Extinction drives recent thermophilization but does not trigger homogenization in forest understorey.

The ongoing climate change is triggering plant community thermophilization. This selection process ought to shift community composition towards species adapted to warmer climates but may also lead to biotic homogenization. The link between thermophilization and homogenization and the community dynamics that drive them (colonization and extinction) remain unknown but is critical for understanding community responses under rapid environmental change. We used 14,167 pairs of plots to study shifts in plant community during 10 years of rising temperature in 80 forest ecoregions of France. We computed community mean thermal optimum (thermophilization) and Δß-diversity (homogenization) for each ecoregion and partitioned these changes into extinction and colonization dynamics of cold- and warm-adapted species. Forest understorey communities thermophilized on average by 0.12 °C per decade and up to 0.20 °C per decade in warm ecoregions. This rate was entirely driven by extinction dynamics. Extinction of cold-adapted species was a driver of homogenization but it was compensated for by the colonization of rare species and the extinction of common species, resulting in the absence of an apparent homogenization trend. Here we show a dieback of present cold-adapted species rather than an adaptation of communities via the arrival of warm-adapted species, with a mutually cancelling effect on ß-diversity. These results suggest that a future loss of biodiversity and delayed biotic homogenization should be considered.

Background mortality drivers of European tree species: climate change matters.

Increases in tree mortality rates have been highlighted in different biomes over the past decades. However, disentangling the effects of climate change on the temporal increase in tree mortality from those of management and forest dynamics remains a challenge. Using a modelling approach taking tree and stand characteristics into account, we sought to evaluate the impact of climate change on background mortality for the most common European tree species. We focused on background mortality, which is the mortality observed in a stand in the absence of abrupt disturbances, to avoid confusion with mortality events unrelated to long-term changes in temperature and rainfall. We studied 372 974 trees including 7312 dead trees from forest inventory data surveyed across France between 2009 and 2015. Factors related to competition, stand characteristics, management intensity, and site conditions were the expected preponderant drivers of mortality. Taking these main drivers into account, we detected a climate change signal on 45% of the 43 studied species, explaining an average 6% of the total modelled mortality. For 18 out of the 19 species sensitive to climate change, we evidenced greater mortality with increasing temperature or decreasing rainfall. By quantifying the mortality excess linked to the current climate change for European temperate forest tree species, we provide new insights into forest vulnerability that will prove useful for adapting forest management to future conditions.

History and environment shape species pools and community diversity in European beech forests.

A central hypothesis of ecology states that regional diversity influences local diversity through species-pool effects. Species pools are supposedly shaped by large-scale factors and then filtered into ecological communities, but understanding these processes requires the analysis of large datasets across several regions. Here, we use a framework of community assembly at a continental scale to test the relative influence of historical and environmental drivers, in combination with regional or local species pools, on community species richness and community completeness. Using 42,173 vegetation plots sampled across European beech forests, we found that large-scale factors largely accounted for species pool sizes. At the regional scale, main predictors reflected historical contingencies related to post-glacial dispersal routes, whereas at the local scale, the influence of environmental filters was predominant. Proximity to Quaternary refugia and high precipitation were the main factors supporting community species richness, especially among beech forest specialist plants. Models for community completeness indicate the influence of large-scale factors, further suggesting community saturation as a result of dispersal limitation or biotic interactions. Our results empirically demonstrate how historical factors complement environmental gradients to provide a better understanding of biodiversity patterns across multiple regions.

Ecological constraints increase the climatic debt in forests.

Biodiversity changes are lagging behind current climate warming. The underlying determinants of this climatic debt are unknown and yet critical to understand the impacts of climate change on the present biota and improve forecasts of biodiversity changes. Here we assess determinants of climatic debt accumulated in French forest herbaceous plant communities between 1987 and 2008 (that is, a 1.05 °C mean difference between the observed and bioindicated temperatures). We show that warmer baseline conditions predispose plant communities to larger climatic debts, and that climate warming exacerbates this response. Forest plant communities, however, are absorbing part of the temperature increase mainly through the species' ability to tolerate changing climate. As climate warming is expected to accelerate during the twenty-first century, plant migration and tolerance to climatic stresses probably will be insufficient to absorb this impact posing threats to the sustainability of forest plant communities.

Early signs of range disjunction of submountainous plant species: an unexplored consequence of future and contemporary climate changes.

Poleward and upward species range shifts are the most commonly anticipated and studied consequences of climate warming. However, these global responses to climate change obscure more complex distribution change patterns. We hypothesize that the spatial arrangement of mountain ranges and, consequently, climatic gradients in Europe, will result in range disjunctions. This hypothesis was investigated for submountainous forest plant species at two temporal and spatial scales: (i) under future climate change (between 1950-2000 and 2061-2080 periods) at the European scale and (ii) under contemporary climate change (between 1914-1987 and 1997-2013 periods) at the French scale. We selected 97 submountainous forest plant species occurring in France, among which distribution data across Europe are available for 25 species. By projecting future distribution changes for the 25 submountainous plant species across Europe, we demonstrated that range disjunction is a likely consequence of future climate change. To assess whether it is already taking place, we used a large forest vegetation-plot database covering the entire French territory over 100 years (1914-2013) and found an average decrease in frequency (-0.01 ± 0.004) in lowland areas for the 97 submountainous species - corresponding to a loss of 6% of their historical frequency - along with southward and upward range shifts, suggesting early signs of range disjunctions. Climate-induced range disjunctions should be considered more carefully since they could have dramatic consequences on population genetics and the ability of species to face future climate changes.

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.

Tree cover at fine and coarse spatial grains interacts with shade tolerance to shape plant species distributions across the Alps.

The role of competition for light among plants has long been recognised at local scales, but its importance for plant species distributions at larger spatial scales has generally been ignored. Tree cover modifies the local abiotic conditions below the canopy, notably by reducing light availability, and thus, also the performance of species that are not adapted to low-light conditions. However, this local effect may propagate to coarser spatial grains, by affecting colonisation probabilities and local extinction risks of herbs and shrubs. To assess the effect of tree cover at both the plot- and landscape-grain sizes (approximately 10-m and 1-km), we fit Generalised Linear Models (GLMs) for the plot-level distributions of 960 species of herbs and shrubs using 6,935 vegetation plots across the European Alps. We ran four models with different combinations of variables (climate, soil and tree cover) at both spatial grains for each species. We used partial regressions to evaluate the independent effects of plot- and landscape-grain tree cover on plot-level plant communities. Finally, the effects on species-specific elevational range limits were assessed by simulating a removal experiment comparing the species distributions under high and low tree cover. Accounting for tree cover improved the model performance, with the probability of the presence of shade-tolerant species increasing with increasing tree cover, whereas shade-intolerant species showed the opposite pattern. The tree cover effect occurred consistently at both the plot and landscape spatial grains, albeit most strongly at the former. Importantly, tree cover at the two grain sizes had partially independent effects on plot-level plant communities. With high tree cover, shade-intolerant species exhibited narrower elevational ranges than with low tree cover whereas shade-tolerant species showed wider elevational ranges at both limits. These findings suggest that forecasts of climate-related range shifts for herb and shrub species may be modified by tree cover dynamics.

Light and competition gradients fail to explain the coexistence of shade-tolerant Fagus sylvatica and shade-intermediate Quercus petraea seedlings.

BACKGROUND AND AIMS: The coexistence of forest tree species has often been linked to differences among species in terms of their response to light availability during the regeneration stage. From this perspective, species coexistence results from growth-growth or mortality-growth trade-offs along spatial light gradients. Experimental evidence of growth-growth trade-offs in natural conditions is sparse due to various confounding factors that potentially hinder the relationship. This study examined growth hierarchies along light gradients between two tree species with contrasting shade tolerance by controlling potential confounding factors such as seedling size, seedling status, seedling density and species composition. METHODS: Natural regenerated shade-tolerant Fagus sylvatica and shade-intermediate Quercus petraea seedlings were used, and growth rankings over a 4-year period were compared in 8- to 10-year-old tree seedlings. KEY RESULTS: No rank reversal occurs between the two species along the light gradient, or along the density, mixture or seedling size gradients. The shade-tolerant species was always the more competitive of the two. Pronounced effects of initial size on seedling growth were observed, whereas the effects of light and competition by neighbours were of secondary importance. The paramount effect of size, which results from the asymmetric nature of interseedling competition, gives a strong advantage to tall seedlings over the long term. CONCLUSIONS: This study extends previous efforts to identify potential drivers of rank reversals in young tree mixtures. It does not support the classical assumption that spatial heterogeneity in canopy opening explains the coexistence of the two species studied. It suggests that spatial variation in local size hierarchies among seedlings that may be caused by seedling emergence time or seedling initial performance is the main driver of the dynamics of these mixed stands.

Changes in plant community composition lag behind climate warming in lowland forests.

Climate change is driving latitudinal and altitudinal shifts in species distribution worldwide, leading to novel species assemblages. Lags between these biotic responses and contemporary climate changes have been reported for plants and animals. Theoretically, the magnitude of these lags should be greatest in lowland areas, where the velocity of climate change is expected to be much greater than that in highland areas. We compared temperature trends to temperatures reconstructed from plant assemblages (observed in 76,634 surveys) over a 44-year period in France (1965-2008). Here we report that forest plant communities had responded to 0.54 °C of the effective increase of 1.07 °C in highland areas (500-2,600 m above sea level), while they had responded to only 0.02 °C of the 1.11 °C warming trend in lowland areas. There was a larger temperature lag (by 3.1 times) between the climate and plant community composition in lowland forests than in highland forests. The explanation of such disparity lies in the following properties of lowland, as compared to highland, forests: the higher proportion of species with greater ability for local persistence as the climate warms, the reduced opportunity for short-distance escapes, and the greater habitat fragmentation. Although mountains are currently considered to be among the ecosystems most threatened by climate change (owing to mountaintop extinction), the current inertia of plant communities in lowland forests should also be noted, as it could lead to lowland biotic attrition.

Cross-scale analysis of the region effect on vascular plant species diversity in southern and northern European mountain ranges.

BACKGROUND: The divergent glacial histories of southern and northern Europe affect present-day species diversity at coarse-grained scales in these two regions, but do these effects also penetrate to the more fine-grained scales of local communities? METHODOLOGY/PRINCIPAL FINDINGS: We carried out a cross-scale analysis to address this question for vascular plants in two mountain regions, the Alps in southern Europe and the Scandes in northern Europe, using environmentally paired vegetation plots in the two regions (n = 403 in each region) to quantify four diversity components: (i) total number of species occurring in a region (total γ-diversity), (ii) number of species that could occur in a target plot after environmental filtering (habitat-specific γ-diversity), (iii) pair-wise species compositional turnover between plots (plot-to-plot ß-diversity) and (iv) number of species present per plot (plot α-diversity). We found strong region effects on total γ-diversity, habitat-specific γ-diversity and plot-to-plot ß-diversity, with a greater diversity in the Alps even towards distances smaller than 50 m between plots. In contrast, there was a slightly greater plot α-diversity in the Scandes, but with a tendency towards contrasting region effects on high and low soil-acidity plots. CONCLUSIONS/SIGNIFICANCE: We conclude that there are strong regional differences between coarse-grained (landscape- to regional-scale) diversity components of the flora in the Alps and the Scandes mountain ranges, but that these differences do not necessarily penetrate to the finest-grained (plot-scale) diversity component, at least not on acidic soils. Our findings are consistent with the contrasting regional Quaternary histories, but we also consider alternative explanatory models. Notably, ecological sorting and habitat connectivity may play a role in the unexpected limited or reversed region effect on plot α-diversity, and may also affect the larger-scale diversity components. For instance, plot connectivity and/or selection for high dispersal ability may increase plot α-diversity and compensate for low total γ-diversity.