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The impact of various environmental stressors, such as plant invasion and extreme drought is increasing in many ecosystems around the globe. Mediterranean ecosystems have especially been affected by both stressors in recent decades. Thus, we conducted a precipitation and shrub invasion manipulation experiment in a Mediterranean cork oak (Quercus suber L.) ecosystem in Portugal. The impact of both stressors (i.e. drought and invasion) as well as their interaction on cork oak functioning was investigated, resulting in four experimental treatments: 1) control trees, 2) trees with a rain exclusion of 45%, 3) trees invaded by the shrub Cistus ladanifer and 4) trees invaded by C. ladanifer combined with rain exclusion. Each of the four treatments was replicated in three spatially separated blocks. In total 36 trees (9 per treatment) and 18 shrubs (9 per treatment) were selected randomly for measurements between October 2017 and March 2020.Invaded trees, independent of the rain exclusion, had lower leaf area index (LAI) and growth rates (trunk increment). Further changes in the relationship of pre-dawn and midday leaf water potential elucidated that competition by shrubs shifted the hydraulic behaviour of invaded trees to a more anisohydric strategy compared to non-invaded cork oaks during summer drought. These negative impacts of shrub invasion had also an effect on transpiration rates of trees, but the extent was dependent on annual precipitation. In the wet year 2018, transpiration rates of trees in all four treatments were similar due to replenished soil water resources. However, in 2019, when precipitation was strongly reduced (ambient: –25%, rain exclusion: –50%), trees under invasion and rain exclusion reduced their transpiration by 47% compared to control trees, which was stronger than the reduction caused by a single stressor (amplifying interaction). However, shrubs under the rain exclusion also suffered from the extreme drought and were not able to recover in the following autumn/winter period, releasing trees from a strong competition (buffering interaction). On the other hand, shrubs under ambient precipitation were highly competitive in the same recovery period and strongly delayed tree transpiration recovery (–51% compared to control). Consequently, extreme drought and shrub invasion interacted buffering in this recovery period after the extreme drought event. In conclusion, the lessons learned from this manipulation experiment are 1) that shrub invasion has a negative impact on the functioning of cork oak trees and 2) the magnitude of this negative impact is dependent on the predominant environmental conditions (i.e. precipitation amount), which can lead to amplifying, neutral or buffering effects of drought and invasion.

Mediterranean ecosystems are threatened by climate change and shrub encroachment. An increase in shrub cover can intensify the competition for water, aggravating the impacts of drought on ecosystem functioning. The effects of shrubs can be positive or negative, depending on the shrub species and density. We used a Mediterranean cork oak (Quercus suber) woodland to investigate the effects of increasing gum rockrose (Cistus ladanifer) cover on leaf carbon assimilation (Amax) and tree growth. The experiment consisted of a gradient of shrub cover, with four treatments: control, LD, MD, and HD, with 0%, 25%, 45%, and maximum shrub cover (>90%), respectively. Increasing shrub cover significantly decreased Amax in trees from HD (−15%) compared to control treatment, with intermediate effects on trees from LD (−5%) and MD (−12%). There was a large variability in tree growth, resulting in no significant effects of shrub cover, despite higher trunk diameter relative increments in LD (+40%), and lower in MD (−17%) and HD (−32%) compared to the control. The results indicate that a dense shrub cover (>90%) affected cork oak carbon assimilation and growth, while a low-to-medium shrub cover (≤45%) only induced mild intermediate effects. This information is important for the effective management of shrub density to improve the health and productivity of cork oak woodlands.

In Portugal, there are countless historic and legislative references about fire use in rural areas. This popular use of fire to manage the huddled of agricultural and forestry leftovers and small burns to clean the farmed lands continues nowadays, to be used in rural environments. However, these environments are becoming depopulated, with aged populations which are often isolated. Every year, in the social media, there are news about fatalities or injuries from elder people when using fire in the landscape. In 2018 the number of incidents were much higher than usual. For that, the objective of this work was to identify which were the factors that could explain this anomaly. To obtain a systematic data of the incidents, a thorough analysis of the regional and national newspapers, both in paper and online, was performed from the years 2008 to 2020. Variables like date, place, sex, age and other were gathered. All the incidents variables were complemented with meteorological data associated with wildfire risk, sociodemographic variables and the legislative context. From 2008 to 2020 there were 44 incidents, with 89% of fatalities. The anomaly of incidents in 2018 corresponded to 45% of cases. Most of the incidents (from 2008 to 2020) happened with elder (75%) and male (82%) people. The predominant months of occurrence were April and October particularly in the year 2018. The majority of fatalities (69%) occurred when the victim was burning alone. Legislative, social and meteorological analysis showed that the most probable factor that lead to the incidents anomaly was the extreme pressure to manage the vegetation, held by police forces, legislation and fines value if you didn’t comply.

Knowledge of plant hydraulic strategies (isohydric vs anisohydric) is crucial to predict the response of plants to changing environmental conditions, such as climate-change induced extreme drought. Several abiotic factors, such as evaporative demand, have been shown to seasonally modify the isohydricity of plants, however, the impact of biotic factors, such as plant-plant interactions on hydraulic strategies has seldom been explored. Here, we investigated adaptations in hydraulic strategies for two woody species in response to seasonal abiotic conditions, experimental drought and plant invasion in a Mediterranean cork oak (Quercus suber) ecosystem with a combined shrub invasion (Cistus ladanifer) and rain exclusion experiment. From summer to winter, the degree of isohydricity shifted from partial isohydric to anisohydric in Q. suber and inversely from strict anisohydric to partial isohydric for C. ladanifer. During drought, plant invasion significantly modified the hydraulic strategy of invaded Q. suber to a higher degree of anisohydricity with severe negative consequences for tree functioning, implying progressive leaf and xylem damage. The rain exclusion alone led to a non-significant increase in anisohydricity for both species. We demonstrate that the degree of isohydricity of plants is dynamically determined by the interplay of species-specific hydraulic traits and their abiotic and biotic environment.

The impact of interacting global change stressors on terrestrial ecosystems is hard to predict due to non-linear, amplifying, neutral or even buffering interaction effects. We investigated the effects of drought and plant invasion on Mediterranean cork oak (Quercus suber L.) ecosystem functioning and recovery with a combined rain exclusion (30-45 % reduction) and shrub (Cistus ladanifer L.) invasion experiment. As key parameter, we determined tree, shrub and ecosystem transpiration in four treatments: 1) cork oak control stands, 2) cork oaks with rain exclusion, 3) cork oaks invaded by shrubs and 4) cork oaks with rain exclusion and shrub invasion. Rain exclusion and plant invasion led to moderate, but neutral reductions of tree transpiration of 18 % (compared to control) during the mild summer drought in 2018. In 2019, the rain exclusion simulated the second driest year since 1950 for Southwestern (SW) Iberia. The interaction effect of drought and plant invasion was strongly amplifying, reducing tree transpiration by 47 %. Legacy effects on shrubs under the rain exclusion treatment led to a non-linear response during recovery from the severe drought in 2019. Invaded trees showed a delayed transpiration recovery (-51 % vs. control) due to strong competition with shrubs, while invaded trees with rain exclusion recovered to 75 % of the control. This buffering interaction response was caused by a weaker competition from drought-stressed shrubs. Given the projected increase in the frequency, intensity and duration of drought, an increasing non-linear impact on Mediterranean cork oak ecosystems is expected. Our results demonstrate that abiotic stressors modulate biotic interactions thereby impacting ecosystem functioning in a highly dynamic manner. Further efforts are thus needed to model and manage the impact of interacting global change stressors on terrestrial ecosystems.

Interaction effects of different stressors, such as extreme drought and plant invasion, can have detrimental effects on ecosystem functioning and recovery after drought. With ongoing climate change and increasing plant invasion, there is an urgent need to predict the short- and long-term interaction impacts of these stressors on ecosystems. We established a combined precipitation exclusion and shrub invasion (Cistus ladanifer) experiment in a Mediterranean cork oak (Quercus suber) ecosystem with four treatments: (1) Q. suber control; (2) Q. suber with rain exclusion; (3) Q. suber invaded by shrubs; and (4) Q. suber with rain exclusion and shrub invasion. As key parameter, we continuously measured ecosystem water fluxes. In an average precipitation year, the interaction effects of both stressors were neutral. However, the combination of imposed drought and shrub invasion led to amplifying interaction effects during an extreme drought by strongly reducing tree transpiration. Contrarily, the imposed drought reduced the competitiveness of the shrubs in the following recovery period, which buffered the negative effects of shrub invasion on Q. suber. Our results demonstrate the highly dynamic and nonlinear effects of interacting stressors on ecosystems and urges for further investigations on biotic interactions in a context of climate change pressures.

Mediterranean ecosystems, such as the savannah-type cork oak (Quercus suber) woodlands, are hotspots for climate change, as the highest impacts are forecasted for the Mediterranean region, mainly by more frequent and intense severe droughts. These ecosystems are also threatened by shrub encroachment, which might further decrease tree water availability and affect ecosystem functioning and resilience. Nevertheless, the combined effects of drought and shrub encroachment on ecosystems have seldom been investigated. A precipitation manipulation and shrub removal experiment was established in a cork oak woodland located in SE Portugal and invaded by the native shrub gum rockrose (Cistus ladanifer). Here we present and discuss the combined effects of drought and shrub encroachment on litterfall production of cork oak trees, an evergreen species, over two contrasting years, a wet year (2018) and a dry year (2019) and assess the nitrogen and phosphorus resorption efficiencies from senescent to green leaves. A previous study reported significant increases in cork oak’s nitrogen resorption efficiency in response to drought. Our preliminary results also indicate changes in nitrogen and phosphorus resorption efficiencies. An increase in nutrient resorption efficiency is likely to mitigate the limitation in nutrient uptake by the roots during drought, improving tree fitness in the short-term. However, it will probably exert a negative feedback on the nitrogen and phosphorus cycles in the long-term which might affect the ecosystem functioning under the forecasted droughts.

Mediterranean type ecosystems such as cork oak (Quercus suber) woodlands are currently threatened by extreme drought events and shrub encroachment in the Iberian Peninsula. Recently, the frequency of extreme droughts has increased with negative effects on many ecosystems. Decreasing soil water availability reduces growth and fitness of trees, and may eventually induce tree mortality. Shrub encroachment may further increase the competition for soil water, impacting tree vulnerability and resilience negatively. Yet, the synergistic effects of extreme droughts and shrub encroachment on ecosystems have rarely been investigated.We established a precipitation manipulation and shrub encroachment experiment in a cork oak stand to study the combined effects of the two environmental pressures. The cork oak woodland is located in Southeast Portugal and partially invaded by the native shrub gum rockrose (Cistus ladanifer). In December 2017, we installed rainout shelters (30 to 45% of precipitation reduction) in replicated cork oak stands invaded and uninvaded by gum rockrose, complemented by control plots with natural precipitation. In each treatment, the trees (n = 9) and shrubs (n = 9) were measured for water and carbon fluxes to reveal species-specific responses and competition effects under recurrent extreme drought.The hydrological year 2018 was characterised by above-average precipitation mainly caused by large spring rainfall events. Probably due to sufficient water supply, no clear treatment effects were evident. For example, minimum leaf water potentials (ΨPD) of the cork oak trees did not drop below −1.5 ± 0.1 MPa and maximum sap flux density was 2.1 ± 0.2 m3 m−2 day−1. Minimum ΨPD of the shrubs was three times lower (−3.5 ± 0.1 MPa) and maximum sap flux density over four-fold higher (8.8 ± 0.8 m3 m−2 day−1) than those of the trees, suggesting distinct species-specific behaviour. Reduced winter and spring precipitation, combined with a late onset of autumn rainfalls in 2019, led to a decrease in water input down to 66% (control) and 44% (drought) compared to the long-term average of 585 mm. In this dry year, negative synergistic effects of drought and shrub encroachment were expressed during the dry-down and drought period by a lower minimum ΨPD and an average sap flux density reduced by 50% (0.4 ± 0.1 m3 m−2 day−1) of invaded trees exposed to the experimental drought, compared to control trees (0.8 ± 0.1 m3 m−2 day−1). In sum, this resulted in a reduction of sap flux densities of the cork oaks by 25% (invaded), 23% (drought) and 34% (drought and invaded) over the course of the hydrological year 2019. The ongoing investigations aim to further determine the stress tolerance and critical physiological thresholds for both species and the entire ecosystem.

The impact of atmospheric reactive nitrogen (Nr) deposition on carbon (C) sequestration in soils and biomass of unfertilised, natural, semi-natural and forest ecosystems has been much debated. Many previous results of this dC / dN response were based on changes in carbon stocks from periodical soil and ecosystem inventories, associated with estimates of Nr deposition obtained from large-scale chemical transport models. This study and a companion paper (Flechard et al., 2019) strive to reduce uncertainties of N effects on C sequestration by linking multi-annual gross and net ecosystem productivity estimates from 40 eddy covariance flux towers across Europe to local measurement-based estimates of dry and wet Nr deposition from a dedicated collocated monitoring network. To identify possible ecological drivers and processes affecting the interplay between C and Nr inputs and losses, these data were also combined with in situ flux measurements of NO, N2O and CH4 fluxes, soil NO3− leaching sampling, as well as results of soil incubation experiments for N and greenhouse gas (GHG) emissions, surveys of available data from online databases and from the literature, together with forest ecosystem (BASFOR) modelling. Multi-year averages of net ecosystem productivity (NEP) in forests ranged from −70 to 826 g (C) m−2 yr−1 at total wet + dry inorganic Nr deposition rates (Ndep) of 0.3 to 4.3 g (N) m−2 yr−1; and from −4 to 361 g (C) m−2 yr−1 at Ndep rates of 0.1 to 3.1 g (N) m−2 yr−1 in short semi-natural vegetation (moorlands, wetlands and unfertilised extensively managed grasslands). The GHG budgets of the forests were strongly dominated by CO2 exchange, while CH4 and N2O exchange comprised a larger proportion of the GHG balance in short semi-natural vegetation. Nitrogen losses in the form of NO, N2O and especially NO3− were of the order of 10–20 % of Ndep at sites with Ndep 3 g (N) m−2 yr−1, indicating that perhaps one third of the sites were in a state of early to advanced N saturation. Net ecosystem productivity increased with Nr deposition up to 2–2.5 g (N) m−2 yr−1, with large scatter associated with a wide range in carbon sequestration efficiency (CSE, defined as the NEP / GPP ratio). At elevated Ndep levels (> 2.5 g (N) m−2 yr−1), where inorganic Nr losses were also increasingly large, NEP levelled off and then decreased. The apparent increase in NEP at low to intermediate Ndep levels was partly the result of geographical cross-correlations between Ndep and climate, indicating that the actual mean dC / dN response at individual sites was significantly lower than would be suggested by a simple, straightforward regression of NEP vs. Ndep.

Mediterranean ecosystems are hotspots for climate change, as the highest impacts are forecasted for the Mediterranean region, mainly by more frequent and intense severe droughts. Plant phenology is a good indicator of species' responses to climate change. In this study, we compared the spring phenology of cork oak trees (Quercus suber), an evergreen species, over 2 contrasting years, a mild year (2004) and a dry year (2005), which was the most severe drought since records exist. We evaluated the timing of occurrence, duration, and intensity of bud development, budburst, shoot elongation, trunk growth, and leaf senescence (phenophases) and assessed the nitrogen resorption efficiency from senescent to green leaves. The temperature was the main driver of budburst. Nevertheless, water had the main role of constraining all the other phenophases by strongly reducing the growing season length (−48 %) and consequently the tree growth. Basal area increment was the most affected growth variable (−36 %), although it occurred at a similar rate in the 2 years. Shoot elongation was also reduced (−21 %), yet elongation occurred at a higher rate in the dry year compared to the mild year. Leaf senescence during the bulk period was higher in the dry year, in which leaves were shed at the same rate over a longer period. Nitrogen concentrations in green and senescent leaves were affected by drought and nitrogen resorption efficiency increased remarkably (+22 %). Our results highlight the importance of studying different phenological metrics to improve our understanding of the ecosystem's responses to climate change. The faster dynamics observed in shoot elongation, while all other phenophases developed at the same rate, indicate that leaf area development is privileged in cork oak. Water availability was the main driver of spring growth in this Mediterranean ecosystem; however, growth may be affected by complex interplays between precipitation and temperature, such as higher temperatures during dry winters or heatwaves during spring, that are likely to result in water stress. Longer studies are needed to disentangle those interplays. Finally, a higher nitrogen resorption efficiency in response to drought appears to be an adaptive trait that mitigates the limitation in nitrogen uptake by the roots during drought and contributes to improving tree fitness in the short term but will probably exert a negative feedback on the nitrogen cycle in the long term, which might affect the ecosystem functioning under the forecasted droughts.

Mediterranean vegetation emits a wide range of biogenic volatile organic compounds (BVOCs) among which isoprenoids present quantitatively the most important compound class. Here, we investigated the isoprenoid emission from two Mediterranean Cistaceae shrubs, Halimium halimifolium and Cistus ladanifer, under controlled and natural conditions, respectively. For the first time, diurnal emission patterns of the diterpene kaurene were detected in real-time by Proton-Transfer-Reaction-Time-of-Flight-Mass-Spectrometer. Kaurene emissions were strongly variable among H. halimifolium plants, ranging from 0.01 ± 0.003 to 0.06 ± 0.01 nmol m−2 s−1 in low and high emitting individuals, respectively. They were in the same order of magnitude as monoterpene (0.01 ± 0.01 to 0.11 ± 0.04 nmol m−2 s−1) and sesquiterpene (0.01 ± 0.01 to 0.52 nmol m−2 s−1) emission rates. Comparable range and variability was found for C. ladanifer under natural conditions. Labelling with 13C-pyruvate suggested that emitted kaurene was not derived from de novo biosynthesis. The high kaurene content in leaves, the weak relationship with ecophysiological parameters and the tendency of higher emissions with increasing temperatures in the field indicate an emission from storage pools. This study highlights significant emissions of kaurene from two Mediterranean shrub species, indicating that the release of diterpenes into the atmosphere should probably deserve more attention in the future.

Trees scale leaf (A(L)) and xylem (A(X)) areas to couple leaf transpiration and carbon gain with xylem water transport. Some species are known to acclimate in A(L) : A(X) balance in response to climate conditions, but whether trees of different species acclimate in A(L) : A(X) in similar ways over their entire (continental) distributions is unknown. We analyzed the species and climate effects on the scaling of A(L) vs A(X) in branches of conifers (Pinus sylvestris, Picea abies) and broadleaved (Betula pendula, Populus tremula) sampled across a continental wide transect in Europe. Along the branch axis, A(L) and A(X) change in equal proportion (isometric scaling: b similar to 1) as for trees. Branches of similar length converged in the scaling of A(L) vs A(X) with an exponent of b = 0.58 across European climates irrespective of species. Branches of slow- growing trees from Northern and Southern regions preferentially allocated into new leaf rather than xylem area, with older xylem rings contributing to maintaining total xylem conductivity. In conclusion, trees in contrasting climates adjust their functional balance between water transport and leaf transpiration by maintaining biomass allocation to leaves, and adjusting their growth rate and xylem production to maintain xylem conductance.

International audience; Many studies have reported that hydraulic properties vary considerably between tree species, but little is known about their intraspecific variation and, therefore, their capacity to adapt to a warmer and drier climate. Here, we quantify phenotypic divergence and clinal variation for embolism resistance, hydraulic conductivity and branch growth, in four tree species, two angiosperms (Betula pendula, Populus tremula) and two conifers (Picea abies, Pinus sylvestris), across their latitudinal distribution in Europe. Growth and hydraulic efficiency varied widely within species and between populations. The variability of embolism resistance was in general weaker than that of growth and hydraulic efficiency, and very low for all species but Populus tremula. In addition, no and weak support for a safety vs. efficiency trade-off was observed for the angiosperm and conifer species, respectively. The limited variability of embolism resistance observed here for all species except Populus tremula, suggests that forest populations will unlikely be able to adapt hydraulically to drier conditions through the evolution of embolism resistance.

The objective of this study was to evaluate the litterfall deposition, the decomposition rate (KL) and the water holding capacity (WHC) of litter stored in the soil in two areas degraded by mining submitted to different methods of forest restoration: induction of natural regeneration (NR) and planting seedlings of native forest species (PS) with a forest fragment (FF) used as reference. The litterfall deposition was collected monthly over 12 months in collectors of 0.25 m2. The KL was estimated by the relation between annual litter production and litter stock in the soil surface. WHC was determined in the rainy and dry period. The litterfall deposition was lower in PS with values of 6.61 ± 0.20, 10.75 ± 0.52 and 11.83 ± 0.72 Mg ha-1 yr-1 for PS, NR and FF respectively. The KL and WHC were significantly lower in PS when compared to NR and FF, and WHC decreased significantly from the rainy season to the dry season. The induction of natural regeneration was the more effective restoration method for degraded areas regarding litterfall deposition, decomposition and water retention capacity, surpassing planting native tree species and approaching a native forest fragment info:eu-repo/semantics/publishedVersion

Background and aims During the recent decades, cork oak (Q. suber) mortality has been increasing in Mediterranean oak woodland endangering the economical and environmental sustainability of the “montado” ecosystem. This fact in combination with climate change and conversion of forestland to pasture may significantly affect the soil-atmosphere greenhouse gases (GHGs) exchange. Our study evaluates the impact of oak trees as compared to pasture on net ecosystem GHG (CH4, N2O, and CO2) exchange as well as the main environmental factors influencing this exchange.

A 10–20% decrease in annual precipitation is predicted in the Mediterranean basin, and in particular to the Iberian Peninsula, with foreseen effects on the exchange of soil-atmosphere greenhouse gases (GHGs; CO2, CH4, and N2O). To simulate this scenario, we setup an experimental design in the particularly dry period of 2008–2009 using rainfall exclusion and irrigation, to obtain plots receiving 110% (538 mm), 100% (493 mm) and 74% (365 mm) of the natural precipitation. Soil CO2 fluxes showed a strong increase from summer to autumn as a consequence of increasing soil heterotrophic respiration that resulted from rewetting. Fluxes of N2O were negligible. According to our data, soil was a permanent CH4 sink independent of the soil water content (in the range between 6–26% WFPS – water-filled pore space) and of soil temperature (in the range of 7–28°C), supporting the concept that seasonally dry ecosystems (Mediterranean) may represent a significant sink of atmospheric CH4. The study provides evidence that the 26% decrease or 10% increase in the ambient rainfall from annual precipitation of ca 500 mm did not significantly affect soil functionality and had a limited impact on soil-atmosphere net GHGs exchange in evergreen oak woodlands in southern Portugal.

Understanding the responses of cork oak (Quercus suber L.) to actual and predicted summer conditions is essential to determine the future sustainability of cork oak woodlands in Iberia. Thermal imaging may provide a rapid method for monitoring the extent of stress. The ecophysiology of cork trees was studied over three years. Three treatments were applied by means of rainfall capture and irrigation, with plots receiving 120%, 100%, or 80% of natural precipitation. Despite stomatal closure, detected using both thermal imaging and porometry, leaf water potential fell during the summer, most drastically during the third year of accumulative stress. The quantum efficiency (FPSII) and the maximum efficiency F V # =F M # of photosystem II also fell more intensely over the third summer, while nonphotochemical quenching (NPQ) increased. The reduced precipitation treatment sporadically further reduced leaf water potential, stomatal conductance (gs), IG (an index of gs derived from thermal imaging), UPSII ,a ndF V # =F M # ,a nd increased leaf temperature and NPQ. It is concluded that these are very resilient trees since they were only severely affected in the third year of severe drought (the third year registering 45% less rainfall than average), and removing 20% of rainfall had a limited impact.

*The effects of drought on the Amazon rainforest are potentially large but remain poorly understood. Here, carbon (C) cycling after 5 yr of a large-scale through-fall exclusion (TFE) experiment excluding about 50% of incident rainfall from an eastern Amazon rainforest was compared with a nearby control plot. *Principal C stocks and fluxes were intensively measured in 2005. Additional minor components were either quantified in later site measurements or derived from the available literature. *Total ecosystem respiration (R(eco)) and total plant C expenditure (PCE, the sum of net primary productivity (NPP) and autotrophic respiration (R(auto))), were elevated on the TFE plot relative to the control. The increase in PCE and R(eco) was mainly caused by a rise in R(auto) from foliage and roots. Heterotrophic respiration did not differ substantially between plots. NPP was 2.4 +/- 1.4 t C ha(-1) yr(-1) lower on the TFE than the control. Ecosystem carbon use efficiency, the proportion of PCE invested in NPP, was lower in the TFE plot (0.24 +/- 0.04) than in the control (0.32 +/- 0.04). *Drought caused by the TFE treatment appeared to drive fundamental shifts in ecosystem C cycling with potentially important consequences for long-term forest C storage.

P>1. The Amazon region may experience increasing moisture limitation over this century. Leaf dark respiration (R) is a key component of the Amazon rain forest carbon (C) cycle, but relatively little is known about its sensitivity to drought. 2. Here, we present measurements of R standardized to 25 degrees C and leaf morphology from different canopy heights over 5 years at a rain forest subject to a large-scale through-fall reduction (TFR) experiment, and nearby, unmodified Control forest, at the Caxiuana reserve in the eastern Amazon. 3. In all five post-treatment measurement campaigns, mean R at 25 degrees C was elevated in the TFR forest compared to the Control forest experiencing normal rainfall. After 5 years of the TFR treatment, R per unit leaf area and mass had increased by 65% and 42%, respectively, relative to pre-treatment means. In contrast, leaf area index (L) in the TFR forest was consistently lower than the Control, falling by 23% compared to the pre-treatment mean, largely because of a decline in specific leaf area (S). 4. The consistent and significant effects of the TFR treatment on R, L and S suggest that severe drought events in the Amazon, of the kind that may occur more frequently in future, could cause a substantial increase in canopy carbon dioxide emissions from this ecosystem to the atmosphere.

Tree mortality is a key factor influencing forest functions and dynamics, but our understanding of the mechanisms leading to mortality and the associated changes in tree growth rates are still limited. We compiled a new pan-continental tree-ring width database from sites where both dead and living trees were sampled (2970 dead and 4224 living trees from 190 sites, including 36 species), and compared early and recent growth rates between trees that died and those that survived a given mortality event. We observed a decrease in radial growth before death in ca. 84% of the mortality events. The extent and duration of these reductions were highly variable (1–100 years in 96% of events) due to the complex interactions among study species and the source(s) of mortality. Strong and long-lasting declines were found for gymnosperms, shade- and drought-tolerant species, and trees that died from competition. Angiosperms and trees that died due to biotic attacks (especially bark-beetles) typically showed relatively small and short-term growth reductions. Our analysis did not highlight any universal trade-off between early growth and tree longevity within a species, although this result may also reflect high variability in sampling design among sites. The intersite and interspecific variability in growth patterns before mortality provides valuable information on the nature of the mortality process, which is consistent with our understanding of the physiological mechanisms leading to mortality. Abrupt changes in growth immediately before death can be associated with generalized hydraulic failure and/or bark-beetle attack, while long-term decrease in growth may be associated with a gradual decline in hydraulic performance coupled with depletion in carbon reserves. Our results imply that growth-based mortality algorithms may be a powerful tool for predicting gymnosperm mortality induced by chronic stress, but not necessarily so for angiosperms and in case of intense drought or bark-beetle outbreaks., GGI by the Spanish Ministry of Economy and Competitiveness (Project AGL2014-61175-JIN)., AMH by the Spanish Ministry of Science and Innovation (Projects CGL2007-60120 and CSD2008-0040) and by the Spanish Ministry of Education via a FPU Scholarship., JJC, JCLC, and GSB by the Spanish Ministry of Economy (Projects CGL2015-69186-C2-1-R, CGL2013-48843-C2-2-R, and CGL2012-32965) and the EU (Project FEDER 0087 TRANSHABITAT).

Cork oak woodlands (montado) are agroforestry systems distributed all over the Mediterranean basin with a very important social, economic and ecological value. A generalized cork oak decline has been occurring in the last decades jeopardizing its future sustainability. It is unknown how loss of tree cover affects microbial processes that are consuming greenhouse gases in the montado ecosystem. The study was conducted under two different conditions in the natural understory of a cork oak woodland in center Portugal: under tree canopy (UC) and open areas without trees (OA). Fluxes of methane and nitrous oxide were measured with a static chamber technique. In order to quantify methanotrophs and bacteria capable of nitrous oxide consumption, we used quantitative real-time PCR targeting the pmoA and nosZ genes encoding the subunit of particulate methane mono-oxygenase and catalytic subunit of the nitrous oxide reductase, respectively. A significant seasonal effect was found on CH4 and N2O fluxes and pmoA and nosZ gene abundance. Tree cover had no effect on methane fluxes; conversely, whereas the UC plots were net emitters of nitrous oxide, the loss of tree cover resulted in a shift in the emission pattern such that the OA plots were a net sink for nitrous oxide. In a seasonal time scale, the UC had higher gene abundance of Type I methanotrophs. Methane flux correlated negatively with abundance of Type I methanotrophs in the UC plots. Nitrous oxide flux correlated negatively with nosZ gene abundance at the OA plots in contrast to that at the UC plots. In the UC soil, soil organic matter had a positive effect on soil extracellular enzyme activities, which correlated positively with the N2O flux. Our results demonstrated that tree cover affects soil properties, key enzyme activities and abundance of microorganisms and, consequently net CH4 and N2O exchange.

The aim of this study was to identify the sources and depth of water uptake by 15-years old Quercus suber L. trees in southern Portugal under a Mediterranean climate, measuring d 18 O and dD in the soil–plantatmosphere continuum. Evidence for hydraulic lift was substantiated by the daily fluctuations observed in Ys at 0.4 and 1 m depth and supported by similar d 18 O values found in tree xylem sap, soil water in the rhizosphere and groundwater. From 0.25 m down to a depth of 1 m, dD trends differed according to vegetation type, showing a more depleted value in soil water collected under the evergreen trees ()47&) than under dead grasses ()35&). The hypothesis of a fractionation process occurring in the soil due to diffusion of water vapour in the dry soil is proposed to explain the more depleted soil dD signature observed under trees. Hydraulically lifted water was estimated to account for 17–81% of the water used during the following day by tree transpiration at the peak of the drought season, i.e., 0.1–14 L tree )1 day )1 . Significant relationships found between xylem sap isotopic composition and leaf water potential in early September emphasized the positive impact of the redistribution of groundwater in the rhizosphere on tree water status.

Climate modelling studies predict that the rain forests of the Eastern Amazon basin are likely to experience reduc- tions in rainfall of up to 50% over the next 50-100 years. Efforts to predict the effects of changing climate, especially drought stress, on forest gas exchange are currently limited by uncertainty about the mechanism that controls stomatal closure in response to low soil moisture. At a through-fall exclusion experiment in Eastern Amazonia where water was experimentally excluded from the soil, we tested the hypothesis that plants are isohydric, that is, when water is scarce, the stomata act to prevent leaf water potential from dropping below a critical threshold level. We made diurnal measurements of leaf water potential ( Ψ l ), stomatal con- ductance ( g s ), sap flow and stem water potential ( Ψ stem ) in the wet and dry seasons. We compared the data with the predictions of the soil-plant-atmosphere (SPA) model, which embeds the isohydric hypothesis within its stomatal conductance algorithm. The model inputs for meteorology, leaf area index (LAI), soil water potential and soil-to-leaf hydraulic resistance ( R ) were altered between seasons in accordance with measured values. No optimization param- eters were used to adjust the model. This 'mechanistic' model of stomatal function was able to explain the individ- ual tree-level seasonal changes in water relations ( r 2 = 0.85, 0.90 and 0.58 for Ψ l , sap flow and g s , respectively). The model indicated that the measured increase in R was the dominant cause of restricted water use during the dry sea- son, resulting in a modelled restriction of sap flow four times greater than that caused by reduced soil water poten- tial. Higher resistance during the dry season resulted from an increase in below-ground resistance (including root and soil-to-root resistance) to water flow.

Determining climate change feedbacks from tropical rainforests requires an understanding of how carbon gain through photosynthesis and loss through respiration will be altered. One of the key changes that tropical rainforests may experience under future climate change scenarios is reduced soil moisture availability. In this study we examine if and how both leaf photosynthesis and leaf dark respiration acclimate following more than 12 years of experimental soil moisture deficit, via a through-fall exclusion experiment (TFE) in an eastern Amazonian rainforest. We find that experimentally drought-stressed trees and taxa maintain the same maximum leaf photosynthetic capacity as trees in corresponding control forest, independent of their susceptibility to drought-induced mortality. We hypothesise that photosynthetic capacity is maintained across all treatments and taxa to take advantage of short-lived periods of high moisture availability, when stomatal conductance (gs ) and photosynthesis can increase rapidly, potentially compensating for reduced assimilate supply at other times. Average leaf dark respiration (Rd ) was elevated in the TFE-treated forest trees relative to the control by 28.2±2.8% (mean ± one standard error). This mean Rd value was dominated by a 48.5±3.6% increase in the Rd of drought-sensitive taxa, and likely reflects the need for additional metabolic support required for stress-related repair, and hydraulic or osmotic maintenance processes. Following soil moisture deficit that is maintained for several years, our data suggest that changes in respiration drive greater shifts in the canopy carbon balance, than changes in photosynthetic capacity. This article is protected by copyright. All rights reserved. (Less)

a b s t r a c t This study details the physiological responses of cork oak (Quercus suber L.) to manipulated water inputs. Treatments named as dry, ambient and wet, which received 80, 100 and 120% of the annual precipitation, respectively, were applied to a Mediterranean woodland in southern Portugal. Tree ecophysiology and growth were monitored from 2003 to 2005. The impacts of the water manipulation were primarily observed in tree transpiration, especially dur- ing summer drought. Rainfall exclusion reduced the annual stand canopy transpiration by 10% over the 2-year study period, while irrigation increased it by 11%. The accumulated tree transpiration matched precipitation in spring 2004 and 2005 at the stand level, suggesting that cork oak trees rely on precip- itation water sources during the peak of the growing season. However, during the summer droughts, groundwater was the main water source for trees. Despite the significant differences in soil water content and tree transpiration, no treatment effects could be detected in leaf water potential and leaf gas exchange, except for a single event after spring irri- gations in the very dry year 2005. These irrigations were intentionally delayed to reduce dry spell duration during the peak of tree growing season. They resulted in an acute positive physiological response of trees from the wet treatment one week after the last irrigation event leading to a 32% raise of stem diame- ter increment the following months. Our results suggest that in a semi-arid environment precipitation changes in spring (amount and timing) have a stronger impact on cork oak physiology and growth than an overall change in the total annual precipitation. The extreme drought of 2005 had a negative impact on tree growth. The annual increment of tree trunk diameter in the ambient and dry treatments was reduced, while it increased for trees from the wet treatment. Water shortage also significantly reduced leaf area. The latter dropped by 10.4% in response to the extreme drought of 2005 in trees from the ambient treatment. The reduction was less pronounced in trees of the wet treatment (−7.6%), and more pronounced in trees of the dry treatment (−14.7%). Cork oak showed high resiliency to inter-annual precipitation variability. The annual accumulated tree transpiration, the minimum midday leaf water potential and the absolute amount of groundwater used Abbreviations: (A), carbon assimilation; (Amax), maximum carbon assimilation; (CV), crown volume; (DBH), diameter at breast height; (DBHinc), trunk diameter increment; (E), sap flux, tree transpiration; (E/DBH), normalized tree transpiration; (ET), potential evapotranspiration; (gs), stomatal conductance; (gsmax), maximum stomatal conduc- tance; (Gt), whole tree specific hydraulic conductance; (LAD), leaf area density; (LAI), leaf area index; (PAR), photosynthetically active radiation; (PCA), projected crown area; (SLA), specific leaf area; (SWC), soil water content; (T), air temperature; (VPD), vapour pressure deficit; (� ), leaf water potential; (� md), midday leaf water potential; (� pd), predawn leaf water potential; (�� ), sap flow driving force; (� ), osmotic potential; (� p), leaf turgor; (� s), leaf solute content; (RSC), regression slope comparison; (SEM), standard error of the mean. ∗ Corresponding author at: Instituto Dom Luiz, Centro de Geofisica da Universidade de Lisboa, Faculdade de Ciencias, Campo Grande, Ed. C8, piso 3, Sala 26, 1749-016

International audience; The static chamber method (non-flow-through-non-steady-state chambers) is the most common method to measure fluxes of methane (CH4) from soils. Laboratory comparisons to quantify errors resulting from chamber design, operation and flux calculation methods are rare. We tested fifteen chambers against four flux levels (FL) ranging from 200 to 2300 μg CH4 m−2 h−1. The measurements were conducted on a calibration tank using three quartz sand types with soil porosities of 53% (dry fine sand, S1), 47% (dry coarse sand, S2), and 33% (wetted fine sand, S3). The chambers tested ranged from 0.06 to 1.8 m in height, and 0.02 to 0.195 m3 in volume, 7 of them were equipped with a fan, and 1 with a vent-tube. We applied linear and exponential flux calculation methods to the chamber data and compared these chamber fluxes to the reference fluxes from the calibration tank. The chambers underestimated the reference fluxes by on average 33% by the linear flux calculation method (Rlin), whereas the chamber fluxes calculated by the exponential flux calculation method (Rexp) did not significantly differ from the reference fluxes (p < 0.05). The flux under- or overestimations were chamber specific and independent of flux level. Increasing chamber height, area and volume significantly reduced the flux underestimation (p < 0.05). Also, the use of non-linear flux calculation method significantly improved the flux estimation; however, simultaneously the uncertainty in the fluxes was increased. We provide correction factors, which can be used to correct the under- or overestimation of the fluxes by the chambers in the experiment.