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There is ample evidence for ecological responses to recent climate change. Most studies to date have concentrated on the effects of climate change on individuals and species, with particular emphasis on the effects on phenology and physiology of organisms as well as changes in the distribution and range shifts of species. However, responses by individual species to climate change are not isolated; they are connected through interactions with others at the same or adjacent trophic levels. Also from this more complex perspective, recent case studies have emphasized evidence on the effects of climate change on biotic interactions and ecosystem services. This review highlights the 'knowns' but also 'unknowns' resulting from recent climate impact studies and reveals limitations of (linear) extrapolations from recent climate-induced responses of species to expected trends and magnitudes of future climate change. Hence, there is need not only to continue to focus on the impacts of climate change on the actors in ecological networks but also and more intensively to focus on the linkages between them, and to acknowledge that biotic interactions and feedback processes lead to highly complex, nonlinear and sometimes abrupt responses.

Recent global warming is acting across marine, freshwater, and terrestrial ecosystems to favor species adapted to warmer conditions and/or reduce the abundance of cold-adapted organisms (i.e., "thermophilization" of communities). Lack of community responses to increased temperature, however, has also been reported for several taxa and regions, suggesting that "climatic lags" may be frequent. Here we show that microclimatic effects brought about by forest canopy closure can buffer biotic responses to macroclimate warming, thus explaining an apparent climatic lag. Using data from 1,409 vegetation plots in European and North American temperate forests, each surveyed at least twice over an interval of 12-67 y, we document significant thermophilization of ground-layer plant communities. These changes reflect concurrent declines in species adapted to cooler conditions and increases in species adapted to warmer conditions. However, thermophilization, particularly the increase of warm-adapted species, is attenuated in forests whose canopies have become denser, probably reflecting cooler growing-season ground temperatures via increased shading. As standing stocks of trees have increased in many temperate forests in recent decades, local microclimatic effects may commonly be moderating the impacts of macroclimate warming on forest understories. Conversely, increases in harvesting woody biomass-e.g., for bioenergy-may open forest canopies and accelerate thermophilization of temperate forest biodiversity.climate change | forest management | understory | climatic debt | range shifts B iological signals of recent global warming are increasingly evident across a wide array of ecosystems (1-7). However, the temperature experienced by organisms at ground level (microclimate) can substantially differ from the atmospheric temperature due to local land cover and terrain variation in terms of vegetation structure, shading, topography, or slope orientation (8-15). The daytime or nighttime surface temperature in rough mountain terrain, for instance, can deviate by up to 9°C from the air temperature (10). Likewise, forest structure creates substantial temperature heterogeneity, with the interior daytime temperature in dense forests being commonly several degrees cooler than in more open habitats during the growing season (12-15). Spatial microclimatic temperature variation can thus be substantial relative to projected changes in average temperature over time, and biotic SignificanceAround the globe, climate warming is increasing the dominance of warm-adapted species-a process described as "thermophilization." However, thermophilization often lags behind warming of the climate itself, with some recent studies showing no response at all. Using a unique database of more than 1,400 resurveyed vegetation plots in forests across Europe and North America, we document significant thermophilization of understory vegetation. However, the response to macroclimate warming was attenuated in forests whose canopies have become denser. This microclima...

Summary1. Atmospheric nitrogen (N) deposition is expected to change forest understorey plant community composition and diversity, but results of experimental addition studies and observational studies are not yet conclusive. A shortcoming of observational studies, which are generally based on resurveys or sampling along large deposition gradients, is the occurrence of temporal or spatial confounding factors. 2. We were able to assess the contribution of N deposition versus other ecological drivers on forest understorey plant communities by combining a temporal and spatial approach. Data from 1205 (semi-)permanent vegetation plots taken from 23 rigorously selected understorey resurvey studies along a large deposition gradient across deciduous temperate forest in Europe were compiled and related to various local and regional driving factors, including the rate of atmospheric N deposition, the change in large herbivore densities and the change in canopy cover and composition. 3. Although no directional change in species richness occurred, there was considerable floristic turnover in the understorey plant community and a shift in species composition towards more shade-tolerant and nutrient-demanding species. However, atmospheric N deposition was not important in explaining the observed eutrophication signal. This signal seemed mainly related to a *Correspondence author. E-mail: kris.verheyen@ugent.be Ó 2011 The Authors. Journal of Ecology Ó 2011 British Ecological Society Journal of Ecology 2012Ecology , 100, 352-365 doi: 10.1111Ecology /j.1365Ecology -2745Ecology .2011 shift towards a denser canopy cover and a changed canopy species composition with a higher share of species with more easily decomposed litter. 4. Synthesis. Our multi-site approach clearly demonstrates that one should be cautious when drawing conclusions about the impact of atmospheric N deposition based on the interpretation of plant community shifts in single sites or regions due to other, concurrent, ecological changes. Even though the effects of chronically increased N deposition on the forest plant communities are apparently obscured by the effects of canopy changes, the accumulated N might still have a significant impact. However, more research is needed to assess whether this N time bomb will indeed explode when canopies will open up again.

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