TY - JOUR A1 - De Frenne, Pieter A1 - Baeten, Lander A1 - Graae, Bente J. A1 - Brunet, Jorg A1 - Wulf, Monika A1 - Orczewska, Anna A1 - Kolb, Annette A1 - Jansen, Ivy A1 - Jamoneau, Aurelien A1 - Jacquemyn, Hans A1 - Hermy, Martin A1 - Diekmann, Martin A1 - De Schrijver, An A1 - De Sanctis, Michele A1 - Decocq, Guillaume A1 - Cousins, Sara A. O. A1 - Verheyen, Kris T1 - Interregional variation in the floristic recovery of post-agricultural forests JF - The journal of ecology N2 - 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. KW - ancient forest KW - colonization capacity KW - forest herbs KW - functional traits KW - habitat fragmentation KW - habitat loss KW - life-history traits KW - meta-analysis KW - plant population and community dynamics KW - secondary succession Y1 - 2011 U6 - https://doi.org/10.1111/j.1365-2745.2010.01768.x SN - 0022-0477 VL - 99 IS - 2 SP - 600 EP - 609 PB - Wiley-Blackwell CY - Malden ER - TY - JOUR A1 - De Frenne, Pieter A1 - Brunet, Jorg A1 - Shevtsova, Anna A1 - Kolb, Annette A1 - Graae, Bente J. A1 - Chabrerie, Olivier A1 - Cousins, Sara Ao A1 - Decocq, Guillaume A1 - De Schrijver, An A1 - Diekmann, Martin A1 - Gruwez, Robert A1 - Heinken, Thilo A1 - Hermy, Martin A1 - Nilsson, Christer A1 - Stanton, Sharon A1 - Tack, Wesley A1 - Willaert, Justin A1 - Verheyen, Kris T1 - Temperature effects on forest herbs assessed by warming and transplant experiments along a latitudinal gradient JF - Global change biology N2 - Slow-colonizing forest understorey plants are probably not able to rapidly adjust their distribution range following large-scale climate change. Therefore, the acclimation potential to climate change within their actual occupied habitats will likely be key for their short-and long-term persistence. We combined transplant experiments along a latitudinal gradient with open-top chambers to assess the effects of temperature on phenology, growth and reproductive performance of multiple populations of slow-colonizing understorey plants, using the spring flowering geophytic forb Anemone nemorosa and the early summer flowering grass Milium effusum as study species. In both species, emergence time and start of flowering clearly advanced with increasing temperatures. Vegetative growth (plant height, aboveground biomass) and reproductive success (seed mass, seed germination and germinable seed output) of A. nemorosa benefited from higher temperatures. Climate warming may thus increase future competitive ability and colonization rates of this species. Apart from the effects on phenology, growth and reproductive performance of M. effusum generally decreased when transplanted southwards (e. g., plant size and number of individuals decreased towards the south) and was probably more limited by light availability in the south. Specific leaf area of both species increased when transplanted southwards, but decreased with open-top chamber installation in A. nemorosa. In general, individuals of both species transplanted at the home site performed best, suggesting local adaptation. We conclude that contrasting understorey plants may display divergent plasticity in response to changing temperatures which may alter future understorey community dynamics. KW - climate change KW - common garden experiment KW - forest understorey KW - latitude KW - local adaptation KW - open-top chambers KW - phenotypic plasticity KW - pot experiment Y1 - 2011 U6 - https://doi.org/10.1111/j.1365-2486.2011.02449.x SN - 1354-1013 VL - 17 IS - 10 SP - 3240 EP - 3253 PB - Wiley-Blackwell CY - Hoboken ER - TY - JOUR A1 - Verheyen, Kris A1 - Baeten, Lander A1 - De Frenne, Pieter A1 - Bernhardt-Römermann, Markus A1 - Brunet, Jorg A1 - Cornelis, Johnny A1 - Decocq, Guillaume A1 - Dierschke, Hartmut A1 - Eriksson, Ove A1 - Hedl, Radim A1 - Heinken, Thilo A1 - Hermy, Martin A1 - Hommel, Patrick A1 - Kirby, Keith J. A1 - Naaf, Tobias A1 - Peterken, George A1 - Petrik, Petr A1 - Pfadenhauer, Joerg A1 - Van Calster, Hans A1 - Walther, Gian-Reto A1 - Wulf, Monika A1 - Verstraeten, Gorik T1 - Driving factors behind the eutrophication signal in understorey plant communities of deciduous temperate forests JF - The journal of ecology N2 - 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. KW - atmospheric deposition KW - determinants of plant community diversity and structure KW - Ellenberg indicator values KW - forest herbs KW - forest management KW - large herbivores KW - north-western Europe KW - resurveys KW - (semi-)permanent plots Y1 - 2012 U6 - https://doi.org/10.1111/j.1365-2745.2011.01928.x SN - 0022-0477 VL - 100 IS - 2 SP - 352 EP - 365 PB - Wiley-Blackwell CY - Hoboken ER - TY - JOUR A1 - Graae, Bente J. A1 - De Frenne, Pieter A1 - Kolb, Annette A1 - Brunet, Jorg A1 - Chabrerie, Olivier A1 - Verheyen, Kris A1 - Pepin, Nick A1 - Heinken, Thilo A1 - Zobel, Martin A1 - Shevtsova, Anna A1 - Nijs, Ivan A1 - Milbau, Ann T1 - On the use of weather data in ecological studies along altitudinal and latitudinal gradients JF - Oikos N2 - Global warming has created a need for studies along climatic gradients to assess the effects of temperature on ecological processes. Altitudinal and latitudinal gradients are often used as such, usually in combination with air temperature data from the closest weather station recorded at 1.52 m above the ground. However, many ecological processes occur in, at, or right above the soil surface. To evaluate how representative the commonly used weather station data are for the microclimate relevant for soil surface biota, we compared weather station temperatures for an altitudinal (500900 m a.s.l.) and a latitudinal gradient (4968 degrees N) with data obtained by temperature sensors placed right below the soil surface at five sites along these gradients. The mean annual temperatures obtained from weather stations and adjusted using a lapse rate of -5.5 degrees C km-1 were between 3.8 degrees C lower and 1.6 degrees C higher than those recorded by the temperature sensors at the soil surface, depending on the position along the gradients. The monthly mean temperatures were up to 10 degrees C warmer or 5 degrees C colder at the soil surface. The within-site variation in accumulated temperature was as high as would be expected from a 300 m change in altitude or from a 4 degrees change in latitude or a climate change scenario corresponding to warming of 1.63.8 degrees C. Thus, these differences introduced by the decoupling are significant from a climate change perspective, and the results demonstrate the need for incorporating microclimatic variation when conducting studies along altitudinal or latitudinal gradients. We emphasize the need for using relevant temperature data in climate impact studies and further call for more studies describing the soil surface microclimate, which is crucial for much of the biota. Y1 - 2012 U6 - https://doi.org/10.1111/j.1600-0706.2011.19694.x SN - 0030-1299 VL - 121 IS - 1 SP - 3 EP - 19 PB - Wiley-Blackwell CY - Hoboken ER - TY - JOUR A1 - Plue, Jan A1 - De Frenne, Pieter A1 - Acharya, Kamal P. A1 - Brunet, Jorg A1 - Chabrerie, Olivier A1 - Decocq, Guillaume A1 - Diekmann, Martin A1 - Graae, Bente J. A1 - Heinken, Thilo A1 - Hermy, Martin A1 - Kolb, Annette A1 - Lemke, Isgard A1 - Liira, Jaan A1 - Naaf, Tobias A1 - Shevtsova, Anna A1 - Verheyen, Kris A1 - Wulf, Monika A1 - Cousins, Sara A. O. T1 - Climatic control of forest herb seed banks along a latitudinal gradient JF - Global ecology and biogeography : a journal of macroecology N2 - Aim Seed banks are central to the regeneration strategy of many plant species. Any factor altering seed bank density thus affects plant regeneration and population dynamics. Although seed banks are dynamic entities controlled by multiple environmental drivers, climatic factors are the most comprehensive, but still poorly understood. This study investigates how climatic variation structures seed production and resulting seed bank patterns. Location Temperate forests along a 1900km latitudinal gradient in north-western (NW) Europe. Methods Seed production and seed bank density were quantified in 153 plots along the gradient for four forest herbs with different seed longevity: Geum urbanum, Milium effusum, Poa nemoralis and Stachys sylvatica. We tested the importance of climatic and local environmental factors in shaping seed production and seed bank density. Results Seed production was determined by population size, and not by climatic factors. G.urbanum and M.effusum seed bank density declined with decreasing temperature (growing degree days) and/or increasing temperature range (maximum-minimum temperature). P.nemoralis and S.sylvatica seed bank density were limited by population size and not by climatic variables. Seed bank density was also influenced by other, local environmental factors such as soil pH or light availability. Different seed bank patterns emerged due to differential seed longevities. Species with long-lived seeds maintained constant seed bank densities by counteracting the reduced chance of regular years with high seed production at colder northern latitudes. Main conclusions Seed bank patterns show clear interspecific variation in response to climate across the distribution range. Not all seed banking species may be as well equipped to buffer climate change via their seed bank, notably in short-term persistent species. Since the buffering capacity of seed banks is key to species persistence, these results provide crucial information to advance climatic change predictions on range shifts, community and biodiversity responses. KW - Climate change KW - interspecific variation KW - plant-climate interaction KW - seed longevity KW - seed production KW - temperate deciduous forest KW - temperature Y1 - 2013 U6 - https://doi.org/10.1111/geb.12068 SN - 1466-822X SN - 1466-8238 VL - 22 IS - 10 SP - 1106 EP - 1117 PB - Wiley-Blackwell CY - Hoboken ER - TY - JOUR A1 - Bernhardt-Römermann, Markus A1 - Baeten, Lander A1 - Craven, Dylan A1 - De Frenne, Pieter A1 - Hedl, Radim A1 - Lenoir, Jonathan A1 - Bert, Didier A1 - Brunet, Jorg A1 - Chudomelova, Marketa A1 - Decocq, Guillaume A1 - Dierschke, Hartmut A1 - Dirnboeck, Thomas A1 - Dörfler, Inken A1 - Heinken, Thilo A1 - Hermy, Martin A1 - Hommel, Patrick A1 - Jaroszewicz, Bogdan A1 - Keczynski, Andrzej A1 - Kelly, Daniel L. A1 - Kirby, Keith J. A1 - Kopecky, Martin A1 - Macek, Martin A1 - Malis, Frantisek A1 - Mirtl, Michael A1 - Mitchell, Fraser J. G. A1 - Naaf, Tobias A1 - Newman, Miles A1 - Peterken, George A1 - Petrik, Petr A1 - Schmidt, Wolfgang A1 - Standovar, Tibor A1 - Toth, Zoltan A1 - Van Calster, Hans A1 - Verstraeten, Gorik A1 - Vladovic, Jozef A1 - Vild, Ondrej A1 - Wulf, Monika A1 - Verheyen, Kris T1 - Drivers of temporal changes in temperate forest plant diversity vary across spatial scales JF - Global change biology N2 - Global biodiversity is affected by numerous environmental drivers. Yet, the extent to which global environmental changes contribute to changes in local diversity is poorly understood. We investigated biodiversity changes in a meta-analysis of 39 resurvey studies in European temperate forests (3988 vegetation records in total, 17-75years between the two surveys) by assessing the importance of (i) coarse-resolution (i.e., among sites) vs. fine-resolution (i.e., within sites) environmental differences and (ii) changing environmental conditions between surveys. Our results clarify the mechanisms underlying the direction and magnitude of local-scale biodiversity changes. While not detecting any net local diversity loss, we observed considerable among-site variation, partly explained by temporal changes in light availability (a local driver) and density of large herbivores (a regional driver). Furthermore, strong evidence was found that presurvey levels of nitrogen deposition determined subsequent diversity changes. We conclude that models forecasting future biodiversity changes should consider coarse-resolution environmental changes, account for differences in baseline environmental conditions and for local changes in fine-resolution environmental conditions. KW - atmospheric nitrogen deposition KW - evenness KW - forestREplot KW - forest management KW - game browsing KW - Shannon diversity KW - spatiotemporal resurvey data KW - species richness Y1 - 2015 U6 - https://doi.org/10.1111/gcb.12993 SN - 1354-1013 SN - 1365-2486 VL - 21 IS - 10 SP - 3726 EP - 3737 PB - Wiley-Blackwell CY - Hoboken ER - TY - JOUR A1 - Lemke, Isgard H. A1 - Kolb, Annette A1 - Graae, Bente J. A1 - De Frenne, Pieter A1 - Acharya, Kamal P. A1 - Blandino, Cristina A1 - Brunet, Jorg A1 - Chabrerie, Olivier A1 - Cousins, Sara A. O. A1 - Decocq, Guillaume A1 - Heinken, Thilo A1 - Hermy, Martin A1 - Liira, Jaan A1 - Schmucki, Reto A1 - Shevtsova, Anna A1 - Verheyen, Kris A1 - Diekmann, Martin T1 - Patterns of phenotypic trait variation in two temperate forest herbs along a broad climatic gradient JF - Plant ecology : an international journal N2 - Phenotypic trait variation plays a major role in the response of plants to global environmental change, particularly in species with low migration capabilities and recruitment success. However, little is known about the variation of functional traits within populations and about differences in this variation on larger spatial scales. In a first approach, we therefore related trait expression to climate and local environmental conditions, studying two temperate forest herbs, Milium effusum and Stachys sylvatica, along a similar to 1800-2500 km latitudinal gradient. Within each of 9-10 regions in six European countries, we collected data from six populations of each species and recorded several variables in each region (temperature, precipitation) and population (light availability, soil parameters). For each plant, we measured height, leaf area, specific leaf area, seed mass and the number of seeds and examined environmental effects on within-population trait variation as well as on trait means. Most importantly, trait variation differed both between and within populations. Species, however, differed in their response. Intrapopulation variation in Milium was consistently positively affected by higher mean temperatures and precipitation as well as by more fertile local soil conditions, suggesting that more productive conditions may select for larger phenotypic variation. In Stachys, particularly light availability positively influenced trait variation, whereas local soil conditions had no consistent effects. Generally, our study emphasises that intra-population variation may differ considerably across larger scales-due to phenotypic plasticity and/or underlying genetic diversity-possibly affecting species response to global environmental change. KW - Climate change KW - Global environmental change KW - Milium effusum KW - Phenotypic plasticity KW - Intraspecific variation KW - Stachys sylvatica Y1 - 2015 U6 - https://doi.org/10.1007/s11258-015-0534-0 SN - 1385-0237 SN - 1573-5052 VL - 216 IS - 11 SP - 1523 EP - 1536 PB - Springer CY - Dordrecht ER - TY - JOUR A1 - Maes, Sybryn L. A1 - Perring, Michael P. A1 - Vanhellemont, Margot A1 - Depauw, Leen A1 - Van den Bulcke, Jan A1 - Brumelis, Guntis A1 - Brunet, Jorg A1 - Decocq, Guillaume A1 - den Ouden, Jan A1 - Härdtle, Werner A1 - Hedl, Radim A1 - Heinken, Thilo A1 - Heinrichs, Steffi A1 - Jaroszewicz, Bogdan A1 - Kopecký, Martin A1 - Malis, Frantisek A1 - Wulf, Monika A1 - Verheyen, Kris T1 - Environmental drivers interactively affect individual tree growth across temperate European forests JF - Global change biology N2 - 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. KW - basal area increment KW - climate change KW - Fagus KW - Fraxinus KW - historical ecology KW - nitrogen deposition KW - Quercus KW - tree-ring analysis Y1 - 2018 U6 - https://doi.org/10.1111/gcb.14493 SN - 1354-1013 SN - 1365-2486 VL - 25 IS - 1 SP - 201 EP - 217 PB - Wiley CY - Hoboken ER - TY - JOUR A1 - De Lombaerde, Emiel A1 - Verheyen, Kris A1 - Perring, Michael P. A1 - Bernhardt-Roemermann, Markus A1 - Van Calster, Hans A1 - Brunet, Jorg A1 - Chudomelova, Marketa A1 - Decocq, Guillaume A1 - Diekmann, Martin A1 - Durak, Tomasz A1 - Hedl, Radim A1 - Heinken, Thilo A1 - Hommel, Patrick A1 - Jaroszewicz, Bogdan A1 - Kopecky, Martin A1 - Lenoir, Jonathan A1 - Macek, Martin A1 - Máliš, František A1 - Mitchell, Fraser J. G. A1 - Naaf, Tobias A1 - Newman, Miles A1 - Petřík, Petr A1 - Reczyńska, Kamila A1 - Schmidt, Wolfgang A1 - Swierkosz, Krzysztof A1 - Vild, Ondrej A1 - Wulf, Monika A1 - Baetena, Lander T1 - Responses of competitive understorey species to spatial environmental gradients inaccurately explain temporal changes JF - Basic and applied ecology : Journal of the Gesellschaft für Ökologie N2 - 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. KW - Temperate forest KW - Herb layer KW - Tree regeneration KW - Global change KW - Nitrogen deposition KW - Canopy KW - Spatiotemporal resurvey data KW - Cover abundance KW - Chronosequence KW - forestREplot Y1 - 2018 U6 - https://doi.org/10.1016/j.baae.2018.05.013 SN - 1439-1791 SN - 1618-0089 VL - 30 SP - 52 EP - 64 PB - Elsevier GMBH CY - München ER - TY - JOUR A1 - Vanneste, Thomas A1 - Valdes, Alicia A1 - Verheyen, Kris A1 - Perring, Michael P. A1 - Bernhardt-Roemermann, Markus A1 - Andrieu, Emilie A1 - Brunet, Jorg A1 - Cousins, Sara A. O. A1 - Deconchat, Marc A1 - De Smedt, Pallieter A1 - Diekmann, Martin A1 - Ehrmann, Steffen A1 - Heinken, Thilo A1 - Hermy, Martin A1 - Kolb, Annette A1 - Lenoir, Jonathan A1 - Liira, Jaan A1 - Naaf, Tobias A1 - Paal, Taavi A1 - Wulf, Monika A1 - Decocq, Guillaume A1 - De Frenne, Pieter T1 - Functional trait variation of forest understorey plant communities across Europe JF - Basic and applied ecology : Journal of the Gesellschaft für Ökologie N2 - 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. KW - Agricultural landscapes KW - Biogeography KW - Community ecology KW - Forest understorey KW - Functional trait diversity KW - Fragmentation KW - Global environmental change KW - Landscape connectivity KW - Macroclimatic gradient KW - Multi-scale analysis Y1 - 2018 U6 - https://doi.org/10.1016/j.baae.2018.09.004 SN - 1439-1791 SN - 1618-0089 VL - 34 SP - 1 EP - 14 PB - Elsevier GmbH CY - München ER -