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Institute
QuestionBelow-ground processes are key determinants of above-ground plant population and community dynamics. Still, our understanding of how environmental drivers shape plant communities is mostly based on above-ground diversity patterns, bypassing below-ground plant diversity stored in seed banks. As seed banks may shape above-ground plant communities, we question whether concurrently analysing the above- and below-ground species assemblages may potentially enhance our understanding of community responses to environmental variation. LocationTemperate deciduous forests along a 2000km latitudinal gradient in NW Europe. MethodsHerb layer, seed bank and local environmental data including soil pH, canopy cover, forest cover continuity and time since last canopy disturbance were collected in 129 temperate deciduous forest plots. We quantified herb layer and seed bank diversity per plot and evaluated how environmental variation structured community diversity in the herb layer, seed bank and the combined herb layer-seed bank community. ResultsSeed banks consistently held more plant species than the herb layer. How local plot diversity was partitioned across the herb layer and seed bank was mediated by environmental variation in drivers serving as proxies of light availability. The herb layer and seed bank contained an ever smaller and ever larger share of local diversity, respectively, as both canopy cover and time since last canopy disturbance decreased. Species richness and -diversity of the combined herb layer-seed bank community responded distinctly differently compared to the separate assemblages in response to environmental variation in, e.g. forest cover continuity and canopy cover. ConclusionsThe seed bank is a below-ground diversity reservoir of the herbaceous forest community, which interacts with the herb layer, although constrained by environmental variation in e.g. light availability. The herb layer and seed bank co-exist as a single community by means of the so-called storage effect, resulting in distinct responses to environmental variation not necessarily recorded in the individual herb layer or seed bank assemblages. Thus, concurrently analysing above- and below-ground diversity will improve our ecological understanding of how understorey plant communities respond to environmental variation.
Understorey plant communities play a key role in the functioning of forest ecosystems. Under favourable environmental conditions, competitive understorey species may develop high abundances and influence important ecosystem processes such as tree regeneration. Thus, understanding and predicting the response of competitive understorey species as a function of changing environmental conditions is important for forest managers. In the absence of sufficient temporal data to quantify actual vegetation changes, space-for-time (SFT) substitution is often used, i.e. studies that use environmental gradients across space to infer vegetation responses to environmental change over time. Here we assess the validity of such SFT approaches and analysed 36 resurvey studies from ancient forests with low levels of recent disturbances across temperate Europe to assess how six competitive understorey plant species respond to gradients of overstorey cover, soil conditions, atmospheric N deposition and climatic conditions over space and time. The combination of historical and contemporary surveys allows (i) to test if observed contemporary patterns across space are consistent at the time of the historical survey, and, crucially, (ii) to assess whether changes in abundance over time given recorded environmental change match expectations from patterns recorded along environmental gradients in space. We found consistent spatial relationships at the two periods: local variation in soil variables and overstorey cover were the best predictors of individual species’ cover while interregional variation in coarse-scale variables, i.e. N deposition and climate, was less important. However, we found that our SFT approach could not accurately explain the large variation in abundance changes over time. We thus recommend to be cautious when using SFT substitution to infer species responses to temporal changes.
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.
Topsoil conditions in temperate forests are influenced by several soil-forming factors, such as canopy composition (e.g. through litter quality), land-use history, atmospheric deposition, and the parent material. Many studies have evaluated the effects of single factors on physicochemical topsoil conditions, but few have assessed the simultaneous effects of multiple drivers. Here, we evaluate the combined effects of litter quality, land-use history (past land cover as well as past forest management), and atmospheric deposition on several physicochemical topsoil conditions of European temperate deciduous forest soils: bulk density, proportion of exchangeable base cations, carbon/nitrogen-ratio (C/N), litter mass, bio-available and total phosphorus, pH(KCI)and soil organic matter. We collected mineral soil and litter layer samples, and measured site characteristics for 190 20 x 20 m European mixed forest plots across gradients of litter quality (derived from the canopy species composition) and atmospheric deposition, and for different categories of past land cover and past forest management. We accounted for the effects of parent material on topsoil conditions by clustering our plots into three soil type groups based on texture and carbonate concentration. We found that litter quality was a stronger driver of topsoil conditions compared to land-use history or atmospheric deposition, while the soil type also affected several topsoil conditions here. Plots with higher litter quality had soils with a higher proportion of exchangeable base cations, and total phosphorus, and lower C/N-ratios and litter mass. Furthermore, the observed litter quality effects on the topsoil were independent from the regional nitrogen deposition or the soil type, although the soil type likely (co)-determined canopy composition and thus litter quality to some extent in the investigated plots. Litter quality effects on topsoil phosphorus concentrations did interact with past land cover, highlighting the need to consider land-use history when evaluating canopy effects on soil conditions. We conclude that forest managers can use the canopy composition as an important tool for influencing topsoil conditions, although soil type remains an important factor to consider.
1. Worldwide, the floristic composition of temperate forests bears the imprint of past land use for decades to centuries as forests regrow on agricultural land. Many species, however, display significant interregional variation in their ability to (re)colonize post-agricultural forests. This variation in colonization across regions and the underlying factors remain largely unexplored.
2. We compiled data on 90 species and 812 species x study combinations from 18 studies across Europe that determined species' distribution patterns in ancient (i.e. continuously forested since the first available land use maps) and post-agricultural forests. The recovery rate (RR) of species in each landscape was quantified as the log-response ratio of the percentage occurrence in post-agricultural over ancient forest and related to the species-specific life-history traits and local (soil characteristics and light availability) and regional factors (landscape properties as habitat availability, time available for colonization, and climate).
3. For the herb species, we demonstrate a strong (interactive) effect of species' life-history traits and forest habitat availability on the RR of post-agricultural forest. In graminoids, however, none of the investigated variables were significantly related to the RR.
4. The better colonizing species that mainly belonged to the short-lived herbs group showed the largest interregional variability. Their recovery significantly increased with the amount of forest habitat within the landscape, whereas, surprisingly, the time available for colonization, climate, soil characteristics and light availability had no effect.
5. Synthesis. By analysing 18 independent studies across Europe, we clearly showed for the first time on a continental scale that the recovery of short-lived forest herbs increased with the forest habitat availability in the landscape. Small perennial forest herbs, however, were generally unsuccessful in colonizing post-agricultural forest even in relatively densely forested landscapes. Hence, our results stress the need to avoid ancient forest clearance to preserve the typical woodland flora.
The contemporary state of functional traits and species richness in plant communities depends on legacy effects of past disturbances. Whether temporal responses of community properties to current environmental changes are altered by such legacies is, however, unknown. We expect global environmental changes to interact with land-use legacies given different community trajectories initiated by prior management, and subsequent responses to altered resources and conditions. We tested this expectation for species richness and functional traits using 1814 survey-resurvey plot pairs of understorey communities from 40 European temperate forest datasets, syntheses of management transitions since the year 1800, and a trait database. We also examined how plant community indicators of resources and conditions changed in response to management legacies and environmental change. Community trajectories were clearly influenced by interactions between management legacies from over 200 years ago and environmental change. Importantly, higher rates of nitrogen deposition led to increased species richness and plant height in forests managed less intensively in 1800 (i.e., high forests), and to decreases in forests with a more intensive historical management in 1800 (i.e., coppiced forests). There was evidence that these declines in community variables in formerly coppiced forests were ameliorated by increased rates of temperature change between surveys. Responses were generally apparent regardless of sites’ contemporary management classifications, although sometimes the management transition itself, rather than historic or contemporary management types, better explained understorey responses. Main effects of environmental change were rare, although higher rates of precipitation change increased plant height, accompanied by increases in fertility indicator values. Analysis of indicator values suggested the importance of directly characterising resources and conditions to better understand legacy and environmental change effects. Accounting for legacies of past disturbance can reconcile contradictory literature results and appears crucial to anticipating future responses to global environmental change.
Global environmental changes are expected to alter the functional characteristics of understorey herb-layer communities, potentially affecting forest ecosystem functioning. However, little is known about what drives the variability of functional traits in forest understories. Here, we assessed the role of different environmental drivers in shaping the functional trait distribution of understorey herbs in fragmented forests across three spatial scales. We focused on 708 small, deciduous forest patches located in 16 agricultural landscape windows, spanning a 2500-km macroclimatic gradient across the temperate forest biome in Europe. We estimated the relative effect of patch-scale, landscape-scale and macroclimatic variables on the community mean and variation of plant height, specific leaf area and seed mass. Macroclimatic variables (monthly temperature and precipitation extremes) explained the largest proportion of variation in community trait means (on average 77% of the explained variation). In contrast, patch-scale factors dominated in explaining community trait variation (on average 68% of the explained variation). Notably, patch age, size and internal heterogeneity had a positive effect on the community-level variability. Landscape-scale variables explained only a minor part of the variation in both trait distribution properties. The variation explained by shared combinations of the variable groups was generally negligible. These findings highlight the importance of considering multiple spatial scales in predictions of environmental-change effects on the functionality of forest understories. We propose that forest management sustainability could benefit from conserving larger, historically continuous and internally heterogeneous forest patches to maximise ecosystem service diversity in rural landscapes. (C) 2018 Gesellschaft fur Okologie. Published by Elsevier GmbH. All rights reserved.
Environmental drivers interactively affect individual tree growth across temperate European forests
(2018)
Forecasting the growth of tree species to future environmental changes requires abetter understanding of its determinants. Tree growth is known to respond to global‐change drivers such as climate change or atmospheric deposition, as well as to localland‐use drivers such as forest management. Yet, large geographical scale studiesexamining interactive growth responses to multiple global‐change drivers are relativelyscarce and rarely consider management effects. Here, we assessed the interactiveeffects of three global‐change drivers (temperature, precipitation and nitrogen deposi-tion) on individual tree growth of three study species (Quercus robur/petraea, Fagus syl-vatica and Fraxinus excelsior). We sampled trees along spatial environmental gradientsacross Europe and accounted for the effects of management for Quercus. We collectedincrement cores from 267 trees distributed over 151 plots in 19 forest regions andcharacterized their neighbouring environment to take into account potentially confounding factors such as tree size, competition, soil conditions and elevation. Wedemonstrate that growth responds interactively to global‐change drivers, with species ‐specific sensitivities to the combined factors. Simultaneously high levels of precipita-tion and deposition benefited Fraxinus, but negatively affected Quercus’ growth, high-lighting species‐specific interactive tree growth responses to combined drivers. ForFagus, a stronger growth response to higher temperatures was found when precipita-tion was also higher, illustrating the potential negative effects of drought stress underwarming for this species. Furthermore, we show that past forest management canmodulate the effects of changing temperatures on Quercus’ growth; individuals in plotswith a coppicing history showed stronger growth responses to higher temperatures.Overall, our findings highlight how tree growth can be interactively determined by glo-bal‐change drivers, and how these growth responses might be modulated by past for-est management. By showing future growth changes for scenarios of environmentalchange, we stress the importance of considering multiple drivers, including past man-agement and their interactions, when predicting tree growth.
Aim In response to environmental changes and to avoid extinction, species may either track suitable environmental conditions or adapt to the modified environment. However, whether and how species adapt to environmental changes remains unclear. By focusing on the realized niche (i.e. the actual space that a species inhabits and the resources it can access as a result of limiting biotic factors present in its habitat), we here examine shifts in the realized-niche width (i.e. ecological amplitude) and position (i.e. ecological optimum) of 26 common and widespread forest understorey plants across their distributional ranges.
Location Temperate forests along a ca. 1800-km-long latitudinal gradient from northern France to central Sweden and Estonia.
Methods We derived species' realized-niche width from a -diversity metric, which increases if the focal species co-occurs with more species. Based on the concept that species' scores in a detrended correspondence analysis (DCA) represent the locations of their realized-niche positions, we developed a novel approach to run species-specific DCAs allowing the focal species to shift its realized-niche position along the studied latitudinal gradient while the realized-niche positions of other species were held constant.
Results None of the 26 species maintained both their realized-niche width and position along the latitudinal gradient. Few species (9 of 26: 35%) shifted their realized-niche width, but all shifted their realized-niche position. With increasing latitude, most species (22 of 26: 85%) shifted their realized-niche position for soil nutrients and pH towards nutrient-poorer and more acidic soils.
Main conclusions Forest understorey plants shifted their realized niche along the latitudinal gradient, suggesting local adaptation and/or plasticity. This macroecological pattern casts doubt on the idea that the realized niche is stable in space and time, which is a key assumption of species distribution models used to predict the future of biodiversity, hence raising concern about predicted extinction rates.
1. 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 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.