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A major component of climate change is an increase in temperature and precipitation variability. Over the last few decades, an increase in the frequency of extremely warm temperatures and drought severity has been observed across Europe. These warmer and drier conditions may reduce productivity and trigger compositional shifts in forest communities. However, we still lack a robust, biogeographical characterization of the negative impacts of climate extremes, such as droughts on forests. In this context, we investigated the impact of the 2017 summer drought on European forests. The normalized difference vegetation index (NDVI) was used as a proxy of forest productivity and was related to the standardized precipitation evapotranspiration index, which accounts for the temperature effects of the climate water balance. The spatial pattern of NDVI reduction in 2017 was largely driven by the extremely warm summer for parts of the central and eastern Mediterranean Basin (Italian and Balkan Peninsulas). The vulnerability to the 2017 summer drought was heterogeneously distributed over Europe, and topographic factors buffered some of the negative impacts. Mediterranean forests dominated by oak species were the most negatively impacted, whereas Pinus pinaster was the most resilient species. The impact of drought on the NDVI decreased at high elevations and mainly on east and north-east facing slopes. We illustrate how an adequate characterization of the coupling between climate conditions and forest productivity (NDVI) allows the determination of the most vulnerable areas to drought. This approach could be widely used for other extreme climate events and when considering other spatially resolved proxies of forest growth and health.
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Secas , Temperatura Alta , Mudança Climática , Europa (Continente) , Florestas , ÁrvoresRESUMO
The future trajectory of atmospheric CO2 concentration depends on the development of the terrestrial carbon sink, which in turn is influenced by forest dynamics under changing environmental conditions. An in-depth understanding of model sensitivities and uncertainties in non-steady-state conditions is necessary for reliable and robust projections of forest development and under scenarios of global warming and CO2 enrichment. Here, we systematically assessed if a biogeochemical process-based model (3D-CMCC-CNR), which embeds similarities with many other vegetation models, applied in simulating net primary productivity (NPP) and standing woody biomass (SWB), maintained a consistent sensitivity to its 55 input parameters through time, during forest ageing and structuring as well as under climate change scenarios. Overall, the model applied at three contrasting European forests showed low sensitivity to the majority of its parameters. Interestingly, model sensitivity to parameters varied through the course of >100 yr of simulations. In particular, the model showed a large responsiveness to the allometric parameters used for initialize forest carbon and nitrogen pools early in forest simulation (i.e., for NPP up to ~37%, 256 g C·m-2 ·yr-1 and for SWB up to ~90%, 65 Mg C/ha, when compared to standard simulation), with this sensitivity decreasing sharply during forest development. At medium to longer time scales, and under climate change scenarios, the model became increasingly more sensitive to additional and/or different parameters controlling biomass accumulation and autotrophic respiration (i.e., for NPP up to ~30%, 167 g C·m-2 ·yr-1 and for SWB up to ~24%, 64 Mg C/ha, when compared to standard simulation). Interestingly, model outputs were shown to be more sensitive to parameters and processes controlling stand development rather than to climate change (i.e., warming and changes in atmospheric CO2 concentration) itself although model sensitivities were generally higher under climate change scenarios. Our results suggest the need for sensitivity and uncertainty analyses that cover multiple temporal scales along forest developmental stages to better assess the potential of future forests to act as a global terrestrial carbon sink.
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Carbono , Mudança Climática , Biomassa , Ciclo do Carbono , FlorestasRESUMO
In this study, we investigated the role of climatic variability and atmospheric nitrogen deposition in driving long-term tree growth in canopy beech trees along a geographic gradient in the montane belt of the Italian peninsula, from the Alps to the southern Apennines. We sampled dominant trees at different developmental stages (from young to mature tree cohorts, with tree ages spanning from 35 to 160 years) and used stem analysis to infer historic reconstruction of tree volume and dominant height. Annual growth volume (GV ) and height (GH ) variability were related to annual variability in model simulated atmospheric nitrogen deposition and site-specific climatic variables, (i.e. mean annual temperature, total annual precipitation, mean growing period temperature, total growing period precipitation, and standard precipitation evapotranspiration index) and atmospheric CO2 concentration, including tree cambial age among growth predictors. Generalized additive models (GAM), linear mixed-effects models (LMM), and Bayesian regression models (BRM) were independently employed to assess explanatory variables. The main results from our study were as follows: (i) tree age was the main explanatory variable for long-term growth variability; (ii) GAM, LMM, and BRM results consistently indicated climatic variables and CO2 effects on GV and GH were weak, therefore evidence of recent climatic variability influence on beech annual growth rates was limited in the montane belt of the Italian peninsula; (iii) instead, significant positive nitrogen deposition (Ndep ) effects were repeatedly observed in GV and GH ; the positive effects of Ndep on canopy height growth rates, which tended to level off at Ndep values greater than approximately 1.0 g m-2 y-1 , were interpreted as positive impacts on forest stand above-ground net productivity at the selected study sites.
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Fagus/crescimento & desenvolvimento , Nitrogênio/metabolismo , Teorema de Bayes , Demografia , Florestas , Itália , Modelos Biológicos , Nitrogênio/análise , Temperatura , Fatores de TempoRESUMO
We present a global assessment of the relationships between the short-wave surface albedo of forests, derived from the MODIS satellite instrument product at 0.5° spatial resolution, with simulated atmospheric nitrogen deposition rates (Ndep ), and climatic variables (mean annual temperature Tm and total annual precipitation P), compiled at the same spatial resolution. The analysis was performed on the following five forest plant functional types (PFTs): evergreen needle-leaf forests (ENF); evergreen broad-leaf forests (EBF); deciduous needle-leaf forests (DNF); deciduous broad-leaf forests (DBF); and mixed-forests (MF). Generalized additive models (GAMs) were applied in the exploratory analysis to assess the functional nature of short-wave surface albedo relations to environmental variables. The analysis showed evident correlations of albedo with environmental predictors when data were pooled across PFTs: Tm and Ndep displayed a positive relationship with forest albedo, while a negative relationship was detected with P. These correlations are primarily due to surface albedo differences between conifer and broad-leaf species, and different species geographical distributions. However, the analysis performed within individual PFTs, strengthened by attempts to select 'pure' pixels in terms of species composition, showed significant correlations with annual precipitation and nitrogen deposition, pointing toward the potential effect of environmental variables on forest surface albedo at the ecosystem level. Overall, our global assessment emphasizes the importance of elucidating the ecological mechanisms that link environmental conditions and forest canopy properties for an improved parameterization of surface albedo in climate models.
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Atmosfera/química , Clima , Florestas , Modelos Teóricos , Nitrogênio/metabolismo , Luz Solar , Simulação por Computador , Análise de Regressão , Especificidade da EspécieRESUMO
Partially mycoheterotrophic (mixotrophic) plants gain carbon from both photosynthesis and their mycorrhizal fungi. This is considered an ancestral state in the evolution of full mycoheterotrophy, but little is known about this nutrition, and especially about the physiological balance between photosynthesis and fungal C gain. To investigate possible compensation between photosynthesis and mycoheterotrophy in the Mediterranean mixotrophic orchid Limodorum abortivum, fungal colonization was experimentally reduced in situ by fungicide treatment. We measured photosynthetic pigments of leaves, stems, and ovaries, as well as the stable C isotope compositions (a proxy for photosynthetic C gain) of seeds and the sizes of ovaries and seeds. We demonstrate that (1) in natural conditions, photosynthetic pigments are most concentrated in ovaries; (2) pigments and photosynthetic C increase in ovaries when fungal C supply is impaired, buffering C limitations and allowing the same development of ovaries and seeds as in natural conditions; and (3) responses to light of pigment and (13)C contents in ovaries shift from null responses in natural conditions to responses typical of autotrophic plants in treated L. abortivum, demonstrating photoadaptation and enhanced use of light in the latter. L. abortivum thus preferentially feeds on fungi in natural conditions, but employs compensatory photosynthesis to buffer fungal C limitations and allow seed development.
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Fenômenos Fisiológicos da Nutrição , Orchidaceae/fisiologia , Evolução Biológica , Isótopos de Carbono/metabolismo , Micorrizas/fisiologia , Orchidaceae/metabolismo , Orchidaceae/microbiologia , Fotossíntese/fisiologia , Pigmentos Biológicos/metabolismo , Estruturas Vegetais/metabolismoRESUMO
Water transport, mechanical support and storage are the vital functions provided by the xylem. These functions are carried out by different cells, exhibiting significant anatomical variation not only within species but also within individual trees. In this study, we used a comprehensive dataset to investigate the consistency of predicted hydraulic vessel diameter widening values in relation to the distance from the tree apex, represented by the relationship Dh â Lß (where Dh is the hydraulic vessel diameter, L the distance from the stem apex and ß the scaling exponent). Our analysis involved 10 Fagus sylvatica L. trees sampled at two distinct sites in the Italian Apennines. Our results strongly emphasize that vessel diameter follows a predictable pattern with the distance from the stem apex and ß ~ 0.20 remains consistent across cambial age and climates. This finding supports the hypothesis that trees do not alter their axial configuration represented by scaling of vessel diameter to compensate for hydraulic limitations imposed by tree height during growth. The study further indicates that within-tree variability significantly contributes to the overall variance of the vessel diameter-stem length exponent. Understanding the factors that contribute to the intraindividual variability in the widening exponent is essential, particularly in relation to interspecific responses and adaptations to drought stress.
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Câmbio , Clima , Fagus , Caules de Planta , Xilema , Fagus/crescimento & desenvolvimento , Fagus/fisiologia , Fagus/anatomia & histologia , Xilema/crescimento & desenvolvimento , Xilema/anatomia & histologia , Xilema/fisiologia , Câmbio/crescimento & desenvolvimento , Caules de Planta/crescimento & desenvolvimento , Caules de Planta/anatomia & histologia , Caules de Planta/fisiologia , Água/metabolismo , Árvores/crescimento & desenvolvimento , Árvores/fisiologia , Árvores/anatomia & histologia , ItáliaRESUMO
Previous favorable climate conditions stimulate tree growth making some forests more vulnerable to hotter droughts. This so-called structural overshoot may contribute to forest dieback, but there is little evidence on its relative importance depending on site conditions and tree species because of limited field data. Here, we analyzed remote sensing (NDVI) and tree-ring width data to evaluate the impacts of the 2017 drought on canopy cover and growth in mixed Mediterranean forests (Fraxinus ornus, Quercus pubescens, Acer monspessulanum, Pinus pinaster) located in southern Italy. Legacy effects were assessed by calculating differences between observed and predicted basal area increment (BAI). Overall, the growth response of the study stands to the 2017 drought was contingent on site conditions and species characteristics. Most sites presented BAI and canopy cover reductions during the drought. Growth decline was followed by a quick recovery and positive legacy effects, particularly in the case of F. ornus. However, we found negative drought legacies in some species (e.g., Q. pubescens, A. monspessulanum) and sites. In those sites showing negative legacies, high growth rates prior to drought in response to previous wet winter-spring conditions may have predisposed trees to drought damage. Vice versa, the positive drought legacy found in some F. ornus site was linked to post-drought growth release due to Q. pubescens dieback and mortality. Therefore, we found evidences of structural drought overshoot, but it was restricted to specific sites and species. Our findings highlight the importance of considering site settings such as stand composition, pre-drought conditions and different tree species when studying structural overshoot. Droughts contribute to modify the composition and dynamics in mixed forests.
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Secas , Florestas , Árvores , Árvores/fisiologia , Itália , Quercus/crescimento & desenvolvimento , Quercus/fisiologia , Mudança Climática , Pinus/fisiologia , Pinus/crescimento & desenvolvimento , Monitoramento Ambiental , Fraxinus/fisiologia , Fraxinus/crescimento & desenvolvimento , Acer/crescimento & desenvolvimento , Acer/fisiologiaRESUMO
Temperate and boreal forests in the Northern Hemisphere cover an area of about 2 x 10(7) square kilometres and act as a substantial carbon sink (0.6-0.7 petagrams of carbon per year). Although forest expansion following agricultural abandonment is certainly responsible for an important fraction of this carbon sink activity, the additional effects on the carbon balance of established forests of increased atmospheric carbon dioxide, increasing temperatures, changes in management practices and nitrogen deposition are difficult to disentangle, despite an extensive network of measurement stations. The relevance of this measurement effort has also been questioned, because spot measurements fail to take into account the role of disturbances, either natural (fire, pests, windstorms) or anthropogenic (forest harvesting). Here we show that the temporal dynamics following stand-replacing disturbances do indeed account for a very large fraction of the overall variability in forest carbon sequestration. After the confounding effects of disturbance have been factored out, however, forest net carbon sequestration is found to be overwhelmingly driven by nitrogen deposition, largely the result of anthropogenic activities. The effect is always positive over the range of nitrogen deposition covered by currently available data sets, casting doubts on the risk of widespread ecosystem nitrogen saturation under natural conditions. The results demonstrate that mankind is ultimately controlling the carbon balance of temperate and boreal forests, either directly (through forest management) or indirectly (through nitrogen deposition).
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Carbono/metabolismo , Clima , Ecossistema , Atividades Humanas , Árvores/metabolismo , Nitrogênio/metabolismoRESUMO
The link between biodiversity and ecosystem function can depend on environmental conditions. This contingency can impede our ability to predict how biodiversity-ecosystem function (BEF) relationships will respond to future environmental change, causing a clear need to explore the processes underlying shifts in BEF relationships across large spatial scales and broad environmental gradients. We compiled a dataset on five functional traits (maximum height, wood density, specific leaf area [SLA], seed size, and xylem vulnerability to embolism [P50]), covering 78%-90% of the tree species in the National Forest Inventory from Italy, to test (i) how a water limitation gradient shapes the functional composition and diversity of forests, (ii) how functional composition and diversity of trees relate to forest annual increment via mass ratio and complementarity effects, and (iii) how the relationship between functional diversity and annual increment varies between Mediterranean and temperate climate regions. Functional composition varied with water limitation; tree communities tended to have more conservative traits in sites with higher levels of water limitation. The response of functional diversity differed among traits and climatic regions but among temperate forest plots, we found a consistent increase of functional diversity with water limitation. Tree diversity was positively associated with annual increment of Italian forests through a combination of mass ratio and niche complementarity effects, but the relative importance of these effects depended on the trait and range of climate considered. Specifically, niche complementarity effects were more strongly associated with annual increment in the Mediterranean compared to temperate forests. Synthesis: Overall, our results suggest that biodiversity mediates forest annual increment under water-limited conditions by promoting beneficial interactions between species and complementarity in resource use. Our work highlights the importance of conserving functional diversity for future forest management to maintain forest annual increment under the expected increase in intensity and frequency of drought.
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We investigated the dendroclimatic response of a Pinus heldreichii metapopulation distributed over a wide elevation interval (from 882 to 2143 m a.s.l.), spanning from low mountain to upper subalpine vegetation belts in the southern Italian Apennines. The tested hypothesis is that wood growth along an elevational gradient is non-linearly related to air temperature. During three years of fieldwork (2012-2015) at 24 sites, we collected wood cores from a total of 214 pine trees with diameter at breast height from 19 to 180 cm (average 82.7 ± 32.9 cm). We used a combination of tree-ring and genetic methods to reveal factors involved in growth acclimation using a space-for-time approach. Scores from canonical correspondence analysis were used to combine individual tree-ring series into four composite chronologies related to air temperature along the elevation gradient. Overall, the June dendroclimatic response followed a bell-shaped thermal niche curve, increasing until a peak around 13-14 °C. A similarly bell-shaped response was found with previous autumn air temperature, and both dendroclimatic signals interacted with stem size and growth rates, generating a divergent growth response between the top and the bottom of the elevation gradient. Increased tree growth in the upper subalpine belt was consistent with the consequences of increasing air temperature under no drought stress. A positive link was uncovered between pine growth at all elevations and April mean temperature, with trees growing at the lowest elevations showing the strongest growth response. No elevational genetic differences were found, hence long-lived tree species with small geographical ranges may reverse their climatic response between the lower and upper bioclimatic zones of their environmental niche. Our study revealed a high resistance and acclimation capability of Mediterranean forest stands, and such low vulnerability to changing climatic conditions highlights the potential to store carbon in these ecosystems for the coming decades.
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Pinus , Árvores , Temperatura , Ecossistema , FlorestasRESUMO
The objective of this study is to globally assess the effects of atmospheric nitrogen deposition and climate, associated with rising levels of atmospheric CO2 , on the variability of carbon isotope discrimination (Δ(13) C), and intrinsic water-use efficiency (iWUE) of angiosperm and conifer tree species. Eighty-nine long-term isotope tree-ring chronologies, representing 23 conifer and 13 angiosperm species for 53 sites worldwide, were extracted from the literature, and used to obtain long-term time series of Δ(13) C and iWUE. Δ(13) C and iWUE were related to the increasing concentration of atmospheric CO2 over the industrial period (1850-2000) and to the variation of simulated atmospheric nitrogen deposition and climatic variables over the period 1950-2000. We applied generalized additive models and linear mixed-effects models to predict the effects of climatic variables and nitrogen deposition on Δ(13) C and iWUE. Results showed a declining Δ(13) C trend in the angiosperm and conifer species over the industrial period and a 16.1% increase of iWUE between 1850 and 2000, with no evidence that the increased rate was reduced at higher ambient CO2 values. The temporal variation in Δ(13) C supported the hypothesis of an active plant mechanism that maintains a constant ratio between intercellular and ambient CO2 concentrations. We defined linear mixed-effects models that were effective to describe the variation of Δ(13) C and iWUE as a function of a set of environmental predictors, alternatively including annual rate (Nrate ) and long-term cumulative (Ncum ) nitrogen deposition. No single climatic or atmospheric variable had a clearly predominant effect, however, Δ(13) C and iWUE showed complex dependent interactions between different covariates. A significant association of Nrate with iWUE and Δ(13) C was observed in conifers and in the angiosperms, and Ncum was the only independent term with a significant positive association with iWUE, although a multi-factorial control was evident in conifers.
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Wildfires have large-scale and profound effects on forest ecosystems, and they force burned forest areas toward a wide range of post-fire successional trajectories from simple reduction of ecosystem functions to transitions to other stable non-forest states. Fire disturbances represent a key driver of changes in forest structure and composition due to post-fire succession processes, thus contributing to modify ecosystem resilience to subsequent disturbances. Here, we aimed to provide useful insights into wildfire severity and post-fire recovery processes at the European continental scale, contributing to improved description and interpretation of large-scale wildfire spatial patterns and their effects on forest ecosystems in the context of climate change. We analyzed fire severity and short-term post-fire vegetation recovery patterns across the European forests between 2004 and 2015 using Corine Land Cover Forest classes and bioregions, based on MODIS-derived spectral metrics of the relativized burn ratio (RBR), normalized difference vegetation index (NDVI) and relative recovery indicator (RRI). The RBR-based fire severity showed geographic differences and interannual variability in the Boreal bioregion compared to that in other biogeographic regions. The NBR-based RRI showed a slower post-fire vegetation recovery rate with respect to the NDVI, highlighting the differential sensitivities of the analyzed remote sensing-spectral metrics. Moreover, the RRI showed a significant decreasing trend during the observation period, suggesting a growing lag in post-fire vegetation recovery across European forests.
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Incêndios , Incêndios Florestais , Benchmarking , Ecossistema , Florestas , Tecnologia de Sensoriamento RemotoRESUMO
Past anthropogenic disturbances lowered the altitudinal distribution of the Mediterranean Fagus sylvatica forests below 2,000 m a.s.l. Accordingly, our current understanding of the southern distribution range of F. sylvatica forests is restricted to managed stands below this elevation, neglecting relic forests growing above. This study has shed light on the structure and species assemblage of an unmanaged relict subalpine F. sylvatica stand growing within the core of its southernmost glacial refugia and at its highest species range elevation limit (2,140 m a.s.l.) in southern Apennines (Italy). Here, tree biometric attributes and understory species abundances were assessed in eight permanent plots systematically positioned from 1,650 to 2,130 m a.s.l. In the subalpine belt, F. sylvatica had formed a dense clonal stem population that was layered downward on the steepest slopes. The density and spatial aggregation of the stems were increased, while their stature and crown size were decreased. Above 2,000 m, changes in tree growth patterns, from upright single-stemmed to procumbent multi-stemmed, and canopy layer architecture, with crowns packed and closer to the floor, were allowed for the persistence of understory herbaceous species of biogeographic interest. Clonal layering represents an adaptive regeneration strategy for the subalpine belt environmental constraints not previously recognized in managed Mediterranean F. sylvatica forests. The clonal structure and unique species assemblage of this relic forest highlight the value of its inclusion in the priority areas networks, representing a long-term management strategy of emblematic glacial and microclimatic refugia.
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Understanding processes controlling forest dynamics has become particularly important in the context of ongoing climate change, which is altering the ecological fitness and resilience of species worldwide. However, whether forest communities would be threatened by projected macroclimate change or unaffected due to the controlling effect of local site conditions is still a matter for debate. After all, forest canopy buffer climate extremes and promote microclimatic conditions, which matters for functional plant response, and act as refugia for understory species in a changing climate. Yet precisely how microclimatic conditions change in response to climate warming will depend on the extent to which vegetation structure and local topography shape air and soil temperature. In this study, we posited that forest microclimatic buffering is sensitive to local topographic conditions and canopy cover, and using meteorological stations equipped with data-loggers we measured this effect during 1 year across a climate gradient (considering aspect as a surrogate of local topography) in a Mediterranean beech treeline growing in contrasting aspects in southern Italy. During the growing season, the below-canopy near-ground temperatures were, on average, 2.4 and 1.0 °C cooler than open-field temperatures for south and north-west aspects, respectively. Overall, the temperature offset became more negative (that is, lower under-canopy temperatures at the treeline) as the open-field temperature increased, and more positive (that is, higher under-canopy temperatures at the treeline) as the open-field temperature decreased. The buffering effect was particularly evident for the treeline on the south-facing slope, where cooling of near-ground temperature was as high as 8.6 °C for the maximum temperature (in August the offset peaked at 10 °C) and as high as 2.5 °C for the average temperature. In addition, compared to the south-facing slope, the northern site exhibited less decoupling from free-air environment conditions and low variability in microclimate trends that closely track the free-air biophysical environment. Although such a decoupling effect cannot wholly isolate forest climatic conditions from macroclimate regional variability in the south-facing treeline, it has the potential to partly offset the regional macroclimatic warming experienced in the forest understory due to anthropogenic climate change.
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Old-growth mountain forests represent an ideal setting for studying long-term impacts of climate change. We studied the few remnants of old-growth forests located within the Pollino massif (southern Italy) to evaluate how the growth of conspecific young and old trees responded to climate change. We investigated two conifer species (Abies alba and Pinus leucodermis) and two hardwood species (Fagus sylvatica and Quercus cerris). We sampled one stand per species along an altitudinal gradient, ranging from a drought-limited low-elevation hardwood forest to a cold-limited subalpine pine forest. We used a dendrochronological approach to characterize the long-term growth dynamics of old (age > 120 years) versus young (age < 120 years) trees. Younger trees grew faster than their older conspecifics during their juvenile stage, regardless of species. Linear mixed effect models were used to quantify recent growth trends (1950-2015) and responses to climate for old and young trees. Climate sensitivity, expressed as radial growth responses to climate during the last three decades, partially differed between species because high spring temperatures enhanced conifer growth, whereas F. sylvatica growth was negatively affected by warmer spring conditions. Furthermore, tree growth was negatively impacted by summer drought in all species. Climate sensitivity differed between young and old trees, with younger trees tending to be more sensitive in P. leucodermis and A. alba, whereas older F. sylvatica trees were more sensitive. In low-elevation Q. cerris stands, limitation of growth due to drought was not related to tree age, suggesting symmetric water competition. We found evidence for a fast-growth trend in young individuals compared with that in their older conspecifics. Notably, old trees tended to have relatively stable growth rates, showing remarkable resistance to climate warming. These responses to climate change should be recognized when forecasting the future dynamics of old-growth forests for their sustainable management.
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Fagus , Florestas , Idoso de 80 Anos ou mais , Mudança Climática , Secas , Humanos , ÁrvoresRESUMO
We analyzed the Italian National Forest Inventory data set to evaluate the interdependence of forest productivity, tree species richness (used to indicate biodiversity), climate, and soil factors. We tested the hypotheses that the relationship between biodiversity and forest productivity is positive and significant for all forests in Italy and whether the relationship is the same for forests growing in the temperate and Mediterranean bioclimatic domains (regions) of Italy. We used generalized additive models to explore the univariate response curves for the data and then performed structural equation modeling (SEM) and multi-group SEM analyses to evaluate the relationship between biodiversity and productivity. We found that the SEM model for the entire dataset explained about 60% of the variation in forest productivity. In addition, the variation associated with species richness was greater than variation due to climatic factors and the variation in climate factors was greater than the variation in soil factors (all relative to their contributions to productivity). The multi-group SEM showed a more predominant effect of biodiversity and climate on productivity in Mediterranean compared to temperate forests. In both cases, we observed a moderate effect of soil (factors) on forest productivity. Our results support the hypothesis that increasing tree diversity in forests could help reduce the effects of climate warming and enhance ecosystem productivity in the Mediterranean region.
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Biodiversidade , Clima , Florestas , Árvores/classificação , Biomassa , Mudança Climática , Ecossistema , Monitoramento Ambiental , Itália , Região do Mediterrâneo , Árvores/fisiologiaRESUMO
Forest decline induced by climate change is a global phenomenon that affects many tree species, mainly in drought-prone areas as the Mediterranean region. In southern Italy, several oak species have shown decline symptoms and elevated mortality since the 2000s due to drought stress. However, it remains to be answered whether decline occurred alone or whether a pathogen was also involved. To this aim, we compared two coexisting oak species in a forest located in southern Italy which are assumed to be less (Quercus cerris) and more tolerant to drought (Quercus pubescens). We sampled fifteen couples of neighboring declining (D) and non-declining (ND) trees of both species. Wood cores were taken from all trees to perform dendrochronological analyses to detect the decline onset and link it to potential climatic drivers. Carbon isotope ratios (d13C) were analyzed in wood of the two vigor classes to compare their water-use efficiency. Phytophthora presence was also assessed in soil samples from ten D-ND couples of trees per species. The oak species most affected by drought-induced decline in terms of leaf shedding and mortality was Q. cerris, i.e., the least tolerant to drought. In both species, the D trees showed a reduced growth rate compared with ND trees from 2000 onward when drought and warming intensified. Q. pubescens showed higher growth sensitivity to precipitation, temperature and drought than Q. cerris. This sensitivity to climate was magnified in D trees whose growth decreased in response to warm and dry conditions during the prior winter and the late summer. The Q. pubescens D trees were more efficient in their water use than ND trees before the growth divergence between D and ND trees amplified. In the studied area, Phytophthora quercina was isolated from 40% of the sampled trees, and tended to be more frequent amongst ND than amongst D trees. Our data suggests that droughts and warm summer conditions triggered oak decline. The high prevalence of P. quercina in the studied area warrants further study as a potential predisposing factor.
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Observations on the net carbon exchange of forests in the European Mediterranean region, measured recently by the eddy covariance method, have revived interest in a phenomenon first characterized on agricultural and forest soils in East Africa in the 1950s and 1960s by H. F. Birch and now often referred to as the "Birch effect." When soils become dry during summer because of lack of rain, as is common in regions with Mediterranean climate, or are dried in the laboratory in controlled conditions, and are then rewetted by precipitation or irrigation, there is a burst of decomposition, mineralization and release of inorganic nitrogen and CO(2). In forests in Mediterranean climates in southern Europe, this effect has been observed with eddy covariance techniques and soil respiration chambers at the stand and small plot scales, respectively. Following the early work of Birch, laboratory incubations of soils at controlled temperatures and water contents have been used to characterize CO(2) release following the rewetting of dry soils. A simple empirical model based on laboratory incubations demonstrates that the amount of carbon mineralized over one year can be predicted from soil temperature and precipitation regime, provided that carbon lost as CO(2) is taken into account. We show that the amount of carbon returned to the atmosphere following soil rewetting can reduce significantly the annual net carbon gain by Mediterranean forests.
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Dióxido de Carbono/metabolismo , Solo , Água/metabolismo , Dióxido de Carbono/química , Ecossistema , Região do Mediterrâneo , Modelos Biológicos , Estações do Ano , Árvores/crescimento & desenvolvimento , Árvores/metabolismoRESUMO
Hydraulic theory suggests that tall trees are at greater risk of drought-triggered death caused by hydraulic failure than small trees. In addition the drop in growth, observed in several tree species prior to death, is often interpreted as an early-warning signal of impending death. We test these hypotheses by comparing size, growth, and wood-anatomy patterns of living and now-dead trees in two Italian oak forests showing recent mortality episodes. The mortality probability of trees is modeled as a function of recent growth and tree size. Drift-diffusion-jump (DDJ) metrics are used to detect early-warning signals. We found that the tallest trees of the anisohydric Italian oak better survived drought contrary to what was predicted by the theory. Dead trees were characterized by a lower height and radial-growth trend than living trees in both study sites. The growth reduction of now-dead trees started about 10 years prior to their death and after two severe spring droughts during the early 2000s. This critical transition in growth was detected by DDJ metrics in the most affected site. Dead trees were also more sensitive to drought stress in this site indicating different susceptibility to water shortage between trees. Dead trees did not form earlywood vessels with smaller lumen diameter than surviving trees but tended to form wider latewood vessels with a higher percentage of vessel area. Since living and dead trees showed similar competition we did not expect that moderate thinning and a reduction in tree density would increase the short-term survival probability of trees.
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The objectives of this study were to provide a quantitative description of the long-term effects of environmental variability on xylem functional traits, in order to better assess xylem capacity to change in response to climate change. Twenty-six sites throughout the world, primarily in Europe, were chosen where results from long-term measurements of anatomical traits were previously published. Published data on long-term xylem anatomy (conduit size and density) and ring width variability were compiled across a range of tree species, which was subsequently related to variability in temperature, precipitation and nitrogen deposition rates across the study sites using generalized additive models and Bayesian methods. We found some appreciable relationships between xylem traits (conduit area Ac and conduit density Dc) and environmental variables; whereas combined trait indices (lumen fraction: Ac × Dc and vessel composition: Ac/Dc) were found to be rather constant across a wide range of environmental conditions and to be decoupled from tree growth rates. Overall, results suggested xylem traits coordinated towards a homeostasis in xylem function, which appeared to act across a wide range of environmental conditions. Results showed also nitrogen deposition was associated with xylem traits and vessel composition: increased nitrogen availability due to nitrogen deposition might facilitate construction of a xylem structure efficient for water transport, and concurrently provide capacity to withstand the risks of drought-induced embolism.