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Aim: The microclimate and light conditions on the forest floor are strongly modified by tree canopies. Therefore, we need to better consider the micro‐environment when quantifying trait–environment relationships for forest understorey plants. Here, we quantify relationships between micro‐environmental conditions and plant functional traits at the community level, including intraspecific trait variation, and their relationship with microclimate air temperature, light and soil properties. Location: Deciduous temperate forests across Europe. Time period: 2018. Major taxa studied: Herbaceous vegetation. Methods: We sampled 225 plots across 15 regions along four complementary gradients capturing both macro‐ and microclimatic conditions including latitude, elevation, forest management and distance to forest edges. We related the community‐weighted mean of five plant functional traits (plant height, specific leaf area [SLA], plant carbon [C], plant nitrogen [N] and plant C:N ratio) across 150 vascular plant species to variation in local microclimate air temperature, light and soil properties. We tested the effect of accounting for intraspecific variation in trait–environment relationships and performed variation partitioning to identify major drivers of trait variation. Results: Microclimate temperature, light availability and soil properties were all important predictors of community‐weighted mean functional traits. When light availability and variation in temperature were higher, the herb community often consisted of taller plants with a higher C:N ratio. In more productive environments (e.g. with high soil nitrogen availability), the community was dominated by individuals with resource‐acquisitive traits: high SLA and N but low C:N. Including intraspecific trait variation increased the strength of the trait–micro‐environment relationship, and increased the importance of light availability. Main conclusions: The trait–environment relationships were much stronger when the micro‐environment and intraspecific trait variation were considered. By locally steering light availability and temperature, forest managers can potentially impact the functional signature of the forest herb‐layer community. [ABSTRACT FROM AUTHOR]

Tree diversity and species identity are known to play an important role in modulating forest productivity and its capacity to buffer the effects of climatic events. The FunDivEurope exploratory platform allowed us to analyse this modulating effect in a medium‐term time span, after an abrupt rise to a new stable level of the average summer temperature of ca 2°C, in Mediterranean forests in central Italy. This paper aims to answer the following questions: 1) did increasing temperature and drought events affect the growth of thermophilous deciduous forests? 2) Was this effect buffered in mixed stands compared to monocultures? 3) Did co‐occurring tree species with different ecological characteristics, from more mesophilous to more xerophilous, have different responses? In 2012 and 2017, wood cores were collected from 659 trees in 36 plots representative of thermophilous deciduous forests. The selected tree species were Castanea sativa, Ostrya carpinifolia, Quercus cerris, Quercus ilex and Quercus petraea. In the sampling plots, they were present in pure stands and mixtures from two to four species. After measuring annual rings on cores, chronologies of basal area increment were built, and inventory data were used to estimate tree growth. Results showed a strong reduction of growth, lasting at least 18 years, after the temperature rise. Tree diversity significantly reduced the growth drop after the sudden and stable rise in summer average temperature. Tree mixture effect on growth stability appeared to be dependent on the tree species present in the mixture. Temperature rise and associated drought events, even without changes in rainfall, are one of the main challenges that European forests will face in the current scenarios of climate change. Tree diversity can buffer the effects of climate change over periods of at least 15 years and should be considered in forest management plans. [ABSTRACT FROM AUTHOR]

Aim: The amount of forest edges is increasing globally due to forest fragmentation and land‐use changes. However, edge effects on the soil seed bank of temperate forests are still poorly understood. Here, we assessed edge effects at contrasting spatial scales across Europe and quantified the extent to which edges can preserve the seeds of forest specialist plants. Location: Temperate European deciduous forests along a 2,300‐km latitudinal gradient. Time period: 2018–2021. Major taxa studied: Vascular plants. Methods: Through a greenhouse germination experiment, we studied how edge effects alter the density, diversity, composition and functionality of forest soil seed banks in 90 plots along different latitudes, elevations and forest management types. We also assessed which environmental conditions drive the seed bank responses at the forest edge versus interior and looked at the relationship between the seed bank and the herb layer species richness. Results: Overall, 10,108 seedlings of 250 species emerged from the soil seed bank. Seed density and species richness of generalists (species not only associated with forests) were higher at edges compared to interiors, with a negative influence of C : N ratio and litter quality. Conversely, forest specialist species richness did not decline from the interior to the edge. Also, edges were compositionally, but not functionally, different from interiors. The correlation between the seed bank and the herb layer species richness was positive and affected by microclimate. Main conclusions Our results underpin how edge effects shape species diversity and composition of soil seed banks in ancient forests, especially increasing the proportion of generalist species and thus potentially favouring a shift in community composition. However, the presence of many forest specialists suggests that soil seed banks still play a key role in understorey species persistence and could support the resilience of our fragmented forests. [ABSTRACT FROM AUTHOR]

Summary: Forests harbour large spatiotemporal heterogeneity in canopy structure. This variation drives the microclimate and light availability at the forest floor. So far, we do not know how light availability and sub‐canopy temperature interactively mediate the impact of macroclimate warming on understorey communities.We therefore assessed the functional response of understorey plant communities to warming and light addition in a full factorial experiment installed in temperate deciduous forests across Europe along natural microclimate, light and macroclimate gradients. Furthermore, we related these functional responses to the species' life‐history syndromes and thermal niches.We found no significant community responses to the warming treatment. The light treatment, however, had a stronger impact on communities, mainly due to responses by fast‐colonizing generalists and not by slow‐colonizing forest specialists. The forest structure strongly mediated the response to light addition and also had a clear impact on functional traits and total plant cover.The effects of short‐term experimental warming were small and suggest a time‐lag in the response of understorey species to climate change. Canopy disturbance, for instance due to drought, pests or logging, has a strong and immediate impact and particularly favours generalists in the understorey in structurally complex forests. [ABSTRACT FROM AUTHOR]

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Aim: Variation in plant defence traits has been frequently assessed along large‐scale macroclimatic clines. In contrast, local‐scale changes in the environment have recently been proposed to also modulate plant defence traits. Yet, the relative importance of drivers at both scales has never been tested. We aimed to quantify the relative importance of environmental drivers inherent to large and small spatial scales on the physical and chemical defence and tolerance to herbivory in understorey plant species of deciduous forests of Europe. Location: Deciduous forests in Europe. Time period: Present. Major taxa studied: Forest understorey plants. Methods: We sampled four typical ancient forest herbs (Anemone nemorosa, Oxalis acetosella, Deschampsia cespitosa, Milium effusum) along small and large spatial scale gradients (those driven by latitude, elevation, forest management and distance to the forest edge), and analysed a suite of nine constitutively expressed traits associated with overall resistance to herbivory, and their multivariate response to environmental clines. Results: Although our study included a large gradient in macroclimate, we found variation in the local environment at small spatial scales (i.e. soil nutrient concentration and forest structural complexity) to be more important in predicting plant resistance to herbivory. Main conclusions: In addition to macroclimatic conditions, subtle differences in forest microclimate and soil characteristics also played a major role in modulating plant defence phenotypes. These findings highlight the importance of the local habitat structure and environmental conditions in modulating plant resistance to herbivory. [ABSTRACT FROM AUTHOR]

Questions: Does the influence of forest edges on plant species richness and composition depend on forest management? Do forest specialists and generalists show contrasting patterns? Location: Mesic, deciduous forests across Europe. Methods: Vegetation surveys were performed in forests with three management types (unthinned, thinned 5–10 years ago and recently thinned) along a macroclimatic gradient from Italy to Norway. In each of 45 forests, we established five vegetation plots along a south‐facing edge‐to‐interior gradient (n = 225). Forest specialist, generalist and total species richness, as well as evenness and proportion of specialists, were tested as a function of the management type and distance to the edge while accounting for several environmental variables (e.g. landscape composition and soil characteristics). Magnitude and distance of edge influence were estimated for species richness per management type. Results: Greatest total species richness was found in thinned forests. Edge influence on generalist plant species richness was contingent on the management type, with the smallest decrease in species richness from the edge‐to‐interior in unthinned forests. In addition, generalist richness increased with the proportion of forests in the surrounding landscape and decreased in forests dominated by tree species that cast more shade. Forest specialist species richness, however, was not affected by management type or distance to the edge, and only increased with pH and increasing proportion of forests in the landscape. Conclusions: Forest thinning affects the plant community composition along edge‐to‐interior transects of European forests, with richness of forest specialists and generalists responding differently. Therefore, future studies should take the forest management into account when interpreting edge‐to‐interior because both modify the microclimate, soil processes and deposition of polluting aerosols. This interaction is key to predict the effects of global change on forest plants in landscapes characterized by the mosaic of forest patches and agricultural land that is typical for Europe. [ABSTRACT FROM AUTHOR]

• An extreme drought event affected Italian thermophilous deciduous forests in 2017. • In the short term, this reduced understorey cover by 50% and species richness. • Evenness and diversity increased in the monospecific plots but not in the mixtures. • Drought increased non-forest specialists and non-native species in monocultures. • Management should favour mixtures to buffer understorey changes soon after drought. Understorey vegetation is a key component of plant biodiversity in forests. Events of extreme drought are a potential major driver of change of this layer. Although these are predicted to increase in frequency and duration due to climate change, their impacts on understorey communities are still unknown. In this paper, we re-surveyed a set of 36 plots in mature thermophilous deciduous forests eight months after the end of an extremely dry period in 2016–2017. The plots are part of a European network representing a gradient of overstorey richness from 1- to 5-tree species mixtures, with variable composition. We took advantage of this model system to test the hypothesis that species richness and identity of dominant trees in the overstorey may modulate the impact of drought on understorey cover, diversity and composition. In spring 2018, a ca. 50% reduction of understorey cover occurred in both mono-specific and mixed plots. Species evenness and Shannon α-diversity increased in the monospecific plots, due to a reduction of abundance of tree species at juvenile stage and the appearance of non-forest specialists that did not occur in the mixtures. Overstorey species identity effects on understorey diversity were negligible. Overall compositional changes were moderate, though larger in the mixed plots and in those with the less drought-adapted tree species in the overstorey. Ellenberg ecological profile of the community did not change significantly but light values tended to increase in the monospecific plots, contrary to the mixtures. Overall, our findings provide circumstantial support to a higher diversity stability in the understorey of mixed plots shortly after drought, despite compositional variations. More studies and monitoring programs are needed to assess understorey recovery capacity and resilience after such events in the long-term. [ABSTRACT FROM AUTHOR]

Forest fragmentation increases the proportion of edge area and this, in turn, induces changes in forest structure, species composition and microclimate. These factors are also strongly determined by the forest management regime. Although the interactive effects of edges and density on forest plant communities have been extensively studied, little is known about the response of the belowground communities. Here we investigated the variation of soil microbiota in 45 deciduous broadleaved forests along a latitudinal gradient from Italy to Norway at a continental scale across Europe. Phospholipid fatty acid (PLFA) and neutral lipid fatty acid (NLFA) were used to map the microbial community in the forest edge and interior across three forest densities (dense, intermediate, open forest). Microbial community composition was only affected by forest edge effects and not by forest density. We did not find any interaction effects between forest density and distance-to-edge. Arbuscular mycorrhizal fungi (AMF) were significantly more abundant in edges and Gram-negative bacteria more abundant in interiors, respectively. The microbial community composition was closely related to soil pH, soil potassium and nitrogen, texture (percent sand) and soil temperature. Soil pH was positively correlated with the saprotrophic fungi and potassium was positively correlated with Gram-negative bacteria but negatively correlated with Actinobacteria. In sum, we reveal the notable effects of forest edges on the soil AMF abundance. This result indicated that AMF could possess a stronger affinity with species growing in the edges, which may help to improve plant performance under hostile conditions herein. [ABSTRACT FROM AUTHOR]

• We quantified evaporation and soil and air temperature offsets in forest edges across Europe. • Roughly 10% of European broadleaved forests are affected by altered temperature regimes. • Macroclimate and management affected edge-to-interior temperature offset gradients. • Forest structure and the forest-floor biomass are important drivers of temperature offsets. • Dense forest edges can help mitigate edge influences and protect forest interior microclimates. Global forest cover is heavily fragmented. Due to high edge-to-surface ratios in small forest patches, a large proportion of forests is affected by edge influences involving steep microclimatic gradients. Although forest edges are important ecotones and account for 20% of the global forested area, it remains unclear how biotic and abiotic drivers affect forest edge microclimates at the continental scale. Here we report soil and air temperatures measured in 225 deciduous forest plots across Europe for two years. Forest stands were situated along a latitudinal gradient and subject to a varying vegetation structure as quantified by terrestrial laser scanning. In summer, the average offset of air and soil temperatures in forest edges compared to temperatures outside the forest amounted to -2.8 °C and -2.3 °C, respectively. Edge-to-interior summer temperature gradients were affected by the macroclimate and edge structure. From the edge onwards, larger offsets were observed in dense forest edges and in warmer, southern regions. In open forests and northern Europe, altered microclimatic conditions extended deeper into the forest and gradients were steeper. Canopy closure and plant area index were important drivers of summer offsets in edges, whereas in winter also the forest-floor biomass played a key role. Using high-resolution maps, we estimated that approximately 10% of the European broadleaved forests would be affected by altered temperature regimes. Gradual transition zones between forest and adjacent lands are valuable habitat types for edge species. However, if cool and moist forest interiors are desired, then (i) dense and complex forest edges, (ii) an undisturbed forested buffer zone of at least 12.5 m deep and (iii) trees with a high shade casting ability could all contribute to an increased offset. These findings provide important guidelines to mitigate edge influences, to protect typical forest microclimates and to adapt forest management to climate change. [ABSTRACT FROM AUTHOR]