Decametric-scale buffering of climate extremes in forest understory within a riparian microrefugia: the key role of microtopography.

Riparian corridors often act as low-land climate refugia for temperate tree species in their southern distribution range. A plausible mechanism is the buffering of regional climate extremes by local physiographic and biotic factors. We tested this idea using a 3-year-long microclimate dataset collected along the Ciron river, a refugia for European beech (Fagus sylvatica) in southwestern France. Across the whole network, canopy gap fraction was the main predictor for spatial microclimatic variations, together with two other landscape features (elevation above the river and woodland fraction within a 300m radius). However, within the riparian forest only (canopy gap fraction < 25%, distance to the river < 150m), variations of up to -4°C and + 15% in summertime daily maximum air temperature and minimum relative humidity, respectively, were still found from the plateau to the cooler, moister river banks, only ~ 5-10m below. Elevation above the river was then identified as the main predictor, and explained the marked variations from the plateau to the banks much better than canopy gap fraction. The microclimate measured near the river is as cool but moister than the macroclimate encountered at 700-1000m asl further east in F. sylvatica's main distribution range. Indeed, at all locations, we found that air relative humidity was higher than expected from a temperature-only effect, suggesting that extra moisture is brought by the river. Our results explain well why beech trees in this climate refugium are restricted to the river gorges where microtopographic variations are the strongest and canopy gaps are rare.

The role of ungulates in nowadays temperate forests. A response to Fløjgaard et al. (DOI:10.1111/gcb.14029).

In Boulanger et al. (2018), we investigated the effects of ungulates on forest plant diversity. By suggesting a revisit of our conclusions regarding ecosystem dynamics since the late Pleistocene, Fløjgaard et al. (2018) came to the conclusion that moderate grazing in forest should be a conservation target. Since major points of our paper were mis- or over- interpreted, we put the record straight on our study system and on the scope of our conclusions. Finally, we advocate for an assessment of the conservation issues of ungulates in forests not only regarding hypothetical and still debated states of past ecosystems but also considering timely challenges for forest ecosystems.

Ungulates increase forest plant species richness to the benefit of non-forest specialists.

Large wild ungulates are a major biotic factor shaping plant communities. They influence species abundance and occurrence directly by herbivory and plant dispersal, or indirectly by modifying plant-plant interactions and through soil disturbance. In forest ecosystems, researchers' attention has been mainly focused on deer overabundance. Far less is known about the effects on understory plant dynamics and diversity of wild ungulates where their abundance is maintained at lower levels to mitigate impacts on tree regeneration. We used vegetation data collected over 10 years on 82 pairs of exclosure (excluding ungulates) and control plots located in a nation-wide forest monitoring network (Renecofor). We report the effects of ungulate exclusion on (i) plant species richness and ecological characteristics, (ii) and cover percentage of herbaceous and shrub layers. We also analyzed the response of these variables along gradients of ungulate abundance, based on hunting statistics, for wild boar (Sus scrofa), red deer (Cervus elaphus) and roe deer (Capreolus capreolus). Outside the exclosures, forest ungulates maintained higher species richness in the herbaceous layer (+15%), while the shrub layer was 17% less rich, and the plant communities became more light-demanding. Inside the exclosures, shrub cover increased, often to the benefit of bramble (Rubus fruticosus agg.). Ungulates tend to favour ruderal, hemerobic, epizoochorous and non-forest species. Among plots, the magnitude of vegetation changes was proportional to deer abundance. We conclude that ungulates, through the control of the shrub layer, indirectly increase herbaceous plant species richness by increasing light reaching the ground. However, this increase is detrimental to the peculiarity of forest plant communities and contributes to a landscape-level biotic homogenization. Even at population density levels considered to be harmless for overall plant species richness, ungulates remain a conservation issue for plant community composition.

Soil phosphorus constrains biodiversity across European grasslands.

Nutrient pollution presents a serious threat to biodiversity conservation. In terrestrial ecosystems, the deleterious effects of nitrogen pollution are increasingly understood and several mitigating environmental policies have been developed. Compared to nitrogen, the effects of increased phosphorus have received far less attention, although some studies have indicated that phosphorus pollution may be detrimental for biodiversity as well. On the basis of a dataset covering 501 grassland plots throughout Europe, we demonstrate that, independent of the level of atmospheric nitrogen deposition and soil acidity, plant species richness was consistently negatively related to soil phosphorus. We also identified thresholds in soil phosphorus above which biodiversity appears to remain at a constant low level. Our results indicate that nutrient management policies biased toward reducing nitrogen pollution will fail to preserve biodiversity. As soil phosphorus is known to be extremely persistent and we found no evidence for a critical threshold below which no environmental harm is expected, we suggest that agro-environmental schemes should include grasslands that are permanently free from phosphorus fertilization.

Bird predation enhances tree seedling resistance to insect herbivores in contrasting forest habitats.

According to the associational resistance hypothesis, neighbouring plants are expected to influence both the insect herbivore communities and their natural enemies. However, this has rarely been tested for the effects of canopy trees on herbivory of seedlings. One possible mechanism responsible for associational resistance is the indirect impact of natural enemies on insect herbivory, such as insectivorous birds. But it remains unclear to what extent such trophic cascades are influenced by the composition of plant associations (i.e. identity of 'associated' plants). Here, we compared the effect of bird exclusion on insect leaf damage for seedlings of three broadleaved tree species in three different forest habitats. Exclusion of insectivorous birds affected insect herbivory in a species-specific manner: leaf damage increased on Betula pendula seedlings whereas bird exclusion had no effect for two oaks (Quercus robur and Q. ilex). Forest habitat influenced both the extent of insect herbivory and the effect of bird exclusion. Broadleaved seedlings had lower overall leaf damage within pine plantations than within broadleaved stands, consistent with the resource concentration hypothesis. The indirect effect of bird exclusion on leaf damage was only significant in pine plantations, but not in exotic and native broadleaved woodlands. Our results support the enemies hypothesis, which predicts that the effects of insectivorous birds on insect herbivory on seedlings are greater beneath non-congeneric canopy trees. Although bird species richness and abundance were greater in broadleaved woodlands, birds were unable to regulate insect herbivory on seedlings in forests of more closely related tree species.

The effects of management on vegetation trajectories during the early-stage restoration of previously arable land after hay transfer.

The restoration of floodplain grasslands has benefited from many studies of the underlying mechanisms. Among the operational tools that resulted, hay transfer is now used increasingly to alleviate the effects of limited seed dispersal and recruitment. To improve this method, we still need to understand how it can affect restoration trajectories, and particularly their direction and magnitude during the early stages of restoration. Based on concepts from the field of community ecology theory, we investigated the effects of early-stage management through grazing or mowing on restoration trajectories after soil harrowing and hay transfer. We established a randomized block design experiment and quantified several community-related metrics to formalize restoration trajectories for 3 years after hay transfer on a previously arable alluvial island in southwestern France. Whatever the management treatment, the species richness and evenness were significantly higher in hay-inoculated than in control plots. This effect was linked to the recruitment of species originating not only from the reference grassland through hay transfer, but also from the seed bank, a well-known effect of soil harrowing. Although generally oriented toward the reference grassland, the origin, direction, and magnitude of the trajectory of hay-inoculated plots all depended on the management applied. Sheep grazing applied at the same time as hay transfer enhanced the recruitment of reference species as from the first experimental year, because it controlled aboveground competition and maintained the window of opportunity open for a sufficiently longer period of time. Our findings show that the type of management applied simultaneously to hay transfer influences the origin of a grassland trajectory, while its direction and magnitude are dependent on the management applied in subsequent years. Grazing immediately after hay transfer may be appropriate to accelerate the recruitment of species from the reference grassland.

[Beech (Fagus sylvatica) germination and seedling growth under climatic and allelopathic constraints]. / Germination et croissance des plantules de hêtre (Fagus sylvatica) sous contraintes climatiques et allélopathiques.

Recruitment is a key process for forest sustainability, especially in warm margin of distribution area. The influence of climate (temperate or warm), of soil water availability, and of allelopathic interactions from different forest species have been tested on the germination of Fagus sylvatica in controlled climatic conditions. Germination rates of non-dormant Fagus seeds were improved by relatively warm temperatures (20°C), but reversibly stopped under heat constraint (27°C). The relative growth rate of Fagus seedlings was better under temperate climatic conditions. Foliar extracts of Hedera helix showed the highest allelopathic effect on Fagus recruitment, especially in temperate conditions. Our results suggest a limitation of Fagus recruitment in warm margin of its distribution area, and a modulation of recruitment success according to the identity of plant neighbourhood.

Modelling the impact of climate change and atmospheric N deposition on French forests biodiversity.

A dynamic coupled biogeochemical-ecological model was used to simulate the effects of nitrogen deposition and climate change on plant communities at three forest sites in France. The three sites had different forest covers (sessile oak, Norway spruce and silver fir), three nitrogen loads ranging from relatively low to high, different climatic regions and different soil types. Both the availability of vegetation time series and the environmental niches of the understory species allowed to evaluate the model for predicting the composition of the three plant communities. The calibration of the environmental niches was successful, with a model performance consistently reasonably high throughout the three sites. The model simulations of two climatic and two deposition scenarios showed that climate change may entirely compromise the eventual recovery from eutrophication of the simulated plant communities in response to the reductions in nitrogen deposition. The interplay between climate and deposition was strongly governed by site characteristics and histories in the long term, while forest management remained the main driver of change in the short term.

[Selectivity and environmental variations in herbivory by Orthoptera]. / Sélectivité et variations environnementales de l'herbivorie par les Orthoptères.

The aim of this study is 1 degree) to quantify environmental changes in herbivory due to Orthoptera on two perennial grasses and 2 degrees) to assess the processes involved in the control of herbivory. Herbivory varies strongly according to shade, drought and mowing, and is positively related to vapour pressure deficit and temperature. Besides the hypothesis of a trophic control of herbivory, our results are consistent with a microclimatic control of herbivory by Orthoptera. The coexistence of different hypothesis of herbivory control may depend on the studied system and specifically on the type of herbivore involved.

The impact of nitrogen deposition on acid grasslands in the Atlantic region of Europe.

A survey of 153 acid grasslands from the Atlantic biogeographic region of Europe indicates that chronic nitrogen deposition is changing plant species composition and soil and plant-tissue chemistry. Across the deposition gradient (2-44 kg N ha(-1) yr(-1)) grass richness as a proportion of total species richness increased whereas forb richness decreased. Soil C:N ratio increased, but soil extractable nitrate and ammonium concentrations did not show any relationship with nitrogen deposition. The above-ground tissue nitrogen contents of three plant species were examined: Agrostis capillaris (grass), Galium saxatile (forb) and Rhytidiadelphus squarrosus (bryophyte). The tissue nitrogen content of neither vascular plant species showed any relationship with nitrogen deposition, but there was a weak positive relationship between R. squarrosus nitrogen content and nitrogen deposition. None of the species showed strong relationships between above-ground tissue N:P or C:N and nitrogen deposition, indicating that they are not good indicators of deposition rate.

Nitrogen deposition threatens species richness of grasslands across Europe.

Evidence from an international survey in the Atlantic biogeographic region of Europe indicates that chronic nitrogen deposition is reducing plant species richness in acid grasslands. Across the deposition gradient in this region (2-44 kg N ha(-1) yr(-1)) species richness showed a curvilinear response, with greatest reductions in species richness when deposition increased from low levels. This has important implications for conservation policies, suggesting that to protect the most sensitive grasslands resources should be focussed where deposition is currently low. Soil pH is also an important driver of species richness indicating that the acidifying effect of nitrogen deposition may be contributing to species richness reductions. The results of this survey suggest that the impacts of nitrogen deposition can be observed over a large geographical range.

Respective influence of habitat conditions and management regimes on prealpine calcareous grasslands.

The calcareous grasslands of the south-western French Alps have been poorly studied, although they provide suitable habitat for rare plant species and communities. The separate and combined effects on calcareous grassland communities of habitat conditions (lithology, soil moisture) and management regimes (grazing intensity, cutting regime) were studied using constrained ordination techniques (canonical correspondence analysis with variance partitioning). Among the explanatory variables considered, the most important factor determining floristic composition was lithology, which explained 11.9% of floristic variability, followed by grazing intensity (6.0%). Additive effects of management and lithology explained 23.9% of floristic variability. Species niche amplitude was measured by conditional variances of samples along main ordination axes, in order to define adequate conservation management for the rarest short-lived species with narrow niche breadth on both habitat and management gradients.