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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

Forests worldwide are experiencing fragmentation, with especially important consequences for ecosystems bordering urbanized areas. Urban forests are exposed to local warming due to the urban heat island which affects their biodiversity and ecosystem functioning. A key ecosystem function affecting carbon and nutrient cycling in forests is litter decomposition, a process driven by the local microclimate. Thus, our aim was to clarify the impact of the urban heat island on litter decomposition in urban forests. We studied soil microclimate and litter decomposition in six urban forests across Europe and along local gradients from the urban forest edge to the interior. To quantify decomposition independent from local forest composition and litter quality, we used standardized green tea and rooibos tea litterbags. We determined the role of the soil microclimate and other environmental drivers for litter decomposition. Secondly, we assessed effects of edge proximity and landscape context on the soil microclimate. Soil characteristics were only driving green tea and not rooibos tea decomposition. On the contrary, higher soil temperatures resulted in faster rates of litter decomposition for both green and rooibos tea and were related to the proximity to the forest edge and the proportion of built-up area in the landscape. Via structural equation modelling we detected cascading effects of the urban heat island on litter decomposition. Such changes in litter decomposition have the potential to alter the soil food web, nutrient cycling and carbon drawdown in urban forests, and could result in significant interactions between urbanisation and ongoing climate change.

Forest biodiversity and ecosystem services are hitherto predominantly quantified in forest interiors, well away from edges. However, these edges also represent a substantial proportion of the global forest cover. Here we quantified plant biodiversity and ecosystem service indicators in 225 plots along forest edge-to-interior transects across Europe. We found strong trade-offs: phylogenetic diversity (evolutionary measure of biodiversity), proportion of forest specialists, decomposition and heatwave buffering increased towards the interior, whereas species richness, nectar production potential, stemwood biomass and tree regeneration decreased. These trade-offs were mainly driven by edge-to-interior structural differences. As fragmentation continues, recognizing the role of forest edges is crucial for integrating biodiversity and ecosystem service considerations into sustainable forest management and policy.

Macroclimatic changes are impacting ecosystems worldwide. However, a large portion of terrestrial species live under conditions where impacts of macroclimate change are buffered, such as in the shade of trees, and how this buffering impacts future below-canopy biodiversity redistributions at the continental scale is unknown. Here we show that shady forest floors due to dense tree canopies mitigate severe warming impacts on forest biodiversity, while canopy opening amplifies macroclimate change impacts. A cross-continental transplant experiment in five contrasting biogeographical areas combined with experimental heating and irradiation treatments was used to parametize 25-m resolution mechanistic demographic distribution models and project the current and future distributions of 12 common understorey plant species, considering the effects of forest microclimate and forest cover density. These results highlight microclimates and forest density as powerful tools for forest managers and policymakers to shelter forest biodiversity from climate change. The impacts of microclimate on future plant population dynamics are poorly understood. The authors use large-scale transplant climate change experiments to show the contribution of forest microclimates to population dynamics and project the distributions of 12 common understorey plants.

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.

Context Plant populations in agricultural landscapes are mostly fragmented and their functional connectivity often depends on seed and pollen dispersal by animals. However, little is known about how the interactions of seed and pollen dispersers with the agricultural matrix translate into gene flow among plant populations. Objectives We aimed to identify effects of the landscape structure on the genetic diversity within, and the genetic differentiation among, spatially isolated populations of three temperate forest herbs. We asked, whether different arable crops have different effects, and whether the orientation of linear landscape elements relative to the gene dispersal direction matters. Methods We analysed the species' population genetic structures in seven agricultural landscapes across temperate Europe using microsatellite markers. These were modelled as a function of landscape composition and configuration, which we quantified in buffer zones around, and in rectangular landscape strips between, plant populations. Results Landscape effects were diverse and often contrasting between species, reflecting their association with different pollen- or seed dispersal vectors. Differentiating crop types rather than lumping them together yielded higher proportions of explained variation. Some linear landscape elements had both a channelling and hampering effect on gene flow, depending on their orientation. Conclusions Landscape structure is a more important determinant of the species' population genetic structure than habitat loss and fragmentation per se. Landscape planning with the aim to enhance the functional connectivity among spatially isolated plant populations should consider that even species of the same ecological guild might show distinct responses to the landscape structure.

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.

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.

Aims: Understanding fine-grain diversity patterns across large spatial extents is fundamental for macroecological research and biodiversity conservation. Using the GrassPlot database, we provide benchmarks of fine-grain richness values of Palaearctic open habitats for vascular plants, bryophytes, lichens and complete vegetation (i.e., the sum of the former three groups). Location: Palaearctic biogeographic realm. Methods: We used 126,524 plots of eight standard grain sizes from the GrassPlot database: 0.0001, 0.001, 0.01, 0.1, 1, 10, 100 and 1,000 m(2) and calculated the mean richness and standard deviations, as well as maximum, minimum, median, and first and third quartiles for each combination of grain size, taxonomic group, biome, region, vegetation type and phytosociological class. Results: Patterns of plant diversity in vegetation types and biomes differ across grain sizes and taxonomic groups. Overall, secondary (mostly semi-natural) grasslands and natural grasslands are the richest vegetation type. The open-access file GrassPlot Diversity Benchmarks and the web tool GrassPlot Diversity Explorer are now available online () and provide more insights into species richness patterns in the Palaearctic open habitats. Conclusions: The GrassPlot Diversity Benchmarks provide high-quality data on species richness in open habitat types across the Palaearctic. These benchmark data can be used in vegetation ecology, macroecology, biodiversity conservation and data quality checking. While the amount of data in the underlying GrassPlot database and their spatial coverage are smaller than in other extensive vegetation-plot databases, species recordings in GrassPlot are on average more complete, making it a valuable complementary data source in macroecology.

Context Evidence for effects of habitat loss and fragmentation on the viability of temperate forest herb populations in agricultural landscapes is so far based on population genetic studies of single species in single landscapes. However, forest herbs differ in their life histories, and landscapes have different environments, structures and histories, making generalizations difficult. Objectives We compare the response of three slow-colonizing forest herbs to habitat loss and fragmentation and set this in relation to differences in life-history traits, in particular their mating system and associated pollinators. Methods We analysed the herbs' landscape-scale population genetic structure based on microsatellite markers from replicate forest fragments across seven European agricultural landscapes. Results All species responded to reductions in population size with a decrease in allelic richness and an increase in genetic differentiation among populations. Genetic differentiation also increased with enhanced spatial isolation. In addition, each species showed unique responses. Heterozygosity in the self-compatible Oxalis acetosella was reduced in smaller populations. The genetic diversity of Anemone nemorosa, whose main pollinators are less mobile, decreased with increasing spatial isolation, but not that of the bumblebee-pollinated Polygonatum multiflorum. Conclusions Our study indicates that habitat loss and fragmentation compromise the long-term viability of slow-colonizing forest herbs despite their ability to persist for many decades by clonal propagation. The distinct responses of the three species studied within the same landscapes confirm the need of multi-species approaches. The mobility of associated pollinators should be considered an important determinant of forest herbs' sensitivity to habitat loss and fragmentation.

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 degrees C and -2.3 degrees 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 clima

Forest biodiversity world-wide is affected by climate change, habitat loss and fragmentation, and today 20% of the forest area is located within 100 m of a forest edge. Still, forest edges harbour a substantial amount of terrestrial biodiversity, especially in the understorey. The functional and phylogenetic diversity of forest edges have never been studied simultaneously at a continental scale, in spite of their importance for the forests' functioning and for communities' resilience to future change. We assessed nine metrics of taxonomic, phylogenetic and functional diversity of understorey plant communities in 225 plots spread along edge-to-interior gradients in deciduous forests across Europe. We then derived the relative effects and importance of edaphic, stand and landscape conditions on the diversity metrics. Here, we show that taxonomic, phylogenetic and functional diversity metrics respond differently to environmental conditions. We report an increase in functional diversity in plots with stronger microclimatic buffering, in spite of their lower taxonomic species richness. Additionally, we found increased taxonomic species richness at the forest edge, but in forests with intermediate and high openness, these communities had decreased phylogenetic diversity. Functional and phylogenetic diversity revealed complementary and important insights in community assembly mechanisms. Several environmental filters were identified as potential drivers of the patterns, such as a colder macroclimate and less buffered microclimate for functional diversity. For phylogenetic diversity, edaphic conditions were more important. Interestingly, plots with lower soil pH had decreased taxonomic species richness, but led to increased phylogenetic diversity, challenging the phylogenetic niche conservatism concept. Synthesis. Taxonomic, phylogenetic and functional diversity of understorey communities in forest edges respond differently to environmental conditions, providing insight int

Forests play a key role in global carbon cycling and sequestration. However, the potential for carbon drawdown is affected by forest fragmentation and resulting changes in microclimate, nutrient inputs, disturbance and productivity near edges. Up to 20% of the global forested area lieswithin 100 m of an edge and, even in temperate forests, knowledge on howedge conditions affect carbon stocks and howfar this influence penetrates into forest interiors is scarce. Here we studied carbon stocks in the aboveground biomass, forest floor and the mineral topsoil in 225 plots in deciduous forest edges across Europe and tested the impact of macroclimate, nitrogen deposition and smaller-grained drivers (e.g. microclimate) on these stocks. Total carbon and carbon in the aboveground biomass stock were on average 39% and 95% higher at the forest edge than 100 m into the interior. The increase in the aboveground biomass stock close to the edgewas mainly related to enhanced nitrogen deposition. No edge influence was found for stocks in the mineral topsoil. Edge-to-interior gradients in forest floor carbon changed across latitude: carbon stocks in the forest floor were higher near the edge in southern Europe. Forest floor carbon decreased with increasing litter quality (i.e. high decomposition rate) and decreasing plant area index, whereas higher soil temperatures negatively affected the mineral top soil carbon. Based on high-resolution forest fragmentation maps, we estimate that the additional carbon stored in deciduous forest edges across Europe amounts to not less than 183 Tg carbon, which is equivalent to the storage capacity of 1 million ha of additional forest. This study underpins the importance of including edge influences when quantifying the carbon stocks in temperate forests and stresses the importance of preserving natural forest edges and small forest patches with a high edge-to-interior surface area.

Linear landscape elements such as hedgerows and road verges have the potential to mitigate the adverse effects of habitat fragmentation and climate change on species, for instance, by serving as a refuge habitat or by improving functional connectivity across the landscape. However, so far this hypothesis has not been evaluated at large spatial scales, preventing us from making generalized conclusions about their efficacy and implementation in conservation policies. Here, we assessed plant diversity patterns in 336 vegetation plots distributed along hedgerows and road verges, spanning a macro-environmental gradient across temperate Europe. We compared herb-layer species richness and composition in these linear elements with the respective seed-source (core) habitats, that is, semi-natural forests and grasslands. Next, we assessed how these differences related to several environmental drivers acting either locally, at the landscape level or along the studied macro-ecological gradient. Across all regions, about 55% of the plant species were shared between forests and hedgerows, and 52% between grasslands and road verges. Habitat-specialist richness was 11% lower in the linear habitats than in the core habitats, while generalist richness was 14% higher. The difference in floristic composition between both habitat types was mainly due to species turnover, and not nestedness. Most notably, forest-specialist richness in hedgerows responded positively to tree cover, tree height and the proportion of forests in the surrounding landscape, while generalist richness was negatively affected by tree height and buffering effect of trees on subcanopy temperatures. Grassland and road verge diversity was mainly influenced by soil properties, with positive effects of basic cation levels on the number of specialists and those of bioavailable soil phosphorus on generalist diversity. Synthesis and applications. We demonstrate that linear landscape elements provide a potential habitat for

Hedgerows have the potential to facilitate the persistence and migration of species across landscapes, mostly due to benign microclimatic conditions. This thermal buffering function may become even more important in the future for species migration under climate change. Unfortunately, there is a lack of empirical studies quantifying the microclimate of hedgerows, particularly at broad geographical scales. Here we monitored sub-canopy temperatures using 168 miniature temperature sensors distributed along woodland-hedgerow transects, and spanning a 1600-km macroclimatic gradient across Europe. First, we assessed the variation in the temperature offset (that is, the difference between sub-canopy and corresponding macroclimate temperatures) for minimum, mean and maximum temperatures along the woodland-hedgerow transects. Next, we linked the observed patterns to macroclimate temperatures as well as canopy structure, overstorey composition and hedgerow characteristics. The sub-canopy versus macroclimate temperature offset was on average 0.10 degrees C lower in hedgerows than in woodlands. Minimum winter temperatures were consistently lower by 0.10 degrees C in hedgerows than in woodlands, while maximum summer temperatures were 0.80 degrees C higher, albeit mainly around the woodland-hedgerow ecotone. The temperature offset was often negatively correlated with macroclimate temperatures. The slope of this relationship was lower for maximum temperatures in hedgerows than in woodlands. During summer, canopy cover, tree height and hedgerow width had strong cooling effects on maximum mid-day temperatures in hedgerows. The effects of shrub height, shrub cover and shade-casting ability, however, were not significant. To our knowledge, this is the first study to quantify hedgerow microclimates along a continental-scale environmental gradient. We show that hedgerows are less efficient thermal insulators than woodlands, especially at high ambient temperatures (e.g. on warm summer da

Forest edges are interfaces between forest interiors and adjacent land cover types. They are important elements in the landscape with almost 20% of the global forest area located within 100 m of the edge. Edges are structurally different from forest interiors, which results in unique edge influences on microclimate, functioning and biodiversity. These edge influences have been studied for multiple decades, yet there is only limited information available on how forest edge structure varies at the continental scale, and which factors drive this potential structural diversity. Here we quantified the structural variation along 45 edge-to-interior transects situated along latitudinal, elevational and management gradients across Europe. We combined state-of-the-art terrestrial laser scanning and conventional forest inventory techniques to investigate how the forest edge structure (e.g. plant area index, stem density, canopy height and foliage height diversity) varies and which factors affect this forest edge structural variability. Macroclimate, management, distance to the forest edge and tree community composition all influenced the forest edge structural variability and interestingly we detected interactive effects of our predictors as well. We found more abrupt edge-to-interior gradients (i.e. steeper slopes) in the plant area index in regularly thinned forests. In addition, latitude, mean annual temperature and humidity all affected edge-to-interior gradients in stem density. We also detected a simultaneous impact of both humidity and management, and humidity and distance to the forest edge, on the canopy height and foliage height diversity. These results contribute to our understanding of how environmental conditions and management shape the forest edge structure. Our findings stress the need for site-specific recommendations on forest edge management instead of generalized recommendations as the macroclimate substantially influences the forest edge structure. Only t

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 environmental changes are expected to alter the functional characteristics of understorey herb-layer communities, potentially affecting forest ecosystem functioning. However, little is known about what drives the variability of functional traits in forest understories. Here, we assessed the role of different environmental drivers in shaping the functional trait distribution of understorey herbs in fragmented forests across three spatial scales. We focused on 708 small, deciduous forest patches located in 16 agricultural landscape windows, spanning a 2500-km macroclimatic gradient across the temperate forest biome in Europe. We estimated the relative effect of patch-scale, landscape-scale and macroclimatic variables on the community mean and variation of plant height, specific leaf area and seed mass. Macroclimatic variables (monthly temperature and precipitation extremes) explained the largest proportion of variation in community trait means (on average 77% of the explained variation). In contrast, patch-scale factors dominated in explaining community trait variation (on average 68% of the explained variation). Notably, patch age, size and internal heterogeneity had a positive effect on the community-level variability. Landscape-scale variables explained only a minor part of the variation in both trait distribution properties. The variation explained by shared combinations of the variable groups was generally negligible. These findings highlight the importance of considering multiple spatial scales in predictions of environmental-change effects on the functionality of forest understories. We propose that forest management sustainability could benefit from conserving larger, historically continuous and internally heterogeneous forest patches to maximise ecosystem service diversity in rural landscapes.