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Plant communities are being exposed to changing environmental conditions all around the globe, leading to alterations in plant diversity, community composition, and ecosystem functioning. For herbaceous understorey communities in temperate forests, responses to global change are postulated to be complex, due to the presence of a tree layer that modulates understorey responses to external pressures such as climate change and changes in atmospheric nitrogen deposition rates. Multiple investigative approaches have been put forward as tools to detect, quantify and predict understorey responses to these global-change drivers, including, among others, distributed resurvey studies and manipulative experiments. These investigative approaches are generally designed and reported upon in isolation, while integration across investigative approaches is rarely considered. In this study, we integrate three investigative approaches (two complementary resurvey approaches and one experimental approach) to investigate how climate warming and changes in nitrogen deposition affect the functional composition of the understorey and how functional responses in the understorey are modulated by canopy disturbance, that is, changes in overstorey canopy openness over time. Our resurvey data reveal that most changes in understorey functional characteristics represent responses to changes in canopy openness with shifts in macroclimate temperature and aerial nitrogen deposition playing secondary roles. Contrary to expectations, we found little evidence that these drivers interact. In addition, experimental findings deviated from the observational findings, suggesting that the forces driving understorey change at the regional scale differ from those driving change at the forest floor (i.e., the experimental treatments). Our study demonstrates that different approaches need to be integrated to acquire a full picture of how understorey communities respond to global change.

Brief introduction: What are microclimates and why are they important?: Microclimate science has developed into a global discipline. Microclimate science is increasingly used to understand and mitigate climate and biodiversity shifts. Here, we provide an overview of the current status of microclimate ecology and biogeography in terrestrial ecosystems, and where this field is heading next. Microclimate investigations in ecology and biogeography: We highlight the latest research on interactions between microclimates and organisms, including how microclimates influence individuals, and through them populations, communities and entire ecosystems and their processes. We also briefly discuss recent research on how organisms shape microclimates from the tropics to the poles. Microclimate applications in ecosystem management: Microclimates are also important in ecosystem management under climate change. We showcase new research in microclimate management with examples from biodiversity conservation, forestry and urban ecology. We discuss the importance of microrefugia in conservation and how to promote microclimate heterogeneity. Methods for microclimate science: We showcase the recent advances in data acquisition, such as novel field sensors and remote sensing methods. We discuss microclimate modelling, mapping and data processing, including accessibility of modelling tools, advantages of mechanistic and statistical modelling and solutions for computational challenges that have pushed the state-of-the-art of the field. What's next?: We identify major knowledge gaps that need to be filled for further advancing microclimate investigations, applications and methods. These gaps include spatiotemporal scaling of microclimate data, mismatches between macroclimate and microclimate in predicting responses of organisms to climate change, and the need for more evidence on the outcomes of microclimate management.

• TreeDivNet is a global network for investigating the benefits and drawbacks of mixed-species planted forests. • After 23 years, TreeDivNet is yielding evidence in favour of mixing tree species in stands to increase tree performance. • More research is needed, especially in the global South, to identify optimal species mixes and to translate scientific knowledge into operational guidelines.

Microclimate research gained renewed interest over the last decade and its importance for many ecological processes is increasingly being recognized. Consequently, the call for high-resolution microclimatic temperature grids across broad spatial extents is becoming more pressing to improve ecological models. Here, we provide a new set of open-access bioclimatic variables for microclimate temperatures of European forests at 25 x 25 m2 resolution.

Understanding forest understorey community response to environmental change, including management actions, is vital given the understorey's importance for biodiversity conservation and ecosystem functioning. The Natural World Heritage Hyrcanian temperate forests (Iran) provide an ideal template for furnishing an appreciation of how management actions can mitigate undesired climate change effects, due to the forests’ broad environmental gradients, isolation from colonization sources and varied light environments. We used records of 95 understorey plant species from 512 plots to model their probability of occurrence as a function of contemporary climate and soil variables, and canopy cover. For 65 species with good predictive accuracy, we then projected two climate scenarios in the context of either increasing or decreasing canopy cover, to assess whether overstorey management could mitigate or aggravate climate change effects. Climate variables were the most important predictors for the distribution of all species. Soil and canopy cover varied in importance depending on understorey growth form. Climate change was projected to negatively affect future probabilities of occurrence. However, management, here represented by canopy cover change, is predicted to modify this trajectory for some species groups. Models predict increases in light-adapted and generalist forbs with reduced canopy cover, while graminoids and ferns still decline. Increased canopy cover is projected to buffer an otherwise significant decreasing response of cold-adapted species to climate change. However, increasing canopy cover is not projected to buffer the predicted negative impact of climate change on shade-adapted forest specialists. Inconsistent responses of different species and/or growth forms to climate change and canopy cover reflect their complicated life histories and habitat preferences. Canopy cover management may help prevent the climate change induced loss of some important groups for

Forest canopies buffer macroclimatic temperature fluctuations. However, we do not know if and how the capacity of canopies to buffer understorey temperature will change with accelerating climate change. Here we map the difference (offset) between temperatures inside and outside forests in the recent past and project these into the future in boreal, temperate and tropical forests. Using linear mixed-effect models, we combined a global database of 714 paired time series of temperatures (mean, minimum and maximum) measured inside forests vs. in nearby open habitats with maps of macroclimate, topography and forest cover to hindcast past (1970–2000) and to project future (2060–2080) temperature differences between free-air temperatures and sub-canopy microclimates. For all tested future climate scenarios, we project that the difference between maximum temperatures inside and outside forests across the globe will increase (i.e. result in stronger cooling in forests), on average during 2060–2080, by 0.27 ± 0.16 °C (RCP2.6) and 0.60 ± 0.14 °C (RCP8.5) due to macroclimate changes. This suggests that extremely hot temperatures under forest canopies will, on average, warm less than outside forests as macroclimate warms. This knowledge is of utmost importance as it suggests that forest microclimates will warm at a slower rate than non-forested areas, assuming that forest cover is maintained. Species adapted to colder growing conditions may thus find shelter and survive longer than anticipated at a given forest site. This highlights the potential role of forests as a whole as microrefugia for biodiversity under future climate change.

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.

1. The unwanted development of dense understorey vegetation composed of resource‐acquisitive, tall plant species competing strongly with tree regeneration can pose formidable problems for managers attempting to regenerate temperate forests. Despite many studies on the effects of understorey removal, no comprehensive review has summarised and quantified its effects on subsequent life stages of tree regeneration in temperate forests. 2. We synthesised data from 32 experimental studies from temperate forest regions. We used meta‐analytic techniques to find general patterns in terms of the characteristics of the understorey, overstorey and characteristics of the regenerating tree species, which are most responsible for possible positive understorey removal effects on early life stages of tree regeneration, i.e. emergence, survival and growth. 3. Both seedling survival and growth increased in response to understorey removal; emergence did not show a clear pattern. Seedlings growing free from competition mainly increased their biomass growth (total and above‐ground), whereas diameter and height growth responded less. These positive effects were largest when removing denser understorey vegetation and under more open overstorey conditions. Multiple management options influenced the regeneration responses to understorey removal. For instance, growth of older, planted seedlings responded less to removal, whereas protection against large browsers increased growth responses. 4. Tree species with differing strategies responded differently to understorey removal. Growth and survival responses of early‐successional species responded more strongly to understorey removal than mid‐ or late‐successional tree species. 5. Synthesis and applications. Our study showed that understorey removal can have strong positive effects on tree regeneration across temperate forest contexts. The magnitude of these effects depended on overstorey and understorey conditions, but also on the type of tree

Questions: Light availability at the forest floor affects many forest ecosystem processes, and is often quantified indirectly through easy‐to‐measure stand characteristics. We investigated how three such characteristics, basal area, canopy cover and canopy closure, were related to each other in structurally complex mixed forests. We also asked how well they can predict the light‐demand signature of the forest understorey (estimated as the mean Ellenberg indicator value for light [“EIVLIGHT”] and the proportion of “forest specialists” [“%FS”] within the plots). Furthermore, we asked whether accounting for the shade‐casting ability of individual canopy species could improve predictions of EIVLIGHT and %FS. Location: A total of 192 study plots from nineteen temperate forest regions across Europe. Methods: In each plot, we measured stand basal area (all stems >7.5 cm diameter), canopy closure (with a densiometer) and visually estimated the percentage cover of all plant species in the herb (<1 m), shrub (1–7 m) and tree layer (>7 m). We used linear mixed‐effect models to assess the relationships between basal area, canopy cover and canopy closure. We performed model comparisons, based on R2 and the Akaike Information Criterion (AIC), to assess which stand characteristics can predict EIVLIGHT and %FS best, and to assess whether canopy shade‐casting ability can significantly improve model fit. Results: Canopy closure and cover were weakly related to each other, but showed no relation with basal area. For both EIVLIGHT and %FS, canopy cover was the best predictor. Including the share of high‐shade‐casting species in both the basal‐area and cover models improved the model fit for EIVLIGHT, but not for %FS. Conclusions: The typically expected relationships between basal area, canopy cover and canopy closure were weak or even absent in structurally complex mixed forests. In these forests, easy‐to‐measure structural canopy characteristics were poor predictors of the understorey

1. A central challenge of today's ecological research is predicting how ecosystems will develop under future global change. Accurate predictions are complicated by (a) simultaneous effects of different drivers, such as climate change, nitrogen deposition and management changes; and (b) legacy effects from previous land use. 2. We tested whether herb layer biodiversity (i.e. richness, Shannon diversity and evenness) and functional (i.e. herb cover, specific leaf area [SLA] and plant height) responses to environmental change drivers depended on land‐use history. We used resurvey data from 192 plots across nineteen European temperate forest regions, with large spatial variability in environmental change factors. We tested for interactions between land‐use history, distinguishing ancient and recent (i.e. post‐agricultural) forests and four drivers: temperature, nitrogen deposition, and aridity at the regional scale and light dynamics at the plot‐scale. 3. Land‐use history significantly modulated global change effects on the functional signature of the herb layer (i.e. cover, SLA and plant height). Light availability was the main environmental driver of change interacting with land‐use history. We found greater herb cover and plant height decreases and SLA increases with decreasing light availability in ancient than in recent forests. Furthermore, we found greater decreases in herb cover with increased nitrogen deposition in ancient forests, whereas warming had the strongest decreasing effect on the herb cover in recent forests. Interactive effects between land‐use history and global change on biodiversity were not found, but species evenness increased more in ancient than in recent forests. 4. Synthesis. Our results demonstrate that land‐use history should not be overlooked when predicting forest herb layer responses to global change. Moreover, we found that herb layer composition in semi‐natural deciduous forests is mainly controlled by local canopy characteristics, re