Intraspecific variability of leaf form and function across habitat types.

Trait-based ecology has already revealed main independent axes of trait variation defining trait spaces that summarize plant adaptive strategies, but often ignoring intraspecific trait variability (ITV). By using empirical ITV-level data for two independent dimensions of leaf form and function and 167 species across five habitat types (coastal dunes, forests, grasslands, heathlands, wetlands) in the Italian peninsula, we found that ITV: (i) rotated the axes of trait variation that define the trait space; (ii) increased the variance explained by these axes and (iii) affected the functional structure of the target trait space. However, the magnitude of these effects was rather small and depended on the trait and habitat type. Our results reinforce the idea that ITV is context-dependent, calling for careful extrapolations of ITV patterns across traits and spatial scales. Importantly, our study provides a framework that can be used to start integrating ITV into trait space analyses.

Unpunctual in diversity: The effect of stand species richness on spring phenology of deciduous tree stands varies among species and years.

Climatic drivers alone do not adequately explain the regional variation in budburst timing in deciduous forests across Europe. Stand-level factors, such as tree species richness, might affect budburst timing by creating different microclimates under the same site macroclimate. We assessed different phases of the spring phenology (start, midpoint, end, and overall duration of the budburst period) of four important European tree species (Betula pendula, Fagus sylvatica, Quercus robur and Tilia cordata) in monocultures and four-species mixture stands of a common garden tree biodiversity experiment in Belgium (FORBIO) in 2021 and 2022. Microclimatic differences between the stands in terms of bud chilling, temperature forcing, and soil temperature were considerable, with four-species mixtures being generally colder than monocultures in spring, but not in winter. In the colder spring of 2021, at the stand level, the end of the budburst period was advanced, and its overall duration shortened, in the four-species mixtures. At species level, this response was significant for F. sylvatica. In the warmer spring of 2022, advances in spring phenology in four-species stands were observed again in F. sylvatica and, less markedly, in B. pendula but without a general response at the stand level. Q. robur showed specific patterns with delayed budburst start in 2021 in the four-species mixtures and very short budburst duration for all stands in 2022. Phenological differences between monocultures and four-species mixtures were linked to microclimatic differences in light availability rather than in temperature as even comparatively colder microclimates showed an advanced phenology. Compared to weather conditions, tree species richness had a lower impact on budburst timing, but this impact can be of importance for key species like F. sylvatica and colder springs. These results indicate that forest biodiversity can affect budburst phenology, with wider implications, especially for forest- and land surface models.

Impact of coppicing on microclimate and understorey vegetation diversity in an ancient Mediterranean oak forest.

Coppicing is one of the oldest silvicultural practices and is still widely applied to produce renewable energy from broadleaf forests. However, the consequences on microclimate and understorey vegetation are still poorly understood, especially in Mediterranean oak forests. With the ongoing changes in the climate system and global biodiversity loss, a better understanding of how the forest temperature buffering capacity and below-canopy plant community are impacted by coppicing is crucial. Here we quantify microclimate and understorey vegetation changes in adjacent ancient coppice-with-standards and high forest stands dominated by oaks in Italy, where these systems have been applied for a long time. Air and soil temperatures were recorded for 2.5 years, and nested vegetation plots were used to analyse coppicing effects on species composition, taxonomic, phylogenetic, and functional diversity. Coppicing significantly reduced the forest temperature buffering capacity. The mean of the daily maximum temperatures over the entire period was 1.45 °C higher in the coppiced sites, whereas the mean of the daily minimum temperatures was 0.62 °C lower than in the high forest. Coppicing increased understorey species richness by favouring generalist taxa, but significantly decreased the proportion of forest specialists. The understorey community in coppiced forests consisted of more warm-adapted species. Moreover, coppicing also led to a loss of phylogenetic evenness and to shifts in diversity and community weighted mean Leaf Dry Matter content, pointing to habitat filtering and acclimation processes. In sum, we show that coppicing affects microclimate and understory vegetation in a direction that can exacerbate the effects of climate change, negatively affecting the oak forest specialist flora and its phylogenetic evenness.

Trade-offs in biodiversity and ecosystem services between edges and interiors in European forests.

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.

Limited effects of population age on the genetic structure of spatially isolated forest herb populations in temperate Europe.

Due to multiple land-cover changes, forest herb populations residing in forest patches embedded in agricultural landscapes display different ages and, thus, experience differences in genetic exchange, mutation accumulation and genetic drift. The extent of divergence in present-day population genetic structure among these populations of different ages remains unclear, considering their diverse breeding systems and associated pollinators. Answering this question is essential to understand these species' persistence, maintenance of evolutionary potential and adaptability to changing environments. We applied a multi-landscape setup to compare the genetic structure of forest herb populations across forest patches of different ages (18-338 years). We studied the impact on three common slow-colonizer herb species with distinct breeding systems and associated pollinators: Polygonatum multiflorum (outcrossing, long-distance pollinators), Anemone nemorosa (outcrossing, short-distance pollinators) and Oxalis acetosella (mixed breeding). We aimed to assess if in general older populations displayed higher genetic diversity and lower differentiation than younger ones. We also anticipated that P. multiflorum would show the smallest while O. acetosella the largest difference, between old and young populations. We found that older populations had a higher observed heterozygosity (H o) but a similar level of allelic richness (A r) and expected heterozygosity (H e) as younger populations, except for A. nemorosa, which exhibited higher A r and H e in younger populations. As populations aged, their pairwise genetic differentiation measured by D PS decreased independent of species identity while the other two genetic differentiation measures showed either comparable levels between old and young populations (G" ST) or inconsistency among three species (cGD). The age difference of the two populations did not explain their genetic differentiation. Synthesis: We found restricted evidence that forest herb populations with different ages differ in their genetic structure, indicating that populations of different ages can reach a similar genetic structure within decades and thus persist in the long term after habitat disturbance. Despite their distinct breeding systems and associated pollinators, the three studied species exhibited partly similar genetic patterns, suggesting that their common characteristics, such as being slow colonizers or their ability to propagate vegetatively, are important in determining their long-term response to land-cover change.

Combining multiple investigative approaches to unravel functional responses to global change in the understorey of temperate forests.

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.

Using warming tolerances to predict understory plant responses to climate change.

Climate change is pushing species towards and potentially beyond their critical thermal limits. The extent to which species can cope with temperatures exceeding their critical thermal limits is still uncertain. To better assess species' responses to warming, we compute the warming tolerance (ΔTniche ) as a thermal vulnerability index, using species' upper thermal limits (the temperature at the warm limit of their distribution range) minus the local habitat temperature actually experienced at a given location. This metric is useful to predict how much more warming species can tolerate before negative impacts are expected to occur. Here we set up a cross-continental transplant experiment involving five regions distributed along a latitudinal gradient across Europe (43° N-61° N). Transplant sites were located in dense and open forests stands, and at forest edges and in interiors. We estimated the warming tolerance for 12 understory plant species common in European temperate forests. During 3 years, we examined the effects of the warming tolerance of each species across all transplanted locations on local plant performance, in terms of survival, height, ground cover, flowering probabilities and flower number. We found that the warming tolerance (ΔTniche ) of the 12 studied understory species was significantly different across Europe and varied by up to 8°C. In general, ΔTniche were smaller (less positive) towards the forest edge and in open stands. Plant performance (growth and reproduction) increased with increasing ΔTniche across all 12 species. Our study demonstrated that ΔTniche of understory plant species varied with macroclimatic differences among regions across Europe, as well as in response to forest microclimates, albeit to a lesser extent. Our findings support the hypothesis that plant performance across species decreases in terms of growth and reproduction as local temperature conditions reach or exceed the warm limit of the focal species.

Forest understorey flowering phenology responses to experimental warming and illumination.

Species are altering their phenology to track warming temperatures. In forests, understorey plants experience tree canopy shading resulting in light and temperature conditions, which strongly deviate from open habitats. Yet, little is known about understorey phenology responses to forest microclimates. We recorded flowering onset, peak, end and duration of 10 temperate forest understorey plant species in two mesocosm experiments to understand how phenology is affected by sub-canopy warming and how this response is modulated by illumination, which is related to canopy change. Furthermore, we investigated whether phenological sensitivities can be explained by species' characteristics, such as thermal niche. We found a mean advance of flowering onset of 7.1 d per 1°C warming, more than previously reported in studies not accounting for microclimatic buffering. Warm-adapted species exhibited greater advances. Temperature sensitivity did not differ between early- and later-flowering species. Experimental illumination did not significantly affect species' phenological temperature sensitivities, but slightly delayed flowering phenology independent from warming. Our study suggests that integrating sub-canopy temperature and light availability will help us better understand future understorey phenology responses. Climate warming together with intensifying canopy disturbances will continue to drive phenological shifts and potentially disrupt understorey communities, thereby affecting forest biodiversity and functioning.

Novel light regimes in European forests.

Tree canopies are one of the most recognizable features of forests, providing shelter from external influences to a myriad of species that live within and below the tree foliage. Canopy disturbances are now increasing across European forests, and climate-change-induced drought is a key driver, together with pests and pathogens, storms and fire. These disturbances are opening the canopy and exposing below-canopy biodiversity and functioning to novel light regimes-spatial and temporal characteristics of light distribution at forest floors not found previously. The majority of forest biodiversity occurs in the shade within and below tree canopies, and numerous ecosystem processes are regulated at the forest floor. Altered light regimes, in interaction with other global change drivers, can thus strongly impact forest biodiversity and functioning. As recent European droughts are unprecedented in the past two millennia, and this has initiated probably the largest pulse of forest disturbances in almost two centuries, we urgently need to quantify, understand and predict the impacts of novel light regimes on below-canopy forest biodiversity and functions. This will be a crucial element in delivering much-needed information for policymakers and managers to adapt European forests to future no-analogue conditions.

Evaluating plant lineage losses and gains in temperate forest understories: a phylogenetic perspective on climate change and nitrogen deposition.

Global change has accelerated local species extinctions and colonizations, often resulting in losses and gains of evolutionary lineages with unique features. Do these losses and gains occur randomly across the phylogeny? We quantified: temporal changes in plant phylogenetic diversity (PD); and the phylogenetic relatedness (PR) of lost and gained species in 2672 semi-permanent vegetation plots in European temperate forest understories resurveyed over an average period of 40 yr. Controlling for differences in species richness, PD increased slightly over time and across plots. Moreover, lost species within plots exhibited a higher degree of PR than gained species. This implies that gained species originated from a more diverse set of evolutionary lineages than lost species. Certain lineages also lost and gained more species than expected by chance, with Ericaceae, Fabaceae, and Orchidaceae experiencing losses and Amaranthaceae, Cyperaceae, and Rosaceae showing gains. Species losses and gains displayed no significant phylogenetic signal in response to changes in macroclimatic conditions and nitrogen deposition. As anthropogenic global change intensifies, temperate forest understories experience losses and gains in specific phylogenetic branches and ecological strategies, while the overall mean PD remains relatively stable.

Les changements globaux accélèrent les processus de colonisation et d'extinction locales d'espèces, aboutissant à des gains ou à des pertes de lignées évolutives uniques. Ces gains et pertes se produisent-ils de manière aléatoire dans l'arbre phylogénétique ? Nous avons mesuré: les changements de diversité phylogénétique; et la parenté phylogénétique des espèces végétales gagnées ou perdues dans 2672 placettes semi-permanentes disposées dans le sous-bois de forêts tempérées d'Europe sur une période moyenne de 40 ans. Une fois corrigée par la richesse spécifique, la diversité phylogénétique a légèrement augmenté au cours du temps dans les différentes placettes. Les espèces perdues ont une plus grande parenté phylogénétique que les espèces gagnées. Les espèces gagnées sont donc issues d'un plus grand nombre de lignées évolutives que les espèces perdues. Certaines lignées ont gagné ou perdu davantage d'espèces que ce qui est prédit par le hasard : les Ericaceae, les Fabaceae et les Orchidaceae ayant davantage perdu, tandis que les Amaranthaceae, les Cyperaceae, et les Rosaceae ont plus gagné. Il n'y a pas de signal phylogénétique des gains ou pertes d'espèces en réponse aux changements de conditions macroclimatiques ou des dépôts atmosphériques d'azote. Alors que les changements globaux d'origine anthropique s'intensifient, les sous-bois des forêts tempérées connaissent des gains et des pertes de certaines lignées évolutives et de certaines stratégies écologiques, sans que la diversité phylogénétique moyenne ne s'en trouve véritablement affectée.

El cambio global ha acelerado las extinciones y colonizaciones a escala local, lo que a menudo ha supuesto pérdidas y ganancias de linajes evolutivos con características únicas. Ahora bien, ¿estas pérdidas y ganancias ocurren aleatoriamente a lo largo de la filogenia? Cuantificamos: los cambios temporales en la diversidad filogenética de las plantas; y la relación filogenética de las especies perdidas y ganadas en 2.672 parcelas de vegetación semipermanente en sotobosques templados europeos y re-muestreadas durante un período promedio de 40 años. Al controlar por las diferencias en la riqueza de especies, la diversidad filogenética aumentó ligeramente con el tiempo y entre parcelas. Además, las especies perdidas dentro de las parcelas exhibieron un mayor grado de relación filogenética que las especies ganadas. Esto implica que las especies ganadas se originaron en un conjunto de linajes evolutivos más diversos que las especies perdidas. Ciertos linajes también perdieron y ganaron más especies de las esperadas aleatoriamente: Ericaceae, Fabaceae y Orchidaceae experimentaron pérdidas y Amaranthaceae, Cyperaceae y Rosaceae mostraron ganancias. Las pérdidas y ganancias de especies no mostraron ninguna señal filogenética significativa en respuesta a los cambios en las condiciones macro-climáticas y la deposición de nitrógeno. A medida que se intensifica el cambio global antropogénico, los sotobosques temperados experimentan pérdidas y ganancias en ramas filogenéticas y estrategias ecológicas específicas, mientras que la diversidad filogenética media general permanece relativamente estable.

Microclimate reveals the true thermal niche of forest plant species.

Species distributions are conventionally modelled using coarse-grained macroclimate data measured in open areas, potentially leading to biased predictions since most terrestrial species reside in the shade of trees. For forest plant species across Europe, we compared conventional macroclimate-based species distribution models (SDMs) with models corrected for forest microclimate buffering. We show that microclimate-based SDMs at high spatial resolution outperformed models using macroclimate and microclimate data at coarser resolution. Additionally, macroclimate-based models introduced a systematic bias in modelled species response curves, which could result in erroneous range shift predictions. Critically important for conservation science, these models were unable to identify warm and cold refugia at the range edges of species distributions. Our study emphasizes the crucial role of microclimate data when SDMs are used to gain insights into biodiversity conservation in the face of climate change, particularly given the growing policy and management focus on the conservation of refugia worldwide.

Heading for a fall: The fate of old wind-thrown beech trees (Fagus sylvatica) is detectable in their growth pattern.

Common beech (Fagus sylvatica) is one of the most important deciduous tree species in European forests. However, climate-change-induced drought may threaten its dominant position. The Sonian Forest close to Brussels (Belgium) is home to some of the largest beech trees in the world. This UNESCO world heritage site is famous for its high density of very large beech trees as a result of its climatic suitability, fertile soil conditions, and past management. Here we utilized tree-ring data from increment cores to investigate the growth of these old and monumental beech trees, evaluating their growth trends, response to past climate, and the effect of mast years on 39 living and 16 recently wind-thrown trees. Our analysis reveals that the sampled trees were generally sensitive to spring and summer droughts but recovered quickly after such an extreme climatic event. The growth trend of living trees has remained high and only shows a slight, statistically insignificant, decline over the past 50 years. Although the overall growth rate remains strong (BAI 50 cm2/year), the past five decades have shown strong inter-annual growth variations due to frequent and more intense droughts combined with an increased frequency of mast years. We also found notable differences in growth patterns between the living trees and those that had recently been wind-thrown. While there were no significant differences between living and wind-thrown trees in response to droughts, heatwaves, or mast years when examining year-to-year growth changes, the wind-thrown trees did exhibit considerably lower overall growth rates and a significant downward trend in growth (BAI -0.57 cm2/year). This difference in growth trends has been apparent since at least the 1980s. Overall, the findings of this study can provide valuable insights for understanding the long-term dynamics of lowland beech forests and their responses to climate change.

Impact of patch age and size on forest soil characteristics in European agricultural landscapes.

Many landscapes worldwide are characterized by the presence of a mosaic of forest patches with contrasting age and size embedded in a matrix of agricultural land. However, our understanding of the effects of these key forest patch features on the soil nutrient status (in terms of nitrogen, carbon, and phosphorus) and soil pH is still limited due to a lack of large-scale data. To address this research gap, we analyzed 830 soil samples from nearly 200 forest patches varying in age (recent versus ancient forests) and size (small versus larger patches) along a 2500-km latitudinal gradient across Europe. We also considered environmental covariates at multiple scales to increase the generality of our research, including variation in macroclimate, nitrogen deposition rates, forest cover in a buffer zone, basal area and soil type. Multiple linear mixed-effects models were performed to test the combined effects of patch features and environmental covariates on soil nutrients and pH. Recent patches had higher total soil phosphorus concentrations and stocks in the mineral soil layer, along with a lower nitrogen to phosphorus ratio within that layer. Small patches generally had a higher mineral soil pH. Mineral soil nitrogen stocks were lower in forest patches with older age and larger size, as a result of a significant interactive effect. Additionally, environmental covariates had significant effects on soil nutrients, including carbon, nitrogen, phosphorus, and their stoichiometry, depending on the specific covariates. In some cases, the effect of patch age on mineral soil phosphorus stocks was greater than that of environmental covariates. Our findings underpin the important roles of forest patch age and size for the forest soil nutrient status. Long-term studies assessing edge effects and soil development in post-agricultural forests are needed, especially in a context of changing land use and climate.

ForestClim-Bioclimatic variables for microclimate temperatures of European forests.

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 × 25 m2 resolution.

Forest edge effects on moss growth are amplified by drought.

Forest fragmentation increases the amount of edges in the landscape. Differences in wind, radiation, and vegetation structure create edge-to-interior gradients in forest microclimate, and these gradients are likely to be more pronounced during droughts and heatwaves. Although the effects of climate extremes on edge influences have potentially strong and long-lasting impacts on forest understory biodiversity, they are not well understood and are not often considered in management and landscape planning. Here we used a novel method of retrospectively quantifying growth to assess biologically relevant edge influences likely caused by microclimate using Hylocomium splendens, a moss with annual segments. We examined how spatio-temporal variation in drought across 3 years and 46 sites in central Sweden, affected the depth and magnitude of edge influences. We also investigated whether edge effects during drought were influenced by differences in forest structure. Edge effects were almost twice as strong in the drought year compared to the non-drought years, but we did not find clear evidence that they penetrated deeper into the forest in the drought year. Edge influences were also greater in areas that had fewer days with rain during the drought year. Higher levels of forest canopy cover and tree height buffered the magnitude of edge influence in times of drought. Our results demonstrate that edge effects are amplified by drought, suggesting that fragmentation effects are aggravated when droughts become more frequent and severe. Our results suggest that dense edges and buffer zones with high canopy cover can be important ways to mitigate negative drought impacts in forest edges.

Usefulness of cultivar-level calibration of AquaCrop for vegetables depends on the crop and data availability.

As a result of climate change, climatic extremes are expected to increase. For high-value crops like vegetables, irrigation is a potentially economically viable adaptation measure in western Europe. To optimally schedule irrigation, decision support systems based on crop models like AquaCrop are increasingly used by farmers. High value vegetable crops like cauliflower or spinach are grown in two distinct growth cycles per year and, additionally, have a high turnover rate of new varieties. To successfully deploy the AquaCrop model in a decision support system, it requires a robust calibration. However, it is not known whether parameters can be conserved over both growth periods, nor whether a cultivar dependent model calibration is always required. Furthermore, when data are collected from farmers' fields, there are constraints in data availability and uncertainty. We collected data from commercial cauliflower and spinach fields in Belgium in 2019, 2020 and 2021 during different growing periods and of different cultivars. With the use of a Bayesian calibration, we confirmed the need for a condition or cultivar specific calibration for cauliflower, while for spinach, splitting the data per cultivar or pooling the data together did not improve uncertainty on the model simulations. However, due to uncertainties arising from field specific soil and weather conditions, or measurement errors from calibration data, real time field specific adjustments are advised to simulations when using AquaCrop as decision support tool. Remotely sensed or in situ ground data may be invaluable information to reduce uncertainty on model simulations.

Intra-individual variation in Galium odoratum is affected by experimental drought and shading.

BACKGROUND AND AIMS: Climate-change induced warmer spring temperatures advance tree leaf-out and result in earlier shading of the forest floor. Climate change also leads to more frequent droughts. Forest understorey herbs may respond to these environmental changes by varying traits at different hierarchical levels of organization. While trait mean variation at the inter-individual level in response to environmental changes is well-studied, little is known about how variation at the intra-individual level responds. METHODS: We sampled genets of the forest understorey herb Galium odoratum from 21 populations in three regions in Germany, varying in microclimatic conditions. The genets were transplanted into a common garden, where we applied shading and drought treatments. We measured plant height and leaf length and width, and calculated the coefficient of variation (CV) at different hierarchical levels: intra-population, intra-genet, intra-ramet and intra-shoot. KEY RESULTS: Variance partitioning showed that intra-shoot CV represented most of the total variation, followed by intra-ramet CV. We found significant variation in CV of plant height and leaf width among populations of origin, indicating that CV is at least partly genetically based. The soil temperature at populations' origins correlated negatively with CV in plant height, suggesting adaptation to local conditions. Furthermore, we observed that early shade led to increased intra-ramet CV in leaf length, while drought reduced intra-shoot CV in leaf width. Finally, intra-shoot leaf width mean and CV were independent under control conditions but correlated under drought. CONCLUSIONS: Our experimental results reveal correlations of intra-individual variation with soil temperature, indicating that intra-individual variation can evolve and may be adaptive. Intra-individual variation responded plastically to drought and shading, suggesting functional changes to improve light capture and reduce evapotranspiration. In conclusion, intra-individual variation makes up the majority of total trait variation in this species and can play a key role in plant adaptation to climatic change.

Divergent roles of herbivory in eutrophying forests.

Ungulate populations are increasing across Europe with important implications for forest plant communities. Concurrently, atmospheric nitrogen (N) deposition continues to eutrophicate forests, threatening many rare, often more nutrient-efficient, plant species. These pressures may critically interact to shape biodiversity as in grassland and tundra systems, yet any potential interactions in forests remain poorly understood. Here, we combined vegetation resurveys from 52 sites across 13 European countries to test how changes in ungulate herbivory and eutrophication drive long-term changes in forest understorey communities. Increases in herbivory were associated with elevated temporal species turnover, however, identities of winner and loser species depended on N levels. Under low levels of N-deposition, herbivory favored threatened and small-ranged species while reducing the proportion of non-native and nutrient-demanding species. Yet all these trends were reversed under high levels of N-deposition. Herbivores also reduced shrub cover, likely exacerbating N effects by increasing light levels in the understorey. Eutrophication levels may therefore determine whether herbivory acts as a catalyst for the "N time bomb" or as a conservation tool in temperate forests.

Protected areas provide thermal buffer against climate change.

Climate change is pushing temperatures beyond the thermal tolerance of many species. Whether protected areas (PAs) can serve as climate change refugia for biodiversity has not yet been explored. We find that PAs of natural (seminatural) vegetation effectively cool the land surface temperature, particularly the daily maximum temperature in the tropics, and reduce diurnal and seasonal temperature ranges in boreal and temperate regions, as compared to nonprotected areas that are often disturbed or converted to various land uses. Moreover, protected forests slow the rate of warming more at higher latitudes. The warming rate in protected boreal forests is up to 20% lower than in their surroundings, which is particularly important for species in the boreal where warming is more pronounced. The fact that nonprotected areas with the same type of vegetation as PAs show reduced warming buffer capacity highlights the importance of conservation to stabilize the local climate and safeguard biodiversity.

Forest structure and composition alleviate human thermal stress.

Current climate change aggravates human health hazards posed by heat stress. Forests can locally mitigate this by acting as strong thermal buffers, yet potential mediation by forest ecological characteristics remains underexplored. We report over 14 months of hourly microclimate data from 131 forest plots across four European countries and compare these to open-field controls using physiologically equivalent temperature (PET) to reflect human thermal perception. Forests slightly tempered cold extremes, but the strongest buffering occurred under very hot conditions (PET >35°C), where forests reduced strong to extreme heat stress day occurrence by 84.1%. Mature forests cooled the microclimate by 12.1 to 14.5°C PET under, respectively, strong and extreme heat stress conditions. Even young plantations reduced those conditions by 10°C PET. Forest structure strongly modulated the buffering capacity, which was enhanced by increasing stand density, canopy height and canopy closure. Tree species composition had a more modest yet significant influence: that is, strongly shade-casting, small-leaved evergreen species amplified cooling. Tree diversity had little direct influences, though indirect effects through stand structure remain possible. Forests in general, both young and mature, are thus strong thermal stress reducers, but their cooling potential can be even further amplified, given targeted (urban) forest management that considers these new insights.