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Aim Climate warming reshuffles biological assemblages towards less cold‐adapted but more warm‐adapted species, a process coined thermophilization. However, the velocity at which this process is happening generally lags behind the velocity of climate change, generating a climatic debt the temporal dynamics of which remain misunderstood. Relying on high‐resolution time series of vegetation data from a long‐term monitoring network of permanent forest plots, we aim at quantifying the temporal dynamics – up to a yearly resolution – of the climatic debt in the understorey of temperate forests before identifying the key determinants that modulate it. Location France. Time period 1995–2017. Taxa studied Vascular plants. Methods We used the community temperature index (CTI) to produce a time series of understorey plant community thermophilization, which we subsequently compared to a time series of mean annual temperature changes over the same period and for the same sites. The direction and magnitude of the difference (i.e., the climatic debt) was finally analysed using linear mixed‐effect models to assess the relative contributions of abiotic and biotic determinants, including forest stand characteristics. Results We found a significant increase in CTI values over time (0.08–0.09 °C/decade), whereas the velocity of mean annual temperature changes was three times higher over the same period (0.22–0.28 °C/decade). Hence, the climatic debt increased over time and was greater in forest stands with higher basal area or older trees as well as under warmer macroclimate. By contrast, a greater frequency of anthropogenic disturbances decreased the climatic debt, while natural disturbances and herbivory had no impact. Conclusions Although often overlooked in understanding the climatic debt of forest biodiversity, changes in forest stand characteristics may modulate the climatic debt by locally modifying microclimatic conditions. Notably, the buffering effect of the upper canopy layer implies microclimate dynamics that may provide more time for understorey plant communities to locally adapt.

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.

The number of species (species richness) is certainly the most widely used descriptor of plant diversity. However, estimating richness is a difficult task because plant censuses are prone to overlooking and identification errors that may lead to spurious interpretations. We used calibration data from the French ICP-level II plots (RENECOFOR) to assess the magnitude of the two kinds of errors in large forest plots. Eleven teams of professional botanists recorded all plants on the same eight 100-m 2 plots in 2004 (four plots, eights teams) and 2005 (four plots, nine teams including six from 2004), first independently and then consensually. On average, 15.5% of the shrubs and trees above 2 m were overlooked and 2.3% not identified at the species level or misidentified. On average, 19.2% of the plant species below 2 m in Electronic supplementary material The online version of this article (height were overlooked and 5.3% were misidentified and 1.3% were misidentified at the genus level (especially bryophytes). The overlooking rate also varied with plant species, morphological type, plot and team. It was higher when only one botanist made the census. It rapidly decreased with species cover and increased with plot species richness, the recording time of the census in the tree layer and the number of the censuses carried out during the day in the ground layer. Familiarity of the team with the local flora reduced the risk of overlooking and identification errors, whereas training had little impact. Differences in species richness (over space or time) in large plots should be cautiously interpreted, especially when several botanists participate in the survey. In particular, the quality of the data needs to be evaluated using calibration training and, if necessary, may be improved by involving more experienced botanists working in teams and by fixing a minimum recording time.

Depending on its developmental and morphological characteristics, shrubby or herbaceous understorey vegetation interacts differently with tree seedlings during the regeneration process. In acidic temperate forests, three common understorey plant species-Calluna vulgaris (L.) Hull, Pteridium aquilinum (L.) Kuhn in Kersten, Molinia caerulea (L.) Moench-are known to rapidly colonize forest gaps. Therefore, they often develop at the expense of light-demanding Scots pine (Pinus sylvestris L.) seedlings. An experiment was set up in a nursery in central France to mimic early competition occurring in a newly created gap between Scots pine seedlings and these three common understorey species (young forest-harvested individuals planted at 5 densities from 0 to 57 plants m(-2)). Pine seedling survival and growth (height, diameter, shoot and root biomass) and a functional trait (leaf mass on an area basis, LMA) were measured for 2 years, and cross-analysed against plant density, plant cover and available light. When understorey plant density increased, pine seedling diameter growth and biomass were negatively affected by all three plant species; height growth only slowed beneath Pteridium. These negative effects were closely linked to competition for light beneath Pteridium and Molinia. The application of the Beer-Lambert law gave an extinction coefficient k that was high for Pteridium, intermediate for Molinia and much lower for Calluna. LMA was confirmed as an effective foliar trait to reflect the degree of stress undergone by pine seedlings