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Abstract As major terrestrial carbon sinks, forests play an important role in mitigating climate change. The relationship between the seasonal uptake of carbon and its allocation to woody biomass remains poorly understood, leaving a significant gap in our capacity to predict carbon sequestration by forests. Here, we compare the intra-annual dynamics of carbon fluxes and wood formation across the Northern hemisphere, from carbon assimilation and the formation of non-structural carbon compounds to their incorporation in woody tissues. We show temporally coupled seasonal peaks of carbon assimilation (GPP) and wood cell differentiation, while the two processes are substantially decoupled during off-peak periods. Peaks of cambial activity occur substantially earlier compared to GPP, suggesting the buffer role of non-structural carbohydrates between the processes of carbon assimilation and allocation to wood. Our findings suggest that high-resolution seasonal data of ecosystem carbon fluxes, wood formation and the associated physiological processes may reduce uncertainties in carbon source-sink relationships at different spatial scales, from stand to ecosystem levels.

ABSTRACT New approaches for managing agricultural and forestry systems are needed to bring back inputs to levels that are within planetary boundaries and make greater and better use of ecosystem services based on biodiversity. A new scientific framework informed by ecology, agronomy, forestry, and agroforestry is key to designing resilient plant‐based ecosystems to meet this challenge. Integrating information on plant functional traits, ontogenetic development stages, site characteristics, and structural stand characteristics can unleash the power of diversity (in species traits and structural and temporal arrangements) as a crucial factor for sustaining environmental services in times of global change. To leverage the benefits of diversity, a general theoretical framework and scalable simulation models are needed to understand structural and species diversification effects and interactions at multiple levels, from plant to field/forest stand to landscape. By working across established research boundaries, the scientific community can harness the power of structural and functional diversity to develop resilient, production‐oriented ecosystems. With this integrative approach, our objectives are as follows: (i) to conceptualize processes and methodologies for managing resilient terrestrial ecosystems that can guarantee sustainable and diversified ecosystem services within planetary boundaries, and (ii) to outline the workflow for crafting a system capable of sustaining human well‐being amid space, resource, and energy constraints.

Abstract Tree mortality and forest dieback episodes are increasing due to drought and heat stress. Nevertheless, a comprehensive understanding of mechanisms enabling trees to withstand and survive droughts remains lacking. Our study investigated basal area increment (BAI), and δ13C-derived intrinsic water-use-efficiency (iWUE), to elucidate beech resilience across four healthy stands in Italy with varying climates and soil water availability. Additionally, fist-order autocorrelation (AR1) analysis was performed to detect early warning signals for potential tree dieback risks during extreme drought events. Results reveal a negative link between BAI and vapour pressure deficit (VPD), especially in southern latitudes. After the 2003 drought, BAI decreased at the northern site, with an increase in δ13C and iWUE, indicating conservative water-use. Conversely, the southern sites showed increased BAI and iWUE, likely influenced by rising CO2 and improved water availability. In contrast, the central site sustained higher transpiration rates due to higher soil water holding capacity (SWHC). Despite varied responses, most sites exhibited reduced resilience to future extreme events, indicated by increased AR1. Temperature significantly affected beech iWUE and BAI in northern Italy, while VPD strongly influenced the southern latitudes. The observed increase in BAI and iWUE in southern regions might be attributed to an acclimation response.

Background: Understanding the role of species identity in interactions among individuals is crucial for assessing the productivity and stability of mixed forests over time. However, there is limited knowledge concerning the variation in competitive effect and response of different species along climatic gradients. In this study, we investigated the importance of climate, tree size, and competition on the growth of three tree species: spruce (Picea abies), fir (Abies alba), and beech (Fagus sylvatica), and examined their competitive response and effect along a climatic gradient. Methods: We selected 39 plots distributed across the European mountains with records of the position and growth of 5,759 individuals. For each target species, models relating tree growth to tree size, climate and competition were proposed. Competition was modelled using a neighbourhood competition index that considered the effects of inter- and intraspecific competition on target trees. Competitive responses and effects were related to climate. Likelihood methods and information theory were used to select the best model. Results: Our findings revealed that competition had a greater impact on target species growth than tree size or climate. Climate did influence the competitive effects of neighbouring species, but it did not affect the target species' response to competition. The strength of competitive effects varied along the gradient, contingent on the identity of the interacting species. When the target species exhibited an intermediate competitive effect relative to neighbouring species, both higher inter- than intraspecific competitive effects and competition reduction occurred along the gradient. Notably, species competitive effects were most pronounced when the target species’ growth was at its peak and weakest when growing conditions were far from their maximum. Conclusions: Climate modulates the effects of competition from neighbouring trees on the target tree and not the susceptibility of the target tree to competition. The modelling approach should be useful in future research to expand our knowledge of how competition modulates forest communities across environmental gradients.

Biodiversity monitoring represents a major challenge to supporting proper forest ecosystem management and biodiversity conservation. The latter is indeed shifting in recent years from single-species to multi-taxon approaches. However, multi-taxonomic studies are quite rare due to the effort required for performing field surveys. In this context, remote sensing is a powerful tool, continuously providing consistent and open access data at a different range of spatial and temporal scales. In particular, the Sentinel-2 (S2) mission has great potential to produce reliable proxies for biological diversity. In beech forests of two Italian National Parks, we sampled the beetle fauna, breeding birds, and epiphytic lichens. First, we calculated Shannon’s entropy and Simpson’s diversity. Then, to produce variables for biodiversity assessment, we exploited S2 data acquired in the 4 years 2017–2021. S2 images were used to construct spectral bands and photosynthetic indices time series, from which 91 harmonic metrics were derived. For each taxon and multi-taxon community, we assessed the correlation with S2 harmonic metrics, biodiversity indices, and forest structural variables. Then, to assess the potential of the harmonic metrics in predicting species diversity in terms of Shannon’s and Simpson’s biodiversity indices, we also fit a random forests model between each diversity index and the best 10 harmonic metrics (in terms of absolute correlation, that is, the magnitude of the correlation) for each taxon. The models’ performance was evaluated via the relative root mean squared error (RMSE%). Overall, 241 beetle, 27 bird, and 59 lichen species were recorded. The diversity indices were higher for the multi-taxon community than for the single taxa. They were generally higher in the CVDA site than in GSML, except for the bird community. The highest correlation values between S2 data and biodiversity indices were recorded in CVDA for multi-taxon and beetle communities (| r| = 0.52 and 0.38, respectively), and in GSML for lichen and beetle communities (| r| = 0.34 and 0.26, respectively). RMSE% ranged between 2.53 and 9.99, and between 8.1 and 16.8 for the Simpson and Shannon index, respectively. The most important variables are phase and RMSE of red-Edge bands for bird and lichen communities, while RMSE and time of tassel cap and from EVI indices for beetles and multi-taxon diversity. Our results demonstrate that S2 data can be used for identifying potential biodiversity hotspots, showing that the herein presented harmonic metrics are informative for several taxa inhabiting wood, giving concrete support to cost-effective biodiversity monitoring and nature-based forest management in complex mountain systems.

Background: Gradients in local environmental characteristics may favour the abundance of species with particular traits, while other species decline, or favour species with different traits at the same time, without an increase in average species abundances. Therefore, we asked: do variations in species and traits differ along gradients of deadwood variables? Do species abundance and trait occurrence change with species richness within or between functional groups? Thus, we analysed the beetle assemblages of five forest sites located in Italy, along the Apennines mountains. Methods: From 2012 to 2018 we sampled beetles and five deadwood types in 193 plots to characterise the deadwood gradient: standing dead trees, snags, dead downed trees, coarse woody debris, and stumps. We modelled beetle species relative abundances and trophic traits occurrences against the deadwood variables using joint species distribution models. Results: Out of 462 species, only 77 showed significant responses to at least one deadwood type, with a weak mean response across species. Trophic groups showed mostly negative responses to deadwood variables. Species abundance increased with species richness among sites only for phytophagous and saproxylophagous. Trait occurrence did not increase with species richness among sites, except for phytophagous and saproxylophagous. However, trait occurrence changed significantly with species richness of several trophic groups within some sites. We found that increases in species richness do not result in decreases in species abundance of a given trophic group, but rather null or positive relationships were found suggesting low interspecific competition. Conclusions: Our findings suggest that in Mediterranean mountain forests there is still room for increasing the level of naturalness, at least for what concerns deadwood management. On one side, our findings suggest that competition for deadwood substrates is still low, on the other side they indicate that increasing deadwood volume and types to improve overall beetle richness may increase also beetle abundances.

European beech (Fagus sylvatica L.) is a widespread and economically important temperate tree species in Europe. The warmer temperatures and severe drought events expected in the future, especially in Mediterranean areas, could affect the vitality and productivity of beech stands that have been intensively used in these areas in the past. Here, we aim to assess the wood anatomical responses of beech to environmental variability and silvicultural practices by investigating three beech stands along an elevational gradient (1,200 to 1,950 m a.s.l.) in the Apennines (Italy). Therefore, we quantified several anatomical traits of the xylem vessels related to tree hydraulics from five trees per stand and investigated variability between and within tree rings. Our results suggest generally limited trait plasticity, with higher plasticity of mean vessel lumen area and theoretical hydraulic conductivity, while maximum vessel size and mean hydraulic diameter were less plastic, likely because of the stronger determination by tree height. High-elevation trees were hydraulically more limited than trees at a mid and lower elevation as indicated by the more conservative anatomical configuration, i.e., comparatively smaller vessels and a 50% tighter trait coordination. Cessation of coppicing resulted in a hydraulically safer anatomy with comparatively smaller vessels at the most intensively used site (1,200 m), triggered by increased water demand due to an increase in canopy density, and thus, an increase in stand transpiration. Furthermore, maximum vessel size at the beginning showed different climate sensitivity compared to the rest of the tree ring, while intra-ring anatomical profiles showed little difference between normal and the 5 years with the highest and lowest mean temperature and precipitation. Overall, this study highlights the challenges to separate the externally induced medium- to longer-term responses from ontogenetically determined patterns. We, therefore, call for more comprehensive studies to further explore and verify the plasticity of wood anatomical traits in European beech in response to short- to long-term environmental fluctuations to gain a mechanistic understanding useful for sustainable forest ecosystems.

The relationships between structural complexity, deadwood abundance, microhabitat type and species-diversity indicators are excellent tools to monitor biodiversity in forest ecosystems.In spite of their importance, correlations between structural traits and Coleoptera communities in Mediterranean mountain forests have only rarely been investigated. Consequently, the magnitude and direction of the relationships between forest traits and biodiversity indicators remain poorly understood. In this study, we analyzed whether biodiversity indices of saproxylic and non-saproxylic beetle communities could be influenced by stand structure, microhabitat type, and deadwood abundance in two protected beech forests located in the central and southern Apennines (namely Gran Sasso e Monti della Laga National Park, GSML, and Cilento, Vallo di Diano e Alburni National Park, CVDA). Standard measurements of forest structural traits and quantitative assessment of tree microhabitats and deadwood were carried out. Adult beetles were collected using window flight traps and emergence traps on decaying deadwood. The two beech forests were different in terms of both beetle communities and structural traits. A two-block partial least squares analysis 2B-PLS highlighted differences in biodiversity indices and structural traits between the two forest ecosystems. In GSML, we observed that biodiversity indices were positively correlated with the volume of coarse woody debris and the presence fungal infections, clefts into the sapwood, and woodpecker cavities, while more dominant beetle communities were found under denser canopy cover. In CVDA, Coleoptera abundance was positively correlated with the basal area and crown broken microhabitats. Our results point toward the relevance of ecological attributes in tracking changes in beetle biodiversity in specific forest contexts. In these protected Mediterranean mountain beech stands, in which the main forest management strategies have the primary objective of biodiversity conservation, we suggest to progressively increase the structural diversity and canopy dynamics, as well as the volume of coarse woody debris.

In the Czech Republic, mountain watersheds are mostly forested with dominant Norway spruce (Picea abies) plantations. The aim of this paper is to analyse changes in radial growth and xylem anatomy of Norway spruce trees in the upper plain of the Jizera Mountains, related to changes in climate (air temperature, precipitation), air pollution and acid atmospheric deposition. Data of two neighbouring climate stations were used to detect trends in air temperatures and precipitation. At elevations of 745-1060 m a.s.l., the ring-width growth was significantly affected by mean annual temperature, while impacts of elevation and precipitation were not significant. In the period 1975-1995, the detected drop in tree radial growth (ca 60% of the normal period, prior to the peak of acid atmospheric deposition) corresponded to the increase in atmospheric SO2 concentrations and acid atmospheric deposition. The number of cells in tree rings decreased by ca 30-40% in comparison with the normal period, but the mean size of cells did not change significantly. In the last 20 years, increasing radial growth has been detected simultaneously with rising air temperature, and density of cells decreased by 30% in early wood, and by 10% in late wood, increasing the total number of cells in tree rings by ca 10% in comparison with the normal period. Integrated effects of climate and non-climate variables on the variation of tree radial growth in the Jizera Mountains reflected the legacy of acid atmospheric deposition in the forest ecosystem.

The dynamics of Pinus mugo krummholz during concomitant change in pastoral land use and climate in central Italy since the mid-20th century was investigated. Krummholz dynamics were detected using sequential aerial photography and fitted to a logistic regression model with elevation, grazing, proximity to beech forest, and proximity to krummholz as explanatory variables. Dendrochronological series were correlated with temperature and precipitation and fitted to a linear model. During this period krummholz doubled in extent and migrated 35–65 m upslope. Expansion was positively associated with krummholz proximity, residual pastoral grazing, and proximity to beech forest beyond 10 m and negatively associated with elevation and beech forest closer than 10 m. The logistic regression model forecasts krummholz migration by an additional 30 m upslope by 2060. During the 20th century, winter and spring minimum temperatures increased but did not result in increased radial stem growth of P. mugo. The combined evidence suggests that krummholz dynamics can be explained by the legacy of summer pastoralism and the dispersal limitations of P. mugo, rather than by climate change.

The application of Monteith’s models to estimate the gross primary production (GPP) of forest ecosystems is a common and efficient practice. A modified version of one of these parametric models, C-Fix, has been widely tested in Italy at daily to annual temporal scales. The current paper is a step forward demonstrating the capability of this model to reproduce also inter-annual forest GPP variations. To this aim, dendrochronological measurements collected in two study areas were utilized to validate the model. The first study area is a coastal Mediterranean pine forest and the second is a more temperate ecosystem dominated by the presence of European beech. The experiment was organized into the following main steps. Modified C-Fix was applied to simulate annual GPP over a 15-year period for both sites. Both the main driving factors used by the model and the estimated GPP were then analysed and assessed against the detrended tree-ring width series of the collected cores. The study yields indications on the factors most influential on GPP for the two forest ecosystems and supports the capability of the proposed modelling approach to reproduce relevant inter-annual GPP variations, which is useful for predicting the impacts of ongoing climatic changes.

Impacts of biochar application in combination with organic fertilizer, such as compost, are not fully understood. In this study, we tested the effects of biochar amendment, compost addition, and their combination on lettuce plants grown in a soil poor in nutrients; soil microbiological, chemical, and physical characteristics were analyzed, together with plant growth and physiology. An initial screening was also done to evaluate the effect of biochar and compost toxicity, using cress plants and earthworms. Results showed that compost amendment had clear and positive effects on plant growth and yield and on soil chemical characteristics. However, we demonstrated that also the biochar alone stimulated lettuce leaves number and total biomass, improving soil total nitrogen and phosphorus contents, as well as total carbon, and enhancing related microbial communities. Nevertheless, combining biochar and compost, no positive synergic and summative effects were observed. Our results thus demonstrate that in a soil poor in nutrients the biochar alone could be effectively used to enhance soil fertility and plant growth and biomass yield. However, we can speculate that the combination of compost and biochar may enhance and sustain soil biophysical and chemical characteristics and improve crop productivity over time.

On Mt. Etna (Italy), an enhanced Normalized Difference in Vegetation Index (NDVI) signature was detected in the summers of 2001 and 2002 along a distinct line where, in November 2002, a flank eruption subsequently occurred. These observations suggest that pre-eruptive volcanic activity may have enhanced photosynthesis along the future eruptive fissure. If a direct relation between NDVI and future volcanic eruptions could be established, it would provide a straightforward and low-cost method for early detection of upcoming eruptions. However, it is unclear if, or to what extent, the observed enhancement of NDVI can be attributed to volcanic activity prior to the subsequent eruption. We consequently aimed at determining whether an increase in ambient temperature or additional water availability owing to the rise of magma and degassing of water vapour prior to the eruption could have increased photosynthesis of Mt. Etna's trees. Using dendro-climatic analyses we quantified the sensitivity of tree ring widths to temperature and precipitation at high elevation stands on Mt. Etna. Our findings suggest that tree growth at high elevation on Mt. Etna is weakly influenced by climate, and that neither an increase in water availability nor an increase in temperature induced by pre-eruptive activity is a plausible mechanism for enhanced photosynthesis before the 2002/2003 flank eruption. Our findings thus imply that other, yet unknown, factors must be sought as causes of the pre-eruption enhancement of NDVI on Mt. Etna.

Competitive interactions are important predictors of tree growth. Spatial and temporal changes in resource availability, and variation in species and spatial patterning of trees alter competitive interactions, thus affecting tree growth and, hence, biomass. Competition indices are used to quantify the level of competition among trees. As these indices are normally computed only over small areas, where field measurements are done, it would be useful to have a tool to predict them over large areas. On this regard, remote sensing, and in particular light detection and ranging (lidar) data, could be the perfect tool. The objective of this study was to use lidar metrics to predict competition (on the basis of distance-dependent competition indices) of individual trees and to relate them with tree aboveground biomass (AGB). The selected study area was a mountain forest area located in the Italian Alps. The analyses focused on the two dominant species of the area: Silver fir (Abies alba Mill.) and Norway spruce (Picea abies (L.) H. Karst). The results showed that lidar metrics could be used to predict competition indices of individual trees (R2 above 0.66). Moreover, AGB decreased as competition increased, suggesting that variations in the availability of resources in the soil, and the ability of plants to withstand competition for light may influence the partitioning of biomass.

It is still unclear whether the exponential rise of atmospheric CO2 concentration has produced a fertilization effect on tropical forests, thus incrementing their growth rate, in the last two centuries. As many factors affect tree growth patterns, short -term studies might be influenced by the confounding effect of several interacting environmental variables on plant growth. Long-term analyses of tree growth can elucidate long-term trends of plant growth response to dominant drivers. The study of annual rings, applied to long tree-ring chronologies in tropical forest trees enables such analysis. Long-term tree-ring chronologies of three widespread African species were measured in Central Africa to analyze the growth of trees over the last two centuries. Growth trends were correlated to changes in global atmospheric CO2 concentration and local variations in the main climatic drivers, temperature and rainfall. Our results provided no evidence for a fertilization effect of CO2 on tree growth. On the contrary, an overall growth decline was observed for all three species in the last century, which appears to be significantly correlated to the increase in local temperature. These findings provide additional support to the global observations of a slowing down of C sequestration in the trunks of forest trees in recent decades. Data indicate that the CO2 increase alone has not been sufficient to obtain a tree growth increase in tropical trees. The effect of other changing environmental factors, like temperature, may have overridden the fertilization effect of CO2.

Changes in intrinsic water use efficiency (iWUE) were investigated in Fagus sylvatica and Nothofagus spp. over the last century. We combined dendrochronological methods with dual-isotope analysis to investigate whether atmospheric changes enhanced iWUE of Fagus and Nothofagus and tree growth (basal area increment, BAI) along latitudinal gradients in Italy and Chile. Post-maturation phases of the trees presented different patterns in δ13C, Δ13C, δ18O, Ci (internal CO2 concentration), iWUE, and BAI. A continuous enhancement in isotope-derived iWUE was observed throughout the twentieth century, which was common to all sites and related to changes in Ca (ambient CO2 concentration) and secondarily to increases in temperature. In contrast to other studies, we observed a general increasing trend of BAI, with the exception of F. sylvatica in Aspromonte. Both iWUE and BAI were uncoupled with the estimated drought index, which is in agreement with the absence of enduring decline in tree growth. In general, δ13C and δ18O showed a weak relationship, suggesting the major influence of photosynthetic rate on Ci and δ13C, and the minor contribution of the regulation of stomatal conductance to iWUE. The substantial warming observed during the twentieth century did not result in a clear pattern of increased drought stress along these latitudinal transects, because of the variability in temporal trends of precipitation and in specific responses of populations.

Olive trees are a classic component of Mediterranean environments and some of them are known historically to be very old. In order to evaluate the possibility to use olive tree-rings for dendrochronology, we examined by various methods the reliability of olive tree-rings identification. Dendrochronological analyses of olive trees growing on the Aegean island Santorini (Greece) show that the determination of the number of tree-rings is impossible because of intra-annual wood density fluctuations, variability in tree-ring boundary structure, and restriction of its cambial activity to shifting sectors of the circumference, causing the tree-ring sequences along radii of the same cross section to differ.

The estimation of site water budget is important in Mediterranean areas, where it represents a crucial factor affecting the quantity and quality of traditional crop production. This is particularly the case for spatially fragmented, multi-layer agricultural ecosystems such as olive groves, which are traditional cultivations of the Mediterranean basin. The current paper aims at demonstrating the effectiveness of spatialized meteorological data and remote sensing techniques to estimate the actual evapotranspiration (ETA) and the soil water content (SWC) of an olive orchard in Central Italy. The relatively small size of this orchard (about 0.1 ha) and its two-layer structure (i.e., olive trees and grasses) require the integration of remotely sensed data with different spatial and temporal resolutions (Terra-MODIS, Landsat 8-OLI and Ikonos). These data are used to drive a recently proposed water balance method (NDVI-Cws) and predict ETA and then site SWC, which are assessed through comparison with sap flow and soil wetness measurements taken in 2013. The results obtained indicate the importance of integrating satellite imageries having different spatio-temporal properties in order to properly characterize the examined olive orchard. More generally, the experimental evidences support the possibility of using widely available remotely sensed and ancillary datasets for the operational estimation of ETA and SWC in olive tree cultivation systems.

Plants can remove or immobilize various environmental contaminants; however, little is known about the physiological mechanisms underlying responses to soil amendment with biosolids contaminated with heavy metals. We investigated the responses of cuttings of hybrid poplar clones Eridano and I-214 grown for a season in soil amended with nutrient-rich organic material from tanneries, which contains potentially toxic amounts of heavy elements. Plant growth traits, gas exchange parameters, stomatal density and leaf layer thickness of frozen-hydrated leaves, and foliar concentrations of heavy metals and nitrogen were determined. Overall, soil amendment increased net assimilation rate and growth, but the cuttings accumulated only small quantities of heavy metal soil contaminants. [ABSTRACT FROM AUTHOR]