismo , Metabolismo de los Hidratos de Carbono , Análisis Discriminante , Análisis de los Mínimos Cuadrados , Metabolómica , Modelos Biológicos , Polifenoles/metabolismo , Análisis de Componente Principal , Estaciones del Año , EspañaRESUMEN
Understanding the soil biogeochemical responses to increasing global warming in the near future is essential for improving our capacity to mitigate the impacts of climate change on highly vulnerable Mediterranean ecosystems. Previous studies have primarily focused on the effects of warming on various biogeochemical processes. However, there is limited knowledge about how the changes in water availability associated to high temperatures can alter the bioavailability and dynamics of soil elements, thereby impacting ecosystem productivity, species composition, and pollution through soil biogeochemical and hydrological processes. In this study, we investigated the effects of long-term nighttime warming on the extractable concentrations of organic carbon (EOC), total nitrogen (ETN), total phosphorus (ETP), and 17 mineral elements (arsenic (As), calcium (Ca), cadmium (Cd), chromium (Cr), copper (Cu), iron (Fe), mercury (Hg), potassium (K), magnesium (Mg), manganese (Mn), molybdenum (Mo), nickel (Ni), lead (Pb), sulfur (S), strontium (Sr), vanadium (V), and zinc (Zn)) through environmental experiments in a semi-arid Mediterranean shrubland. We explored the potential biotic and abiotic mechanisms underlying the seasonal and long-term changes in extractable-mobilizable elemental composition and concentrations. Our findings revealed that prolonged warming led to higher mean annual soil temperature (with an average increase of 0.67⯰C from 1999 to 2014), accumulation of soil organic matter (EOC) and extractable concentrations of soil elements (particularly increased ETP and extractable Ca, Mg, Cu, Sr, Mn, and As). These changes were attributed to uniformly higher activities of extracellular soil enzymes and/or lower plant photosynthetic and nutrient uptake capacity linked to more water deficit under warmer conditions. Seasonality unevenly altered element extractable concentrations, with soil microclimate (temperature and water content) and biological (soil microbial and plant) activity being the main drivers of this variability, thus influencing soil element composition. These results suggest significant fluctuations in the extractable concentrations of specific mineral elements in these soils, implying potential future variations in soil element composition as well as the loss of total element concentrations/contents in semi-arid Mediterranean ecosystems due to increasing warming. Therefore, these findings enhance our ability to predict ecosystem management strategies and mitigate the observed negative impacts on plant-soil systems and water quality in the context of climate change.
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Forests respond to increasing intensities and frequencies of drought by reducing growth and with higher tree mortality rates. Little is known, however, about the long-term consequences of generally drier conditions and more frequent extreme droughts. A Holm oak forest was exposed to experimental rainfall manipulation for 13 years to study the effect of increasing drought on growth and mortality of the dominant species Quercus ilex, Phillyrea latifolia, and Arbutus unedo. The drought treatment reduced stem growth of A. unedo (-66.5%) and Q. ilex (-17.5%), whereas P. latifolia remained unaffected. Higher stem mortality rates were noticeable in Q. ilex (+42.3%), but not in the other two species. Stem growth was a function of the drought index of early spring in the three species. Stem mortality rates depended on the drought index of winter and spring for Q. ilex and in spring and summer for P. latifolia, but showed no relation to climate in A. unedo. Following a long and intense drought (2005-2006), stem growth of Q. ilex and P. latifolia increased, whereas it decreased in A. unedo. Q. ilex also enhanced its survival after this period. Furthermore, the effect of drought treatment on stem growth in Q. ilex and A. unedo was attenuated as the study progressed. These results highlight the different vulnerabilities of Mediterranean species to more frequent and intense droughts, which may lead to partial species substitution and changes in forest structure and thus in carbon uptake. The response to drought, however, changed over time. Decreased intra- and interspecific competition after extreme events with high mortality, together with probable morphological and physiological acclimation to drought during the study period, may, at least in the short term, buffer forests against drier conditions. The long-term effects of drought consequently deserve more attention, because the ecosystemic responses are unlikely to be stable over time.Nontechnical summaryIn this study, we evaluate the effect of long-term (13 years) experimental drought on growth and mortality rates of three forest Mediterranean species, and their response to the different intensities and durations of natural drought. We provide evidence for species-specific responses to drought, what may eventually lead to a partial community shift favoring the more drought-resistant species. However, we also report a dampening of the treatment effect on the two drought-sensitive species, which may indicate a potential adaptation to drier conditions at the ecosystem or population level. These results are thus relevant to account for the stabilizing processes that would alter the initial response of ecosystem to drought through changes in plant physiology, morphology, and demography compensation.
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Sequías , Ericaceae/crecimiento & desarrollo , Oleaceae/crecimiento & desarrollo , Quercus/crecimiento & desarrollo , Tallos de la Planta/crecimiento & desarrollo , España , ÁrbolesRESUMEN
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.
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Sequías , Quercus , Fluorescencia , Bosques , Hojas de la Planta , Complejo de Proteína del Fotosistema II , ClorofilaRESUMEN
Soil mineral elements play a crucial role in ecosystem productivity and pollution dynamics. Climate models project an increase in drought severity in the Mediterranean Basin in the coming decades, which could lead to changes in the composition and concentrations of mineral elements in soils. These changes can have significant impacts on the fundamental processes of plant-soil cycles. While previous studies have predominantly focused on carbon, nitrogen, and phosphorus, there is a notable lack of research on the biogeochemical responses of other mineral elements to increasing drought. In this study, we investigated the effects of chronic drought (15 years of experimental rainfall exclusion) and seasonal drought (summer period) on the extractable soil concentrations of 17 mineral elements (arsenic (As), calcium (Ca), cadmium (Cd), chromium (Cr), copper (Cu), iron (Fe), mercury (Hg), potassium (K), magnesium (Mg), manganese (Mn), molybdenum (Mo), nickel (Ni), lead (Pb), sulphur (S), strontium (Sr), vanadium (V) and zinc (Zn)) in a Mediterranean holm oak forest. We also explored the potential biotic and abiotic mechanisms underlying the changes in extractable elemental concentrations under chronic drought conditions. Our findings reveal that soil elemental concentrations varied significantly due to seasonal changes and chronic drought, with soil microclimate, biological activity, and organic matter being the main drivers of this variability. Levels of soil water content primarily explained the observed variations in soil elemental concentrations. Most of the mineral elements (13 out of 17) exhibited higher concentrations during winter-spring (wet seasons) compared to summer-autumn (dry seasons). The chronic drought treatment resulted in K limitation, increasing vegetation vulnerability to drought stress. Conversely, the accumulation of S in soils due to drought may intensify the risk of S losses from the plant-soil system. Under drought conditions, certain trace elements (particularly Mn, V, and Cd) exhibited increased extractability, posing potential risks to plant health and the exportation of these elements into continental waters. Overall, our results suggest that alterations in mineral element concentrations under future drier conditions could promote ecosystem degradation and pollution dispersion in the Mediterranean Basin. Understanding and predicting these changes are essential for effective ecosystem management and mitigating the potential negative impacts on plant health and water quality.