Leaf morpho-anatomical adjustments in a Quercus pubescens forest after 10 years of partial rain exclusion in the field.

In the Mediterranean region, a reduction of annual precipitation and a longer and drier summer season are expected with climate change by the end of the century, eventually endangering forest survival. To cope with such rapid changes, trees may modulate their morpho-anatomical and physiological traits. In the present study, we focused on the variation in leaf gas exchange and different leaf morpho-anatomical functional traits of Quercus pubescens Willd. in summer using a long-term drought experiment in natura consisting of a dynamic rainfall exclusion system where trees have been submitted to amplified drought (AD) (~-30% of annual precipitation) since April 2012 and compared them with trees under natural drought (ND) in a Mediterranean forest. During the study, we analyzed net CO2 assimilation (An), stomatal conductance (gs), transpiration (E), water-use efficiency (WUE), stomatal size and density, density of glandular trichomes and non-glandular trichomes, thickness of the different leaf tissues, specific leaf area and leaf surface. Under AD, tree functioning was slightly impacted, since only An exhibited a 49% drop, while gs, E and WUE remained stable. The decrease in An under AD was regulated by concomitant lower stomatal density and reduced leaf thickness. Trees under AD also featured leaves with a higher non-glandular trichome density and a lower glandular trichome density compared with ND, which simultaneously limits transpiration and production costs. This study points out that Q. pubescens exhibits adjustments of leaf morpho-anatomical traits which can help trees to acclimate to AD scenarios as those expected in the future in the Mediterranean region.

Forest disturbances and climate constrain carbon allocation dynamics in trees.

Forest disturbances such as drought, fire, and logging affect the forest carbon dynamics and the terrestrial carbon sink. Forest mortality after disturbances creates uncertainties that need to be accounted for to understand forest dynamics and their associated C-sink. We combined data from permanent resampling plots and biomass oriented dendroecological plots to estimate time series of annual woody biomass growth (ABI) in several forests. ABI time series were used to benchmark a vegetation model to analyze dynamics in forest productivity and carbon allocation forced by environmental variability. The model implements source and sink limitations explicitly by dynamically constraining carbon allocation of assimilated photosynthates as a function of temperature and moisture. Bias in tree-ring reconstructed ABI increased back in time from data collection and with increasing disturbance intensity. ABI bias ranged from zero, in open stands without recorded mortality, to over 100% in stands with major disturbances such as thinning or snowstorms. Stand leaf area was still lower than in control plots decades after heavy thinning. Disturbances, species life-history strategy and climatic variability affected carbon-partitioning patterns in trees. Resprouting broadleaves reached maximum biomass growth at earlier ages than nonresprouting conifers. Environmental variability and leaf area explained much variability in woody biomass allocation. Effects of stand competition on C-allocation were mediated by changes in stand leaf area except after major disturbances. Divergence between tree-ring estimated and simulated ABI were caused by unaccounted changes in allocation or misrepresentation of some functional process independently of the model calibration approach. Higher disturbance intensity produced greater modifications of the C-allocation pattern, increasing error in reconstructed biomass dynamics. Legacy effects from disturbances decreased model performance and reduce the potential use of ABI as a proxy to net primary productivity. Trait-based dynamics of C-allocation in response to environmental variability need to be refined in vegetation models.

The role of maternal age, growth, and environment in shaping offspring performance in an aerial conifer seed bank.

PREMISE: Maternal effects have been demonstrated to affect offspring performance in many organisms, and in plants, seeds are important mediators of these effects. Some woody plant species maintain long-lasting canopy seed banks as an adaptation to wildfires. Importantly, these seeds stored in serotinous cones are produced by the mother plant under varying ontogenetic and physiological conditions. METHODS: We sampled the canopy seed bank of a highly serotinous population of Pinus pinaster to test whether maternal age and growth and the environmental conditions during each crop year affected seed mass and ultimately germination and early survival. After determining retrospectively the year of each seed cohort, we followed germination and early survival in a semi-natural common garden. RESULTS: Seed mass was related to maternal age and growth at the time of seed production; i.e., slow-growing, older mothers had smaller seeds, and fast-growing, young mothers had larger seeds, which could be interpreted either as a proxy of senescence or as a maternal strategy. Seed mass had a positive effect on germination success, but aside from differences in seed mass, maternal age had a negative effect and diameter had a positive effect on germination timing and subsequent survival. CONCLUSIONS: The results highlight the importance of maternal conditions combined with seed mass in shaping seedling establishment. Our findings open new insights in the offspring performance deriving from long-term canopy seed banks, which may have high relevance for plant adaptation.

Old forests and old carbon: A case study on the stand dynamics and longevity of aboveground carbon.

Most information on the ecology of oak-dominated forests in Europe comes from forests altered for centuries because remnants of old-growth forests are rare. Disturbance and recruitment regimes in old-growth forests provide information on forest dynamics and their effects on long-term carbon storage. In an old-growth Quercus petraea forest in northwestern Spain, we inventoried three plots and extracted cores from 166 live and dead trees across canopy classes (DBH ≥ 5 cm). We reconstructed disturbance dynamics for the last 500 years from tree-ring widths. We also reconstructed past dynamics of above ground biomass (AGB) and recent AGB accumulation rates at stand level using allometric equations. From these data, we present a new tree-ring-based approach to estimate the age of carbon stored in AGB. The oldest tree was at least 568 years, making it the oldest known precisely-dated oak to date and one of the oldest broadleaved trees in the Northern Hemisphere. All plots contained trees over 400 years old. The disturbance regime was dominated by small, frequent releases with just a few more intense disturbances that affected ≤20% of trees. Oak recruitment was variable but rather continuous for 500 years. Carbon turnover times ranged between 153 and 229 years and mean carbon ages between 108 and 167 years. Over 50% of AGB (150 Mg·ha-1) persisted ≥100 years and up to 21% of AGB (77 Mg·ha-1) ≥300 years. Low disturbance rates and low productivity maintained current canopy oak dominance. Absence of management or stand-replacing disturbances over the last 500 years resulted in high forest stability, long carbon turnover times and long mean carbon ages. Observed dynamics and the absence of shade-tolerant species suggest that oak dominance could continue in the future. Our estimations of long-term carbon storage at centennial scales in unmanaged old-growth forests highlights the importance of management and natural disturbances for the global carbon cycle.

Holm oak death is accelerated but not sudden and expresses drought legacies.

The increase in abiotic and biotic stress driven by global change threatens forest ecosystems and challenges understanding of mechanisms producing mortality. Phytophthora spp. like P. cinnamomi (PHYCI) are among the most lethal pathogens for many woody species including Quercus spp. Dynamics of biotic agents and their hosts are complex and influenced by climatic conditions. We analysed radial growth trends of dead and live adult Quercus ilex trees from agrosilvopastoral open woodlands under intense land-use. A pronounced warming trend since the 1980s has coincided in these woodlands with high oak mortality rates generally attributed to PHYCI. Yet, tree mortality and latency of the pathogen could be expressed at variable time spans, whereas, like in many other forests worldwide, tree death could also be explained by other factors like drought. PHYCI was isolated from roots of all dead oaks from one region. Trees were younger than generally believed and ages of dead trees ranged between 38 and 230 years. Growth of dead trees reached a tipping point in 1980 and 1990 coincident with two-year extraordinary droughts. These dates set the start of growth declines up to 30 years before tree death. Live trees did not exhibit any recent growth decline. Tree growth was highly sensitive to climatic variability associated with water stress and climate-growth relationships suggested phenological changes since the 1980s. Live and dead trees showed differences in their sensitivity to moisture availability and temperature. The sensitivity of growth to climate was partially related to site environmental conditions. Simulated gross and net primary productivity were higher in live sites with less atmospheric demand for water. Tree death was not sudden but a slow multiannual process as expressed by radial growth declines likely triggered by drought. Regardless of the causal agent or mechanism, the observed mortality affected trees exhibiting negative drought and land-use legacies.

Climate Change Synchronizes Growth and iWUE Across Species in a Temperate-Submediterranean Mixed Oak Forest.

Tree species have good tolerance to a range of environmental conditions, though their ability to respond and persist to environmental changes is dramatically reduced at the rear-edge distribution limits. At those edges, gene flow conferring adaptation is impaired due to lack of populations at lower latitudes. Thus, trees mainly rely on phenotypic changes to buffer against long-term environmental changes. Interspecific hybridization may offer an alternative mechanism in the generation of novel genetic recombinants that could be particularly valuable to ensure persistence in geographically isolated forests. In this paper, we take advantage of the longevity of a temperate-submediterranean mixed-oak forest to explore the long-term impact of environmental changes on two different oak species and their hybrid. Individual trees were genetically characterized and classified into three groups: pure Quercus petraea (Matt.), Liebl, pure Q. pyrenaica Willd, and hybrids. We calculated basal area increment and intrinsic water-use efficiency (iWUE) from tree-ring width and δ13C per genetic group, respectively. Tree-growth drivers were assessed using correlation analyses and generalized linear mixed models for two contrasting climatic periods: (1880-1915, colder with [CO2] < 303 ppm; and 1980-2015, warmer with [CO2] > 338 ppm). The three genetic groups have increased radial growth and iWUE during the last decades, being the least drought-tolerant QuPe the most sensitive species to water stress. However, no significant differences were found among genetic groups neither in mean growth rate nor in mean iWUE. Furthermore, little differences were found in the response to climate among groups. Genetic groups only differed in the relationship between δ13C and temperature and precipitation during the earlier period, but such a difference disappeared during the recent decades. Climate change may have promoted species-level convergence as a response to environment-induced growth limitations, which translated in synchronized growth and response to climate as well as a tighter stomatal control and increased iWUE across coexisting oak species.

Low growth resilience to drought is related to future mortality risk in trees.

Severe droughts have the potential to reduce forest productivity and trigger tree mortality. Most trees face several drought events during their life and therefore resilience to dry conditions may be crucial to long-term survival. We assessed how growth resilience to severe droughts, including its components resistance and recovery, is related to the ability to survive future droughts by using a tree-ring database of surviving and now-dead trees from 118 sites (22 species, >3,500 trees). We found that, across the variety of regions and species sampled, trees that died during water shortages were less resilient to previous non-lethal droughts, relative to coexisting surviving trees of the same species. In angiosperms, drought-related mortality risk is associated with lower resistance (low capacity to reduce impact of the initial drought), while it is related to reduced recovery (low capacity to attain pre-drought growth rates) in gymnosperms. The different resilience strategies in these two taxonomic groups open new avenues to improve our understanding and prediction of drought-induced mortality.

Water potential control of turgor-driven tracheid enlargement in Scots pine at its xeric distribution edge.

The extent to which water availability can be used to predict the enlargement and final dimensions of xylem conduits remains an open issue. We reconstructed the time course of tracheid enlargement in Pinus sylvestris trees in central Spain by repeated measurements of tracheid diameter on microcores sampled weekly during a 2 yr period. We analyzed the role of water availability in these dynamics empirically through time-series correlation analysis and mechanistically by building a model that simulates daily tracheid enlargement rate and duration based on Lockhart's equation and water potential as the sole input. Tracheid enlargement followed a sigmoid-like time course, which varied intra- and interannually. Our empirical analysis showed that final tracheid diameter was strongly related to water availability during tracheid enlargement. The mechanistic model was calibrated and successfully validated (R2  = 0.92) against the observed tracheid enlargement time course. The model was also able to reproduce the seasonal variations of tracheid enlargement rate, duration and final diameter (R2  = 0.84-0.99). Our results support the hypothesis that tracheid enlargement and final dimensions can be modeled based on the direct effect of water potential on turgor-driven cell expansion. We argue that such a mechanism is consistent with other reported patterns of tracheid dimension variation.

Synergistic abiotic and biotic stressors explain widespread decline of Pinus pinaster in a mixed forest.

Global change potentially increases forest vulnerability. Different abiotic and biotic factors may interact to cause forest decline and accelerated tree mortality. We studied a mixed Mediterranean continental forest where Pinus pinaster Ait. (maritime pine) shows widespread decline to analyse the role of different abiotic and biotic factors on health status and growth dynamics both at the individual and plot levels. We also analysed stand composition and regeneration of tree species to check whether there is a change in species dominance. Fungal pathogens were seldom present and we detected no pervasive fungi or insect infestation and no presence of pathogens like Heterobasidion or Phytophthora. Infection of hemiparasite plants like Viscum album L. (mistletoe) can reduce leaf area and its abundance is generally considered an expression of host decline. Yet, the existence among declining trees of high defoliation levels without mistletoe, but not vice versa, suggests that defoliation in response to some abiotic stressor could be a predisposing factor preceding mistletoe infection. Compared to healthy trees, declining and dead trees exhibited higher defoliation rates, smaller needles and lower recent growth with steeper negative trends. Dead and declining trees showed similar negative growth trends since the early 1990s droughts, which we interpreted as early warning signals anticipating mortality of currently declining trees in the near future. Mortality of maritime pine extending across all size classes, the lower presence of this species in the smallest size classes and its lack of regeneration suggest it is potentially losing its current dominance and being replaced by other co-occurring, more drought-tolerant species. Our results unravelled that maritime pine decline seems to be mainly driven by a combination of predisposing and inciting abiotic factors (microenvironment and drought stress) and biotic factors (mistletoe). The absence of widespread fungal pathogens suggests that they may have a minor role on pine decline acting only eventually as contributing factors. Although there could be other interrelations among factors or other biotic agents at play, our results strongly suggest that water stress plays a major role in the decline process of the dominant species on an ecosystem with strong land-use legacies.

Geographical adaptation prevails over species-specific determinism in trees' vulnerability to climate change at Mediterranean rear-edge forests.

Climate change may reduce forest growth and increase forest mortality, which is connected to high carbon costs through reductions in gross primary production and net ecosystem exchange. Yet, the spatiotemporal patterns of vulnerability to both short-term extreme events and gradual environmental changes are quite uncertain across the species' limits of tolerance to dryness. Such information is fundamental for defining ecologically relevant upper limits of species tolerance to drought and, hence, to predict the risk of increased forest mortality and shifts in species composition. We investigate here to what extent the impact of short- and long-term environmental changes determines vulnerability to climate change of three evergreen conifers (Scots pine, silver fir, Norway spruce) and two deciduous hardwoods (European beech, sessile oak) tree species at their southernmost limits of distribution in the Mediterranean Basin. Finally, we simulated future forest growth under RCP 2.6 and 8.5 emission scenarios using a multispecies generalized linear mixed model. Our analysis provides four key insights into the patterns of species' vulnerability to climate change. First, site climatic marginality was significantly linked to the growth trends: increasing growth was related to less climatically limited sites. Second, estimated species-specific vulnerability did not match their a priori rank in drought tolerance: Scots pine and beech seem to be the most vulnerable species among those studied despite their contrasting physiologies. Third, adaptation to site conditions prevails over species-specific determinism in forest response to climate change. And fourth, regional differences in forests vulnerability to climate change across the Mediterranean Basin are linked to the influence of summer atmospheric circulation patterns, which are not correctly represented in global climate models. Thus, projections of forest performance should reconsider the traditional classification of tree species in functional types and critically evaluate the fine-scale limitations of the climate data generated by global climate models.

Divergent phenological and leaf gas exchange strategies of two competing tree species drive contrasting responses to drought at their altitudinal boundary.

In Mediterranean mountains, Pinus sylvestris L. is expected to be displaced under a warming climate by more drought-tolerant species such as the sub-Mediterranean Quercus pyrenaica Willd. Understanding how environmental factors drive tree physiology and phenology is, therefore, essential to assess the effect of changing climatic conditions on the performance of these species and, ultimately, their distribution. We compared the cambial and leaf phenology and leaf gas exchange of Q. pyrenaica and P. sylvestris at their altitudinal boundary in Central Spain and assessed the environmental variables involved. Results indicate that P. sylvestris cambial phenology was more sensitive to weather conditions (temperature at the onset and water deficit at the end of the growing season) than Q. pyrenaica. On the other hand, Q. pyrenaica cambial and leaf phenology were synchronized and driven by photoperiod and temperatures. Pinus sylvestris showed lower photosynthetic nitrogen-use efficiency and higher intrinsic water-use efficiency than Q. pyrenaica as a result of a tighter stomatal control in response to summer dry conditions, despite its less negative midday leaf water potentials. These phenological and leaf gas exchange responses evidence a stronger sensitivity to drought of P. sylvestris than that of Q. pyrenaica, which may therefore hold a competitive advantage over P. sylvestris under the predicted increase in recurrence and intensity of drought events. On the other hand, both species could benefit from warmer springs through an advanced phenology, although this effect could be limited in Q. pyrenaica if it maintains a photoperiod control over the onset of xylogenesis.

Early-Warning Signals of Individual Tree Mortality Based on Annual Radial Growth.

Tree mortality is a key driver of forest dynamics and its occurrence is projected to increase in the future due to climate change. Despite recent advances in our understanding of the physiological mechanisms leading to death, we still lack robust indicators of mortality risk that could be applied at the individual tree scale. Here, we build on a previous contribution exploring the differences in growth level between trees that died and survived a given mortality event to assess whether changes in temporal autocorrelation, variance, and synchrony in time-series of annual radial growth data can be used as early warning signals of mortality risk. Taking advantage of a unique global ring-width database of 3065 dead trees and 4389 living trees growing together at 198 sites (belonging to 36 gymnosperm and angiosperm species), we analyzed temporal changes in autocorrelation, variance, and synchrony before tree death (diachronic analysis), and also compared these metrics between trees that died and trees that survived a given mortality event (synchronic analysis). Changes in autocorrelation were a poor indicator of mortality risk. However, we found a gradual increase in inter-annual growth variability and a decrease in growth synchrony in the last ∼20 years before mortality of gymnosperms, irrespective of the cause of mortality. These changes could be associated with drought-induced alterations in carbon economy and allocation patterns. In angiosperms, we did not find any consistent changes in any metric. Such lack of any signal might be explained by the relatively high capacity of angiosperms to recover after a stress-induced growth decline. Our analysis provides a robust method for estimating early-warning signals of tree mortality based on annual growth data. In addition to the frequently reported decrease in growth rates, an increase in inter-annual growth variability and a decrease in growth synchrony may be powerful predictors of gymnosperm mortality risk, but not necessarily so for angiosperms.

Endophytic Fungi as Pretreatment to Enhance Enzymatic Hydrolysis of Olive Tree Pruning.

Olive tree pruning, as one of the most abundant lignocellulosic residues in Mediterranean countries, has been evaluated as a source of sugars for fuel and chemicals production. A mild acid pretreatment has been combined with a fungal pretreatment using either two endophytes (Ulocladium sp. and Hormonema sp.) or a saprophyte (Trametes sp. I-62). The use of endophytes is based on the important role that some of them play during the initial stages of wood decomposition. Without acid treatment, fungal pretreatment with Ulocladium sp. provided a nonsignificant enhancement of 4.6% in glucose digestibility, compared to control. When a mild acid hydrolysis was carried out after fungal pretreatments, significant increases in glucose digestibility from 4.9% to 12.0% (compared to control without fungi) were observed for all fungal pretreatments, with maximum values yielded by Hormonema sp. However, despite the observed digestibility boost, the total sugar yields (taking into account solid yield) were not significantly increased by the pretreatments. Nevertheless, based on these preliminary improvements in digestibility, this work proves the potential of endophytic fungi to boost the production of sugar from olive tree pruning, which would add an extra value to the bioeconomy of olive crops.

Effects of Competition, Drought Stress and Photosynthetic Productivity on the Radial Growth of White Spruce in Western Canada.

Understanding the complex interactions of competition, climate warming-induced drought stress, and photosynthetic productivity on the radial growth of trees is central to linking climate change impacts on tree growth, stand structure and in general, forest productivity. Using a mixed modeling approach, a stand-level photosynthetic production model, climate, stand competition and tree-ring data from mixedwood stands in western Canada, we investigated the radial growth response of white spruce [Picea glauca (Moench.) Voss] to simulated annual photosynthetic production, simulated drought stress, and tree and stand level competition. The long-term (~80-year) radial growth of white spruce was constrained mostly by competition, as measured by total basal area, with minor effects from drought. There was no relation of competition and drought on tree growth but dominant trees increased their growth more strongly to increases in modeled photosynthetic productivity, indicating asymmetric competition. Our results indicate a co-limitation of drought and climatic factors inhibiting photosynthetic productivity for radial growth of white spruce in western Canada. These results illustrate how a modeling approach can separate the complex factors regulating both multi-decadal average radial growth and interannual radial growth variations of white spruce, and contribute to advance our understanding on sustainable management of mixedwood boreal forests in western Canada.

Xylem and Leaf Functional Adjustments to Drought in Pinus sylvestris and Quercus pyrenaica at Their Elevational Boundary.

Climatic scenarios for the Mediterranean region forecast increasing frequency and intensity of drought events. Consequently, a reduction in Pinus sylvestris L. distribution range is projected within the region, with this species being outcompeted at lower elevations by more drought-tolerant taxa such as Quercus pyrenaica Willd. The functional response of these species to the projected shifts in water availability will partially determine their performance and, thus, their competitive success under these changing climatic conditions. We studied how the cambial and leaf phenology and xylem anatomy of these two species responded to a 3-year rainfall exclusion experiment set at their elevational boundary in Central Spain. Additionally, P. sylvestris leaf gas exchange, water potential and carbon isotope content response to the treatment were measured. Likewise, we assessed inter-annual variability in the studied functional traits under control and rainfall exclusion conditions. Prolonged exposure to drier conditions did not affect the onset of xylogenesis in either of the studied species, whereas xylem formation ceased 1-3 weeks earlier in P. sylvestris. The rainfall exclusion had, however, no effect on leaf phenology on either species, which suggests that cambial phenology is more sensitive to drought than leaf phenology. P. sylvestris formed fewer, but larger tracheids under dry conditions and reduced the proportion of latewood in the tree ring. On the other hand, Q. pyrenaica did not suffer earlywood hydraulic diameter changes under rainfall exclusion, but experienced a cumulative reduction in latewood width, which could ultimately challenge its hydraulic performance. The phenological and anatomical response of the studied species to drought is consistent with a shift in resource allocation under drought stress from xylem to other sinks. Additionally, the tighter stomatal control and higher intrinsic water use efficiency observed in drought-stressed P. sylvestris may eventually limit carbon uptake in this species. Our results suggest that both species are potentially vulnerable to the forecasted increase in drought stress, although P. sylvestris might experience a higher risk of drought-induced decline at its low elevational limit.

A synthesis of radial growth patterns preceding tree mortality.

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.

Risky future for Mediterranean forests unless they undergo extreme carbon fertilization.

Forest performance is challenged by climate change but higher atmospheric [CO2 ] (ca ) could help trees mitigate the negative effect of enhanced water stress. Forest projections using data assimilation with mechanistic models are a valuable tool to assess forest performance. Firstly, we used dendrochronological data from 12 Mediterranean tree species (six conifers and six broadleaves) to calibrate a process-based vegetation model at 77 sites. Secondly, we conducted simulations of gross primary production (GPP) and radial growth using an ensemble of climate projections for the period 2010-2100 for the high-emission RCP8.5 and low-emission RCP2.6 scenarios. GPP and growth projections were simulated using climatic data from the two RCPs combined with (i) expected ca ; (ii) constant ca  = 390 ppm, to test a purely climate-driven performance excluding compensation from carbon fertilization. The model accurately mimicked the growth trends since the 1950s when, despite increasing ca , enhanced evaporative demands precluded a global net positive effect on growth. Modeled annual growth and GPP showed similar long-term trends. Under RCP2.6 (i.e., temperatures below +2 °C with respect to preindustrial values), the forests showed resistance to future climate (as expressed by non-negative trends in growth and GPP) except for some coniferous sites. Using exponentially growing ca and climate as from RCP8.5, carbon fertilization overrode the negative effect of the highly constraining climatic conditions under that scenario. This effect was particularly evident above 500 ppm (which is already over +2 °C), which seems unrealistic and likely reflects model miss-performance at high ca above the calibration range. Thus, forest projections under RCP8.5 preventing carbon fertilization displayed very negative forest performance at the regional scale. This suggests that most of western Mediterranean forests would successfully acclimate to the coldest climate change scenario but be vulnerable to a climate warmer than +2 °C unless the trees developed an exaggerated fertilization response to [CO2 ].

Stand competition determines how different tree species will cope with a warming climate.

Plant-plant interactions influence how forests cope with climate and contribute to modulate species response to future climate scenarios. We analysed the functional relationships between growth, climate and competition for Pinus sylvestris, Quercus pyrenaica and Quercus faginea to investigate how stand competition modifies forest sensitivity to climate and simulated how annual growth rates of these species with different drought tolerance would change throughout the 21st century. Dendroecological data from stands subjected to thinning were modelled using a novel multiplicative nonlinear approach to overcome biases related to the general assumption of a linear relationship between covariates and to better mimic the biological relationships involved. Growth always decreased exponentially with increasing competition, which explained more growth variability than climate in Q. faginea and P. sylvestris. The effect of precipitation was asymptotic in all cases, while the relationship between growth and temperature reached an optimum after which growth declined with warmer temperatures. Our growth projections indicate that the less drought-tolerant P. sylvestris would be more negatively affected by climate change than the studied sub-Mediterranean oaks. Q. faginea and P. sylvestris mean growth would decrease under all the climate change scenarios assessed. However, P. sylvestris growth would decline regardless of the competition level, whereas this decrease would be offset by reduced competition in Q. faginea. Conversely, Q. pyrenaica growth would remain similar to current rates, except for the warmest scenario. Our models shed light on the nature of the species-specific interaction between climate and competition and yield important implications for management. Assuming that individual growth is directly related to tree performance, trees under low competition would better withstand the warmer conditions predicted under climate change scenarios but in a variable manner depending on the species. Thinning following an exponential rule may be desirable to ensure long-term conservation of high-density Mediterranean woodlands, particularly in drought-limited sites.

Modeling acclimation of photosynthesis to temperature in evergreen conifer forests.

• In this study, we used a canopy photosynthesis model which describes changes in photosynthetic capacity with slow temperature-dependent acclimations. • A flux-partitioning algorithm was applied to fit the photosynthesis model to net ecosystem exchange data for 12 evergreen coniferous forests from northern temperate and boreal regions. • The model accounted for much of the variation in photosynthetic production, with modeling efficiencies (mean > 67%) similar to those of more complex models. The parameter describing the rate of acclimation was larger at the northern sites, leading to a slower acclimation of photosynthesis to temperature. The response of the rates of photosynthesis to air temperature in spring was delayed up to several days at the coldest sites. Overall photosynthesis acclimation processes were slower at colder, northern locations than at warmer, more southern, and more maritime sites. • Consequently, slow changes in photosynthetic capacity were essential to explaining variations of photosynthesis for colder boreal forests (i.e. where acclimation of photosynthesis to temperature was slower), whereas the importance of these processes was minor in warmer conifer evergreen forests.