Drought legacies in mixed Mediterranean forests: Analysing the effects of structural overshoot, functional traits and site factors.

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.

Water-limited environments affect the association between functional diversity and forest productivity.

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.

Bell-shaped tree-ring responses to air temperature drive productivity trends in long-lived mountain Mediterranean pines.

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.

Topoclimate effect on treeline elevation depends on the regional framework: A contrast between Southern Alps (New Zealand) and Apennines (Italy) forests.

Deciphering the spatial patterns of alpine treelines is critical for understanding the ecosystem processes involved in the persistence of tree species and their altitudinal limit. Treelines are thought to be controlled by temperature, and other environmental variables but they have rarely been investigated in regions with different land-use change legacies. Here, we systematically investigated treeline elevation in the Apennines (Italy) and Southern Alps (New Zealand) with contrasting human history but similar biogeographic trajectories, intending to identify distinct drivers that affect their current elevation and highlight their respective peculiarities. Over 3622 km of Apennines, treeline elevation was assessed in 302 mountain peaks and in 294 peaks along 4504 km of Southern Alps. The major difference between the Southern Alps and Apennines treeline limit is associated with their mountain aspects. In the Southern Alps, the scarcely anthropized Nothofagus treeline elevation was higher on the warmer equator-facing slopes than on the pole-facing ones. Contrary to what would be expected based on temperature limitation, the elevation of Fagus sylvatica treelines in the Apennines was higher on colder, pole-facing slopes than on human-shaped equator-facing, warmer mountainsides. Pervasive positive correlations were found between treeline elevation and temperature in the Southern Alps but not in the Apennines. While the position of the Fagus and Nothofagus treelines converge on similar isotherms of annual average temperature, a striking isothermal difference between the temperatures of the hottest month on which the two taxonomic groups grow exists. We conclude that actual treeline elevation reflects the ecological processes driven by a combination of local-scale topoclimatic conditions, and human disturbance legacy. Predicting dynamic processes affecting current and future alpine treeline position requires further insight into the modulating influences that are currently understood at a regional scale.

Clonality drives structural patterns and shapes the community assemblage of the Mediterranean Fagus sylvatica subalpine belt.

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.

Towards sustainable management of forest residues in the southern Apennine Mediterranean mountain forests: a scenario-based approach.

Key message: Managing forest residues according to the carbon content of the soil helps to minimize the ecological footprint of their removal. Context: In Mediterranean mountain ecosystems, unsustainable harvesting of wood residues might contribute to land degradation, carbon, and nutrient depletion in forest soils. Aims: This study aimed to assess the amount of forest biomass residues that should be left on-site to minimize the depletion of soil fertility. Methods: We estimated the availability of biomass residues in the public forest land of the Basilicata region of Southern Italy by collecting stand-scale inventory attributes from forest management plans. Subsequently, we quantified the amount of forest biomass residue released by implementing a scenario-based approach. Results: Approximately 5800 m3 year-1 of forest residues could be potentially available for bio-based industries at the regional scale within the next 10 years. Such residues mainly belong to broadleaved forest types, having a high variability in their soil organic stock (228.5-705.8 Mg C ha-1) and altitudinally spanning from 400 to 1500 m a.s.l. In these forests, the simulated scenarios displayed a wide range of average harvestable residues from 2.5 to 5.5 m3 ha-1, containing approximately 1.1 to 2.1 Mg ha-1 of organic carbon. Conclusion: Our study suggests that forest management plans are a useful source of information to estimate the available forest biomass residues consistently. In southern Mediterranean mountain forests, the management of forest residues according to soil carbon content helps to minimize the environmental impact and increase their sustainability.

Biogeographic variability in wildfire severity and post-fire vegetation recovery across the European forests via remote sensing-derived spectral metrics.

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.

Declines in canopy greenness and tree growth are caused by combined climate extremes during drought-induced dieback.

Several dieback episodes triggered by droughts are revealing the high vulnerability of Mediterranean forests, manifested as declines in growth, increased defoliation, and rising mortality rates. Understanding forest responses to such climate extreme events is of high priority for predicting their future vegetation dynamics. We examined how remotely sensed measures of vegetation activity (NDVI, Normalized Difference Vegetation Index) and radial growth (BAI, basal area increment) responded to climate extreme events. We considered tree (Pinus sylvestris, Quercus pubescens, Quercus frainetto) and shrub (Juniperus phoenicea) populations from Italy and Spain showing recent dieback phenomena. Two components of drought, namely elevated atmospheric demand (VPD, vapor pressure deficit) and low soil moisture were analyzed in nearby stands showing or not showing dieback symptoms. Dieback stands exhibited lower NDVI values than non-dieback stands. NDVI and BAI were positively related in all sites except for the dieback stand of Q. frainetto that was negatively related. Such NDVI-BAI linkages were related to specific time windows, which could be useful for identifying when climatic conditions have the greatest influence on vegetation. Growth decline occurred in response to increasing VPD, but responses differed among species. J. phoenicea was the most negatively impacted by higher VPD, whereas oaks responded to soil moisture. A high VPD was related to stronger growth reduction in dieback P. sylvestris trees regardless of soil moisture changes. We highlighted that coupling between proxies of forest productivity (NDVI, BAI) allows better understanding and forecasting of drought-induced dieback phenomena in forests and shrublands. Scaling up from tree to stand levels might be feasible when using the maximum growing season NDVI, which can be applied for retrospective modeling of the impact of drought stress on forest productivity and tree growth.

Mediterranean old-growth forests exhibit resistance to climate warming.

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.

Topography modulates near-ground microclimate in the Mediterranean Fagus sylvatica treeline.

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.

The impact of drought spells on forests depends on site conditions: The case of 2017 summer heat wave in southern Europe.

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.

Linkage of forest productivity to tree diversity under two different bioclimatic regimes in Italy.

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.

The sensitivity of the forest carbon budget shifts across processes along with stand development and climate change.

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.

Nitrogen deposition outweighs climatic variability in driving annual growth rate of canopy beech trees: Evidence from long-term growth reconstruction across a geographic gradient.

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.

Biochar As Plant Growth Promoter: Better Off Alone or Mixed with Organic Amendments?

Biochar is nowadays largely used as a soil amendment and is commercialized worldwide. However, in temperate agro-ecosystems the beneficial effect of biochar on crop productivity is limited, with several studies reporting negative crop responses. In this work, we studied the effect of 10 biochar and 9 not pyrogenic organic amendments (NPOA), using pure and in all possible combinations on lettuce growth (Lactuca sativa). Organic materials were characterized by 13C-CPMAS NMR spectroscopy and elemental analysis (pH, EC, C, N, C/N and H/C ratios). Pure biochars and NPOAs have variable effects, ranging from inhibition to strong stimulation on lettuce growth. For NPOAs, major inhibitory effects were found with N poor materials characterized by high C/N and H/C ratio. Among pure biochars, instead, those having a low H/C ratio seem to be the best for promoting plant growth. When biochars and organic amendments were mixed, non-additive interactions, either synergistic or antagonistic, were prevalent. However, the mixture effect on plant growth was mainly dependent on the chemical quality of NPOAs, while biochar chemistry played a secondary role. Synergisms were prevalent when N rich and lignin poor materials were mixed with biochar. On the contrary, antagonistic interactions occurred when leaf litter or woody materials were mixed with biochar. Further research is needed to identify the mechanisms behind the observed non-additive effects and to develop biochar-organic amendment combinations that maximize plant productivity in different agricultural systems.

Long-term temporal relationships between environmental conditions and xylem functional traits: a meta-analysis across a range of woody species along climatic and nitrogen deposition gradients.

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.

Interpreting the Climatic Effects on Xylem Functional Traits in Two Mediterranean Oak Species: The Role of Extreme Climatic Events.

In the Mediterranean region, the widely predicted rise in temperature, change in the precipitation pattern, and increase in the frequency of extreme climatic events are expected to alter the shape of ecological communities and to affect plant physiological processes that regulate ecosystem functioning. Although change in the mean values are important, there is increasing evidence that plant distribution, survival, and productivity respond to extremes rather than to the average climatic condition. The present study aims to assess the effects of both mean and extreme climatic conditions on radial growth and functional anatomical traits using long-term tree-ring time series of two co-existing Quercus spp. from a drought-prone site in Southern Italy. In particular, this is the first attempt to apply the Generalized Additive Model for Location, Scale, and Shape (GAMLSS) technique and Bayesian modeling procedures to xylem traits data set, with the aim of (i) detecting non-linear long-term responses to climate and (ii) exploring relationships between climate extreme and xylem traits variability in terms of probability of occurrence. This study demonstrates the usefulness of long-term xylem trait chronologies as records of environmental conditions at annual resolution. Statistical analyses revealed that most of the variability in tree-ring width and specific hydraulic conductivity might be explained by cambial age. Additionally, results highlighted appreciable relationships between xylem traits and climate variability more than tree-ring width, supporting also the evidence that the plant hydraulic traits are closely linked to local climate extremes rather than average climatic conditions. We reported that the probability of extreme departure in specific hydraulic conductivity (Ks) rises at extreme values of Standardized Precipitation Index (SPI). Therefore, changing frequency or intensity of extreme events might overcome the adaptive limits of vascular transport, resulting in substantial reduction of hydraulic functionality and, hence increased incidence of xylem dysfunctions.

Functional adjustments of xylem anatomy to climatic variability: insights from long-term Ilex aquifolium tree-ring series.

The present study assessed the effects of climatic conditions on radial growth and functional anatomical traits, including ring width, vessel size, vessel frequency and derived variables, i.e., potential hydraulic conductivity and xylem vulnerability to cavitation in Ilex aquifolium L. trees using long-term tree-ring time series obtained at two climatically contrasting sites, one mesic site in Switzerland (CH) and one drought-prone site in Italy (ITA). Relationships were explored by examining different xylem traits, and point pattern analysis was applied to investigate vessel clustering. We also used generalized additive models and bootstrap correlation functions to describe temperature and precipitation effects. Results indicated modified radial growth and xylem anatomy in trees over the last century; in particular, vessel frequency increased markedly at both sites in recent years, and all xylem traits examined, with the exception of xylem cavitation vulnerability, were higher at the CH mesic compared with the ITA drought site. A significant vessel clustering was observed at the ITA site, which could contribute to an enhanced tolerance to drought-induced embolism. Flat and negative relationships between vessel size and ring width were observed, suggesting carbon was not allocated to radial growth under conditions which favored stem water conduction. Finally, in most cases results indicated that climatic conditions influenced functional anatomical traits more substantially than tree radial growth, suggesting a crucial role of functional xylem anatomy in plant acclimation to future climatic conditions.