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.

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 understorey communities respond strongly to light in interaction with forest structure, but not to microclimate warming.

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.

Soil seed bank responses to edge effects in temperate European 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.

ForestTemp - Sub-canopy microclimate temperatures of European forests.

Ecological research heavily relies on coarse-gridded climate data based on standardized temperature measurements recorded at 2 m height in open landscapes. However, many organisms experience environmental conditions that differ substantially from those captured by these macroclimatic (i.e. free air) temperature grids. In forests, the tree canopy functions as a thermal insulator and buffers sub-canopy microclimatic conditions, thereby affecting biological and ecological processes. To improve the assessment of climatic conditions and climate-change-related impacts on forest-floor biodiversity and functioning, high-resolution temperature grids reflecting forest microclimates are thus urgently needed. Combining more than 1200 time series of in situ near-surface forest temperature with topographical, biological and macroclimatic variables in a machine learning model, we predicted the mean monthly offset between sub-canopy temperature at 15 cm above the surface and free-air temperature over the period 2000-2020 at a spatial resolution of 25 m across Europe. This offset was used to evaluate the difference between microclimate and macroclimate across space and seasons and finally enabled us to calculate mean annual and monthly temperatures for European forest understories. We found that sub-canopy air temperatures differ substantially from free-air temperatures, being on average 2.1°C (standard deviation ± 1.6°C) lower in summer and 2.0°C higher (±0.7°C) in winter across Europe. Additionally, our high-resolution maps expose considerable microclimatic variation within landscapes, not captured by the gridded macroclimatic products. The provided forest sub-canopy temperature maps will enable future research to model below-canopy biological processes and patterns, as well as species distributions more accurately.

Seasonal drivers of understorey temperature buffering in temperate deciduous forests across Europe.

AIM: Forest understorey microclimates are often buffered against extreme heat or cold, with important implications for the organisms living in these environments. We quantified seasonal effects of understorey microclimate predictors describing canopy structure, canopy composition and topography (i.e., local factors) and the forest patch size and distance to the coast (i.e., landscape factors). LOCATION: Temperate forests in Europe. TIME PERIOD: 2017-2018. MAJOR TAXA STUDIED: Woody plants. METHODS: We combined data from a microclimate sensor network with weather-station records to calculate the difference, or offset, between temperatures measured inside and outside forests. We used regression analysis to study the effects of local and landscape factors on the seasonal offset of minimum, mean and maximum temperatures. RESULTS: The maximum temperature during the summer was on average cooler by 2.1 °C inside than outside forests, and the minimum temperatures during the winter and spring were 0.4 and 0.9 °C warmer. The local canopy cover was a strong nonlinear driver of the maximum temperature offset during summer, and we found increased cooling beneath tree species that cast the deepest shade. Seasonal offsets of minimum temperature were mainly regulated by landscape and topographic features, such as the distance to the coast and topographic position. MAIN CONCLUSIONS: Forest organisms experience less severe temperature extremes than suggested by currently available macroclimate data; therefore, climate-species relationships and the responses of species to anthropogenic global warming cannot be modelled accurately in forests using macroclimate data alone. Changes in canopy cover and composition will strongly modulate the warming of maximum temperatures in forest understories, with important implications for understanding the responses of forest biodiversity and functioning to the combined threats of land-use change and climate change. Our predictive models are generally applicable across lowland temperate deciduous forests, providing ecologically important microclimate data for forest understories.

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.

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.

Multi-scale approach to biodiversity proxies of biological control service in European farmlands.

Intensive agriculture has profoundly altered biodiversity and trophic relationships in agricultural landscapes, leading to the deterioration of many ecosystem services such as pollination or biological control. Information on which spatio-temporal factors are simultaneously affecting crop pests and their natural enemies is required to improve conservation biological control practices. We conducted a study in 80 winter wheat crop fields distributed in three regions of North-western Europe (Brittany, Hauts-de-France and Wallonia), along intra-regional gradients of landscape complexity. Five taxa of major crop pests (aphids and slugs) and natural enemies (spiders, carabids, and parasitoids) were sampled three times a year, for two consecutive years. We analysed the influence of regional (meteorology), landscape (structure in both the years n and n-1) and local factors (hedge or grass strip field boundaries, and distance to boundary) on the abundance and species richness of crop-dwelling organisms, as proxies of the service/disservice they provide. Firstly, there was higher biocontrol potential in areas with mild winter climatic conditions. Secondly, natural enemy communities were less diverse and had lower abundances in landscapes with high crop and wooded continuities (sum of interconnected crop or wood surfaces), contrary to slugs and aphids. Finally, field boundaries with grass strips were more favourable to spiders and carabids than boundaries formed by hedges, while the opposite was found for crop pests, with the latter being less abundant towards the centre of the fields. We also revealed temporal modulation-and sometimes reversion-of the impact of local elements on crop biodiversity. To some extent, these results cause controversy because they show that hedgerows and woodlots should not be the unique cornerstones of agro-ecological landscape design strategies. We point out that combining woody and grassy habitats to take full advantage of the features and ecosystem services they both provide (biological pest control, windbreak effect, soil stabilization) may promote sustainable agricultural ecosystems. It may be possible to both reduce pest pressure and promote natural enemies by accounting for taxa-specific antagonistic responses to multi-scale environmental characteristics.

Estimating crop parameters using Sentinel-1 and 2 datasets and geospatial field data.

Crop monitoring is essential for ensuring food security in a global context of population growth and climate change. Satellite images are commonly used to estimate crop parameters over large areas, and the freely available Synthetic Aperture Radar (SAR) Sentinel-1 (S-1) and optical Sentinel-2 (S-2) images are relevant for that purpose combining high temporal resolution and high spatial resolution. For this data article, field surveys were conducted from January to July 2017 in France to sample wheat and rapeseed crop parameters during the entire crops cycle. Phenological stages were identified in 83 wheat fields and 32 rapeseed fields in Brittany and Picardy regions. Moreover, Leaf Area Index (LAI), wet biomass, dry biomass and water content were sampled in three wheat fields and three rapeseed fields in Brittany. We assigned to each field sample 10 spectral bands and 12 vegetation indices from S-2 images and two backscattering coefficients, one backscattering ratio and four polarimetric indicators from S-1 images. This dataset can be used for crop monitoring in other regions, as well as for modelling development.

Drivers of carbon stocks in forest edges across Europe.

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

Does nitrogen fertilization history affects short-term microbial responses and chemical properties of soils submitted to different glyphosate concentrations?

The use of nitrogen (N) fertilizer and glyphosate-based herbicides is increasing worldwide, with agriculture holding the largest market share. The agronomic and socioeconomic utilities of glyphosate are well established; however, our knowledge of the potential effects of glyphosate applied in the presence or absence of long-term N fertilization on microbial functional activities and the availability of soil nutrients remains limited. Using an ex situ approach with soils that did (N+) or did not (N0) receive synthetic N fertilization for 6 years, we assessed the impact of different rates (no glyphosate, CK; field rate, FR; 100 × field rate, 100FR) of glyphosate application on biological and chemical parameters. We observed that, after immediate application (1 day), the highest dose of glyphosate (100FR) negatively affected the alkaline phosphatase (AlP) activity in soils without N fertilization history and decreased the cation exchange capacity (CEC) in N0 compared to CK and FR treatments with N+. Conversely, the 100FR application increased nitrate (NO3-) and available phosphorus (PO43-) regardless of N fertilization history. Then, after 8 and 15 days, the N+\100FR and N+\FR treatments exhibited the lowest values for dehydrogenase (DH) and AlP activities, respectively, while urease (URE) activity was mainly affected by N fertilization. After 15 days and irrespective of N fertilization history, the FR glyphosate application negatively affected the degradation of carbon substrates by microbial communities (expressed as the average well color development, AWCD). By contrast, the 100FR treatment positively affected AWCD, increasing PO43- by 5 and 16% and NO3- by 126 and 119% in the N+ and N0 treatments, respectively. In addition, the 100FR treatment resulted in an increase in the average net nitrification rate. Principal component analysis revealed that the 100FR glyphosate treatment selected microbial communities that were able to metabolize amine substrates. Overall, the lack of N fertilization in the 6 past years combined with the highest glyphosate application rate (100FR) induced the highest values of AWCD, functional diversity, NO3-, PO43- and nitrification. We concluded that the intensive use of N fertilization for 6 years may change the non-target effects of glyphosate application on enzyme activities. The functional activities, nitrification and nutrient contents were increased by glyphosate only when applied at 100 times the field application rate.

Response of bacterial communities to Pb smelter pollution in contrasting soils.

Anthropogenic inputs of trace elements (TE) into soils constitute a major public and environmental health problem. Bioavailability of TE is strongly related to the soil physicochemical parameters and thus to the ecosystem type. In order to test whether soil parameters influence the response of the bacterial community to TE pollution, we collected soil samples across contrasting ecosystems (hardwood, coniferous and hydromorphic soils), which have been contaminated in TE and especially lead (Pb) over several decades due to nearby industrial smelting activities. Bacterial community composition was analysed using high throughput amplicon sequencing and compared to the soil physicochemical parameters. Multivariate analyses of the pedological and biological data revealed that the bacterial community composition was affected by ecosystem type in the first place. An influence of the contamination level was also evidenced within each ecosystem. Despite the important variability in bacterial community structure, we found that specific bacterial groups such as γ-Proteobacteria, Verrucomicrobia and Chlamydiae showed a consistent response to Pb content across contrasting ecosystems. Verrucomicrobia were less abundant at high contamination level whereas Chlamydiae and γ-Proteobacteria were more abundant. We conclude that such groups and ratio's thereof can be considered as relevant bioindicators of Pb contamination.

Effects of ozone on stomatal responses to environmental parameters (blue light, red light, CO2 and vapour pressure deficit) in three Populus deltoides × Populus nigra genotypes.

The effect of ozone (O(3)) on stomatal regulation was studied in three Euramerican poplar genotypes (Populus deltoides × Populus nigra: Carpaccio, Cima and Robusta). The impact of O(3) on stomatal conductance responses to variations in blue light, red light, CO(2) concentration and vapour pressure deficit (VPD) was studied. Upon O(3) exposure, a sluggish response of stomatal movements was observed, characterized by slower reactions to increases in blue light intensity, CO(2) concentration and VPD, and lower amplitude of the response to variations in light intensity. That sluggish response should be taken into account in stomatal conductance models for phytotoxic ozone dose (POD(Y)) calculations. The speed of the response to variations in environmental parameters appears as a determining factor of genotype-related sensitivity.