Search

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

Plant phenology, i. e. the timing of life cycle events, is related to individual fitness and species distribution ranges. Temperature is one of the most important drivers of plant phenology together with day length. The adaptation of their phenology may be important for the success of invasive plant species. The present study aims at understanding how the performance and the phenology of the invasive legume Lupinus polyphyllus vary with latitude. We sampled data across a >2000 km latitudinal gradient from Central to Northern Europe. We quantified variation in phenology of flowering and fruiting of L. polyphyllus using >1600 digital photos of inflorescences from 220 individual plants observed weekly at 22 sites. The day of the year at which different phenological phases were reached, increased 1.3–1.8 days per degree latitude, whereas the growing degree days (gdd) required for these phenological phases decreased 5–16 gdd per degree latitude. However, this difference disappeared, when the day length of each day included in the calculation of gdd was considered. The day of the year of the earliest and the latest climatic zone to reach any of the three studied phenological phases differed by 23–30 days and temperature requirements to reach these stages differed between 62 and 236 gdd. Probably, the invasion of this species will further increase in the northern part of Europe over the next decades due to climate warming. For invasive species control, our results suggest that in countries with a large latitudinal extent, the mowing date should shift by ca. one week per 500 km at sites with similar elevations.

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.

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.

Plant phenology, i. e. the timing of life cycle events, is related to individual fitness and species distribution ranges. Among the environmental factors, phenology is mostly driven by temperature and day length. Rapid adaptation of their phenology may also be important for the success of invasive plant species. Our main aim was to understand how the performance, timing, and temperature dependence of the phenology of the invasive legume Lupinus polyphyllus varies with latitude. L. polyphyllus is one of the most frequent invasive species in Europe, and the gained information may help to make management more effective by adjustments to latitude and phenology. Methods: We quantified variation in phenology across a >2000 km latitudinal gradient from Central to Northern Europe. We sampled data of flowering and fruiting of L. polyphyllus using >1600 digital photos of inflorescences from 220 individual plants observed weekly at 22 locations. We calculated the accumulated growing degree days for each observation date at each site from the temperature data of the meteorological stations, which are listed in the metadata of the dataset., License:This work is licensed under a CC0 1.0 Universal (CC0 1.0) Public Domain Dedication license.

Question: When evaluating forests in terms of their biodiversity, distinctiveness and naturalness, the affinity of the constituent species to forests is a crucial parameter. Here we ask to what extent are vascular plant species associated with forests, and does species’ affinity to forests vary between European regions?. Location: Temperate and boreal forest biome of Northwestern and Central Europe. Methods: We compiled EuForPlant, a new extensive list of forest vascular plant species in 24 regions spread across 13 European countries using vegetation databases and expert knowledge. Species were region-specifically classified into four categories reflecting the degree of their affinity to forest habitats: 1.1, species of forest interiors; 1.2, species of forest edges and forest openings; 2.1, species that can be found in forest as well as open vegetation; and 2.2, species that can be found partly in forest, but mainly in open vegetation. An additional “O” category was distinguished, covering species typical for non-forest vegetation. Results: EuForPlant comprises 1,726 species, including 1,437 herb-layer species, 159 shrubs, 107 trees, 19 lianas and 4 epiphytic parasites. Across regions, generalist forest species (with 450 and 777 species classified as 2.1 and 2.2, respectively) significantly outnumbered specialist forest species (with 250 and 137 species classified as 1.1 and 1.2, respectively). Even though the degree of shifting between the categories of forest affinity among regions was relatively low (on average, 17.5%), about one-third of the forest species (especially 1.2 and 2.2) swapped categories in at least one of the study regions. Conclusions: The proposed list can be used widely in vegetation science and global change ecology related to forest biodiversity and community dynamics. Shifting of forest affinity among regions emphasizes the importance of a continental-scale forest plant species list with regional specificity.

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

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.

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

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

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