Unexpected westward range shifts in European forest plants link to nitrogen deposition.

Climate change is commonly assumed to induce species' range shifts toward the poles. Yet, other environmental changes may affect the geographical distribution of species in unexpected ways. Here, we quantify multidecadal shifts in the distribution of European forest plants and link these shifts to key drivers of forest biodiversity change: climate change, atmospheric deposition (nitrogen and sulfur), and forest canopy dynamics. Surprisingly, westward distribution shifts were 2.6 times more likely than northward ones. Not climate change, but nitrogen-mediated colonization events, possibly facilitated by the recovery from past acidifying deposition, best explain westward movements. Biodiversity redistribution patterns appear complex and are more likely driven by the interplay among several environmental changes than due to the exclusive effects of climate change alone.

Long-term nitrogen deposition reduces the diversity of nitrogen-fixing plants.

Biological nitrogen fixation is a fundamental part of ecosystem functioning. Anthropogenic nitrogen deposition and climate change may, however, limit the competitive advantage of nitrogen-fixing plants, leading to reduced relative diversity of nitrogen-fixing plants. Yet, assessments of changes of nitrogen-fixing plant long-term community diversity are rare. Here, we examine temporal trends in the diversity of nitrogen-fixing plants and their relationships with anthropogenic nitrogen deposition while accounting for changes in temperature and aridity. We used forest-floor vegetation resurveys of temperate forests in Europe and the United States spanning multiple decades. Nitrogen-fixer richness declined as nitrogen deposition increased over time but did not respond to changes in climate. Phylogenetic diversity also declined, as distinct lineages of N-fixers were lost between surveys, but the "winners" and "losers" among nitrogen-fixing lineages varied among study sites, suggesting that losses are context dependent. Anthropogenic nitrogen deposition reduces nitrogen-fixing plant diversity in ways that may strongly affect natural nitrogen fixation.

The concentrations of microelements in forest mushrooms are influenced by soil pH and C/N ratio and less by stand characteristics.

The fruitbodies or sporocarps formed by mushrooms can accumulate mineral elements, such as selenium, zinc or copper, making them an important source of micronutrients essential to humans. However, the literature about environmental factors affecting mineral composition in mushrooms is scarce and limited to the ambiguous impact of soil properties and region. In our study, we investigated the effects of tree stand characteristics (tree species and tree canopy cover), understory cover, and soil properties (pH and C/N ratio of the soil) on the concentration of minerals in six edible mushroom species: Laccaria laccata, L. proxima, L. amethystina, Lepista nuda, Lycoperdon perlatum, and Calvatia excipuliformis, collected on 20 plots covered by stands of different tree species composition and varying in the understory cover. We estimated the concentration of eight elements (Zn, Se, Mg, Mn, Cu, Co, Cr, Mo) using the ICP-MS (Inductively Coupled Plasma - Mass Spectrometry) technique and compared their concentration between the plots, using ordination and linear regression methods. Our research revealed that mushroom species identity, including its ecological role and phylogenetic affinity, had the greatest effect on the mineral composition of mushrooms. The effect of environmental factors depended also on the micronutrient identity. Some elements were not affected at all (Co, Cr), some only by soil reaction or C/N ratio (Mn, Mg, Cu), while others were influenced by both tree stand characteristics and soil properties (Se, Zn, Mo). This knowledge enables us to maximize the content of minerals in harvested mushrooms by collecting them in specific areas. For example, mushrooms, which are sources of Se and Zn, can be gathered in coniferous forests characterized by acid soils, low canopy cover, and minimal understory cover. This targeted collection approach can effectively increase the mineral content in harvested mushrooms, thereby enhancing their health benefits.

Fungal species and element type modulate the effects of environmental factors on the concentration of potentially toxic elements in mushrooms.

Numerous edible mushrooms accumulate Potentially Toxic Elements (PTE), such as cadmium, mercury, and lead, within their sporocarps. This accumulation poses a potential risk of poisoning for humans and is influenced by factors such as the mushroom species, type of element, and the level of industrialization in the region. In our study, we investigated how soil and tree stand characteristics, including C/N ratio, pH, tree diversity, canopy cover, and the proportion of deciduous trees, influence PTE concentration in mushrooms. We collected edible mushrooms from 20 plots situated in the Bialowieza Primeval Forest, one of Europe's best-preserved lowland forests. Plots varied in terms of tree species composition, with other factors minimized. We used ICP-MS (Inductively Coupled Plasma - Mass Spectrometry) technique to analyze the concentration of eight PTE (Ag, As, Cd, Ni, Pb, Sb, Sr, Tl) in eight edible mushroom species (M.procera, L.perlatum, R. butyracea, R.cyanoxantha, R.heterophylla, L.vellereus, A.mellea, and Xerocomellus chrysenteron). Our research revealed that the presence of the effect of specific factors on concentration of PTE and its direction depends on mushroom species and type of PTE. The proportion of deciduous tree species and pH of the topsoil layer emerged as the most influential factors affecting PTE concentration in mushroom samples. Tree species richness in the canopy layer did not affect PTE concentration in mushrooms, except for the concentration of Pb in X. chrysenteron. We observed a consistent profile of PTE concentration in mushrooms with similar ecological roles (ectomycorrhizal, saprotrophic, parasite mushrooms) and from comparable phylogenetic affinities.

Factors affecting composition of fatty acids in wild-growing forest mushrooms.

The importance of mushrooms as a food source is continually increasing. To investigate how environmental factors affect the nutritional value of mushrooms, we harvested them in eastern Poland, south-central Germany, and northwestern Belgium in plots with similar environmental conditions but varying in tree species composition and richness. We used gas chromatography-mass spectrometry (GC-MS) to analyze the fatty acid (FA) content of the mushrooms. Fungal species identity explained the largest part (40%) of the total variation in FA concentration and composition. Environmental factors accounted for 1-12% of variation. The concentration of FA, especially saturated fatty acids, decreased with increasing understory cover and increasing nitrogen concentration in the topsoil. The effect of tree species richness or tree species identity was negligible. Our results suggest that the nutritional value of mushrooms depends mainly on the species identity of fungi, but that their FA content is slightly higher in forests with less undergrowth and in nitrogen-poor soils.

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.

Threats to conservation from national security interests.

There is a growing trend of nation states invoking national security and emergency declarations to build state-sponsored infrastructure projects for border defense, energy production, and transportation. Established laws, regulations, and agreements for the protection of nature and cultural heritage within and between countries are becoming secondary to national security, compromising the function of protected areas, such as national parks, wilderness areas, and biosphere reserves that safeguard biodiversity, climate, and human health. We considered cases where decades-long multinational cross-border endangered species recovery programs have been jeopardized by waivers of environmental protection laws to facilitate rapid construction of border barriers that impede the movement and migration of animals, such as at the US-Mexico and Poland-Belarus borders. Renewable energy megaprojects, such as the Pinacate solar plant in Mexico, coupled with power transmission lines and road networks likewise cast a large footprint on the land and are being carried out with minimal to no environmental compliance under the guise of national security. National sovereignty likewise has been used as justification for bypassing laws to proceed with similar projects, such as Mexico's Dos Bocas refinery and Poland's Vistula Spit canal. Emphasis on security is also apparent in increasing military expenditure by the world's largest economies, which has created a mismatch with improvement in environmental policy stringency. Decisions to prioritize security can undermine democratic principles and environmental review protocols, trivialize humanity's dependence on functioning ecosystems, and contradict the United Nation's resolution on the human right to a healthy environment. Framing infrastructure projects as matters of national security also foments civil and political unrest by the labeling and casting of dissenters, including conservation scientists and environmental defenders, as antinational. World leaders must refrain from misusing extraordinary powers, adhere to laws and international agreements, and consult experts and local people before taking unilateral action on projects that affect ecological and human communities.

Amenazas para la conservación a partir de los intereses nacionales de seguridad Resumen En los países existe una tendencia creciente por invocar la seguridad nacional y las declaraciones de emergencia para construir infraestructuras financiadas por el estado para la defensa de las fronteras, producción de energía y transporte. Las leyes, regulaciones y acuerdos establecidos para la protección de la naturaleza y el patrimonio cultural dentro y entre los países se están relegando por la seguridad nacional, lo que compromete la función de las áreas protegidas (parques nacionales, áreas silvestres y reservas de la biósfera) que resguardan la biodiversidad, el clima y la salud humana. Consideramos los casos en donde se han puesto en peligro los programas longevos y multinacionales de recuperación de especies en peligro por las exenciones a las leyes de protección ambiental para facilitar la construcción rápida de barreras fronterizas que impiden el movimiento y la migración de animales, como es el caso de las fronteras entre EU y México y Polonia y Bielorrusia. Los megaproyectos de energía renovable, como la planta solar del Pinacate en México, en conjunto con las líneas de transmisión eléctrica y las redes de carreteras también dejan una gran huella sobre la tierra y se realizan con el mínimo o ningún cumplimiento bajo el aspecto de la seguridad nacional. La soberanía nacional también se ha usado para justificar la omisión de las leyes para proceder con proyectos similares, como la refinería de Dos Bocas en México y el canal Vistula Spit en Polonia. El énfasis sobre la seguridad también es evidente con el incremento del gasto militar de las mayores economías mundiales, lo que ha creado una desigualdad con las mejoras en la exigencia de la política ambiental. Las decisiones para priorizar la seguridad pueden debilitar los principios democráticos y los protocolos de revisión ambiental, banalizar la dependencia de la humanidad por los ecosistemas funcionales y contradecir la resolución de las Naciones Unidas sobre el derecho humano a un ambiente saludable. Cuando se denominan los proyectos de infraestructura como asuntos de seguridad nacional, también se fomenta el malestar civil y político al etiquetar como antinacionales a los disidentes, incluidos los defensores ambientales y los científicos de la conservación. Los líderes mundiales deben abstenerse de usar indebidamente los poderes extraordinarios, adherirse a las leyes y acuerdos internacionales y consultar con expertos y personas locales antes de actuar de forma unilateral en cuanto a proyectos que afectan las comunidades humanas y ecológicas.

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.

Metagenomic Analysis of the Composition of Microbial Consortia Involved in Spruce Degradation over Time in Bialowieza Natural Forest.

Deadwood plays an important role in forest ecology; its degradation and, therefore, carbon assimilation is carried out by fungi and bacteria. To quantify the abundance and distribution of microbial taxa inhabiting dead spruce logs fallen over a span of 50 years and the soil beneath, we used taxonomic profiling with NGS sequencing of hypervariable DNA fragments of ITS1 and 16S V3-V4, respectively. The analysis of sequencing data revealed a high level of diversity in microbial communities participating in the degradation of spruce logs. Differences in the relative abundance of microbial taxa between the samples of the wood that died in 1974 and 2014, and of the soil in its immediate vicinity, were visible, especially at the genus level. Based on the Lefse analysis significantly higher numbers of classified bacterial taxa were observed in the wood and soil samples from 2014 (wood: 1974-18 and 2014-28 taxa; soil: 1974-8 and 2014-41 taxa) while the number of classified fungal taxa was significantly higher in the wood and soil samples from 1974 (wood: 1974-17 and 2014-9 taxa; soil: 1974-57 and 2014-28 taxa). Most of the bacterial and fungal amplicon sequence variants (ASVs) unique to wood were found in the samples from 1974, while those unique to soil were detected in the samples from 2014. The ATR-FTIR method supported by CHN analysis revealed physicochemical changes in deadwood induced by the activity of fungal and bacterial organisms.

Common seed dispersers contribute most to the persistence of a fleshy-fruited tree.

Mutualistic interactions are by definition beneficial for each contributing partner. However, it is insufficiently understood how mutualistic interactions influence partners throughout their lives. Here, we used animal species-explicit, microhabitat-structured integral projection models to quantify the effect of seed dispersal by 20 animal species on the full life cycle of the tree Frangula alnus in Bialowieza Forest, Eastern Poland. Our analysis showed that animal seed dispersal increased population growth by 2.5%. The effectiveness of animals as seed dispersers was strongly related to the interaction frequency but not the quality of seed dispersal. Consequently, the projected population decline due to simulated species extinction was driven by the loss of common rather than rare mutualist species. Our results support the notion that frequently interacting mutualists contribute most to the persistence of the populations of their partners, underscoring the role of common species for ecosystem functioning and nature conservation.

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.

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.

Co-limitation towards lower latitudes shapes global forest diversity gradients.

The latitudinal diversity gradient (LDG) is one of the most recognized global patterns of species richness exhibited across a wide range of taxa. Numerous hypotheses have been proposed in the past two centuries to explain LDG, but rigorous tests of the drivers of LDGs have been limited by a lack of high-quality global species richness data. Here we produce a high-resolution (0.025° × 0.025°) map of local tree species richness using a global forest inventory database with individual tree information and local biophysical characteristics from ~1.3 million sample plots. We then quantify drivers of local tree species richness patterns across latitudes. Generally, annual mean temperature was a dominant predictor of tree species richness, which is most consistent with the metabolic theory of biodiversity (MTB). However, MTB underestimated LDG in the tropics, where high species richness was also moderated by topographic, soil and anthropogenic factors operating at local scales. Given that local landscape variables operate synergistically with bioclimatic factors in shaping the global LDG pattern, we suggest that MTB be extended to account for co-limitation by subordinate drivers.

Forest degradation limits the complementarity and quality of animal seed dispersal.

Forest degradation changes the structural heterogeneity of forests and species communities, with potential consequences for ecosystem functions including seed dispersal by frugivorous animals. While the quantity of seed dispersal may be robust towards forest degradation, changes in the effectiveness of seed dispersal through qualitative changes are poorly understood. Here, we carried out extensive field sampling on the structure of forest microhabitats, seed deposition sites and plant recruitment along three characteristics of forest microhabitats (canopy cover, ground vegetation and deadwood) in Europe's last lowland primeval forest (Bialowieza, Poland). We then applied niche modelling to study forest degradation effects on multi-dimensional seed deposition by frugivores and recruitment of fleshy-fruited plants. Forest degradation was shown to (i) reduce the niche volume of forest microhabitat characteristics by half, (ii) homogenize the spatial seed deposition within and among frugivore species, and (iii) limit the regeneration of plants via changes in seed deposition and recruitment. Our study shows that the loss of frugivores in degraded forests is accompanied by a reduction in the complementarity and quality of seed dispersal by remaining frugivores. By contrast, structure-rich habitats, such as old-growth forests, safeguard the diversity of species interactions, forming the basis for high-quality ecosystem functions.

Climatic conditions, not above- and belowground resource availability and uptake capacity, mediate tree diversity effects on productivity and stability.

Tree species diversity promotes multiple ecosystem functions and services. However, little is known about how above- and belowground resource availability (light, nutrients, and water) and resource uptake capacity mediate tree species diversity effects on aboveground wood productivity and temporal stability of productivity in European forests and whether the effects differ between humid and arid regions. We used the data from six major European forest types along a latitudinal gradient to address those two questions. We found that neither leaf area index (a proxy for light uptake capacity), nor fine root biomass (a proxy for soil nutrient and water uptake capacity) was related to tree species richness. Leaf area index did, however, enhance productivity, but negatively affected stability. Productivity was further promoted by soil nutrient availability, while stability was enhanced by fine root biomass. We only found a positive effect of tree species richness on productivity in arid regions and a positive effect on stability in humid regions. This indicates a possible disconnection between productivity and stability regarding tree species richness effects. In other words, the mechanisms that drive the positive effects of tree species richness on productivity do not per se benefit stability simultaneously. Our findings therefore suggest that tree species richness effects are largely mediated by differences in climatic conditions rather than by differences in above- and belowground resource availability and uptake capacity at the regional scales.

Directional turnover towards larger-ranged plants over time and across habitats.

Species turnover is ubiquitous. However, it remains unknown whether certain types of species are consistently gained or lost across different habitats. Here, we analysed the trajectories of 1827 plant species over time intervals of up to 78 years at 141 sites across mountain summits, forests, and lowland grasslands in Europe. We found, albeit with relatively small effect sizes, displacements of smaller- by larger-ranged species across habitats. Communities shifted in parallel towards more nutrient-demanding species, with species from nutrient-rich habitats having larger ranges. Because these species are typically strong competitors, declines of smaller-ranged species could reflect not only abiotic drivers of global change, but also biotic pressure from increased competition. The ubiquitous component of turnover based on species range size we found here may partially reconcile findings of no net loss in local diversity with global species loss, and link community-scale turnover to macroecological processes such as biotic homogenisation.

Forest floor plant diversity drives the use of mature spruce forests by European bison.

The distribution of large ungulates in space is in large extent driven by the availability of forage, which in temperate forests depends on light availability, and associated plant diversity and cover. We hypothesized that the increased number of GPS fixes of European bison (Bison bonasus L.) in usually avoided spruce forests was an effect of higher plant species richness and cover of the forest floor, which developed owing to increased light availability enhanced by spruce mortality. We carried out 80 forest floor plant surveys combined with tree measurement on plots chosen according to the number of GPS locations of GPS-collared European bison. The mean plant species richness per plot was higher on intensively visited plots (IV) than rarely visited (RV) plots (30 ± 5.75 (SD) versus. 26 ± 6.19 (SD)). The frequency of 34 plant species was higher on IV plots, and they were mainly herbaceous species (32 species), while a significant part of 13 species with higher frequency on RV plots was woody plants (5 species). The species richness of forbs was higher on IV plots, while other functional groups of plants did not differ. Tree stem density on the IV plots was lower than on the RV plots (17.94 ± 6.73 (SD) versus 22.9 ± 7.67 (SD)), and the mean value of Ellenberg's ecological indicator for light availability for all forest floor plant species was higher on IV plots. European bison visiting mature spruce forests was driven by higher forest floor plant cover and species richness, and high share and species richness of forbs. The two latter features may be translated into higher quality and diversity of forage. In spite of morphological characteristics suggesting that European bison is a species of mixed (mosaic) habitats, it seems to be well adapted to thrive in diverse forests.

Response to Comment on "Forest microclimate dynamics drive plant responses to warming".

Schall and Heinrichs question our interpretation that the climatic debt in understory plant communities is locally modulated by canopy buffering. However, our results clearly show that the discrepancy between microclimate warming rates and thermophilization rates is highest in forests where canopy cover was reduced, which suggests that the need for communities to respond to warming is highest in those forests.

Response to Comment on "Forest microclimate dynamics drive plant responses to warming".

Bertrand et al question our interpretation about warming effects on the thermophilization in forest plant communities and propose an alternative way to analyze climatic debt. We show that microclimate warming is a better predictor than macroclimate warming for studying forest plant community responses to warming. Their additional analyses do not affect or change our interpretations and conclusions.