@article{MaesPerringVanhellemontetal.2018, author = {Maes, Sybryn L. and Perring, Michael P. and Vanhellemont, Margot and Depauw, Leen and Van den Bulcke, Jan and Brumelis, Guntis and Brunet, Jorg and Decocq, Guillaume and den Ouden, Jan and H{\"a}rdtle, Werner and Hedl, Radim and Heinken, Thilo and Heinrichs, Steffi and Jaroszewicz, Bogdan and Kopeck{\´y}, Martin and Malis, Frantisek and Wulf, Monika and Verheyen, Kris}, title = {Environmental drivers interactively affect individual tree growth across temperate European forests}, series = {Global change biology}, volume = {25}, journal = {Global change biology}, number = {1}, publisher = {Wiley}, address = {Hoboken}, issn = {1354-1013}, doi = {10.1111/gcb.14493}, pages = {201 -- 217}, year = {2018}, abstract = {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.}, language = {en} } @article{PlueDeFrenneAcharyaetal.2017, author = {Plue, Jan and De Frenne, Pieter and Acharya, Kamal and Brunet, J{\"o}rg and Chabrerie, Olivier and Decocq, Guillaume and Diekmann, Martin and Graae, Bente J. and Heinken, Thilo and Hermy, Martin and Kolb, Annette and Lemke, Isgard and Liira, Jaan and Naaf, Tobias and Verheyen, Kris and Wulf, Monika and Cousins, Sara A. O.}, title = {Where does the community start, and where does it end?}, series = {Journal of vegetation science}, volume = {28}, journal = {Journal of vegetation science}, number = {2}, publisher = {Wiley}, address = {Hoboken}, issn = {1100-9233}, doi = {10.1111/jvs.12493}, pages = {424 -- 435}, year = {2017}, abstract = {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.}, language = {en} } @article{DeLombaerdeVerheyenPerringetal.2018, author = {De Lombaerde, Emiel and Verheyen, Kris and Perring, Michael P. and Bernhardt-Roemermann, Markus and Van Calster, Hans and Brunet, Jorg and Chudomelova, Marketa and Decocq, Guillaume and Diekmann, Martin and Durak, Tomasz and Hedl, Radim and Heinken, Thilo and Hommel, Patrick and Jaroszewicz, Bogdan and Kopecky, Martin and Lenoir, Jonathan and Macek, Martin and M{\´a}liš, František and Mitchell, Fraser J. G. and Naaf, Tobias and Newman, Miles and Petř{\´i}k, Petr and Reczyńska, Kamila and Schmidt, Wolfgang and Swierkosz, Krzysztof and Vild, Ondrej and Wulf, Monika and Baetena, Lander}, title = {Responses of competitive understorey species to spatial environmental gradients inaccurately explain temporal changes}, series = {Basic and applied ecology : Journal of the Gesellschaft f{\"u}r {\"O}kologie}, volume = {30}, journal = {Basic and applied ecology : Journal of the Gesellschaft f{\"u}r {\"O}kologie}, publisher = {Elsevier GMBH}, address = {M{\"u}nchen}, issn = {1439-1791}, doi = {10.1016/j.baae.2018.05.013}, pages = {52 -- 64}, year = {2018}, abstract = {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.}, language = {en} } @article{PerringBernhardtRoemermannBaetenetal.2018, author = {Perring, Michael P. and Bernhardt-Roemermann, Markus and Baeten, Lander and Midolo, Gabriele and Blondeel, Haben and Depauw, Leen and Landuyt, Dries and Maes, Sybryn L. and De Lombaerde, Emiel and Caron, Maria Mercedes and Vellend, Mark and Brunet, Joerg and Chudomelova, Marketa and Decocq, Guillaume and Diekmann, Martin and Dirnboeck, Thomas and Doerfler, Inken and Durak, Tomasz and De Frenne, Pieter and Gilliam, Frank S. and Hedl, Radim and Heinken, Thilo and Hommel, Patrick and Jaroszewicz, Bogdan and Kirby, Keith J. and Kopecky, Martin and Lenoir, Jonathan and Li, Daijiang and Malis, Frantisek and Mitchell, Fraser J. G. and Naaf, Tobias and Newman, Miles and Petrik, Petr and Reczynska, Kamila and Schmidt, Wolfgang and Standovar, Tibor and Swierkosz, Krzysztof and Van Calster, Hans and Vild, Ondrej and Wagner, Eva Rosa and Wulf, Monika and Verheyen, Kris}, title = {Global environmental change effects on plant community composition trajectories depend upon management legacies}, series = {Global change biology}, volume = {24}, journal = {Global change biology}, number = {4}, publisher = {Wiley}, address = {Hoboken}, issn = {1354-1013}, doi = {10.1111/gcb.14030}, pages = {1722 -- 1740}, year = {2018}, abstract = {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.}, language = {en} } @article{MaesBlondeelPerringetal.2019, author = {Maes, Sybryn L. and Blondeel, Haben and Perring, Michael P. and Depauw, Leen and Brumelis, Guntis and Brunet, J{\"o}rg and Decocq, Guillaume and den Ouden, Jan and Haerdtle, Werner and Hedl, Radim and Heinken, Thilo and Heinrichs, Steffi and Jaroszewicz, Bogdan and Kirby, Keith J. and Kopecky, Martin and Malis, Frantisek and Wulf, Monika and Verheyen, Kris}, title = {Litter quality, land-use history, and nitrogen deposition effects on topsoil conditions across European temperate deciduous forests}, series = {Forest ecology and management}, volume = {433}, journal = {Forest ecology and management}, publisher = {Elsevier}, address = {Amsterdam}, issn = {0378-1127}, doi = {10.1016/j.foreco.2018.10.056}, pages = {405 -- 418}, year = {2019}, abstract = {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.}, language = {en} } @article{VannesteValdesVerheyenetal.2018, author = {Vanneste, Thomas and Valdes, Alicia and Verheyen, Kris and Perring, Michael P. and Bernhardt-Roemermann, Markus and Andrieu, Emilie and Brunet, Jorg and Cousins, Sara A. O. and Deconchat, Marc and De Smedt, Pallieter and Diekmann, Martin and Ehrmann, Steffen and Heinken, Thilo and Hermy, Martin and Kolb, Annette and Lenoir, Jonathan and Liira, Jaan and Naaf, Tobias and Paal, Taavi and Wulf, Monika and Decocq, Guillaume and De Frenne, Pieter}, title = {Functional trait variation of forest understorey plant communities across Europe}, series = {Basic and applied ecology : Journal of the Gesellschaft f{\"u}r {\"O}kologie}, volume = {34}, journal = {Basic and applied ecology : Journal of the Gesellschaft f{\"u}r {\"O}kologie}, publisher = {Elsevier GmbH}, address = {M{\"u}nchen}, issn = {1439-1791}, doi = {10.1016/j.baae.2018.09.004}, pages = {1 -- 14}, year = {2018}, abstract = {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.}, language = {en} } @article{deFrenneGraaeKolbetal.2010, author = {de Frenne, Pieter and Graae, Bente Jessen and Kolb, Annette and Brunet, J{\"o}rg and Chabrerie, Olivier and Cousins, Sara A. O. and Decocq, Guillaume and Dhondt, Rob and Diekmann, Martin and Eriksson, Olof and Heinken, Thilo and Hermy, Martin and J{\"o}gar, uelle and Saguez, Robert and Shevtsova, Anna and Stanton, Sharon and Zindel, Renate and Zobel, Martin and Verheyen, Kris}, title = {Significant effects of temperature on the reproductive output of the forest herb Anemone nemorosa L.}, issn = {0378-1127}, doi = {10.1016/j.foreco.2009.04.038}, year = {2010}, abstract = {Climate warming is already influencing plant migration in different parts of the world. Numerous models have been developed to forecast future plant distributions. Few studies, however, have investigated the potential effect of warming on the reproductive output of plants. Understorey forest herbs in particular, have received little attention in the debate on climate change impacts. This study focuses on the effect of temperature on sexual reproductive output (number of seeds, seed mass, germination percentage and seedling mass) of Anemone nemorosa L., a model species for slow colonizing herbaceous forest plants. We sampled seeds of A. nemorosa in populations along a 2400 km latitudinal gradient from northern France to northern Sweden during three growing seasons (2005,2006 and 2008). This study design allowed us to isolate the effects of accumulated temperature (Growing Degree Hours; GDH) from latitude and the local abiotic and biotic environment. Germination and seed sowing trials were performed in incubators, a greenhouse and under field conditions in a forest. Finally, we disentangled correlations between the different reproductive traits of A. nemorosa along the latitudinal gradient. We found a clear positive relationship between accumulated temperature and seed and seedling traits: reproductive output of A. nemorosa improved with increasing GDH along the latitudinal gradient. Seed mass and seedling mass, for instance, increased by 9.7\% and 10.4\%, respectively, for every 1000 degrees C h increase in GDH. We also derived strong correlations between several seed and seedling traits both under field conditions and in incubators. Our results indicate that seed mass, incubator-based germination percentage (Germ\%(Inc)) and the output of germinable seeds (product of number of seeds and Germ\%(Inc) divided by 100) from plants grown along a latitudinal gradient (i.e. at different temperature regimes) provide valuable proxies to parameterize key population processes in models. We conclude that (1) climate warming may have a pronounced positive impact on sexual reproduction of A. nemorosa and (2) climate models forecasting plant distributions would benefit from including the temperature sensitivity of key seed traits and population processes.}, language = {en} } @article{PlueDeFrenneAcharyaetal.2013, author = {Plue, Jan and De Frenne, Pieter and Acharya, Kamal P. and Brunet, Jorg and Chabrerie, Olivier and Decocq, Guillaume and Diekmann, Martin and Graae, Bente J. and Heinken, Thilo and Hermy, Martin and Kolb, Annette and Lemke, Isgard and Liira, Jaan and Naaf, Tobias and Shevtsova, Anna and Verheyen, Kris and Wulf, Monika and Cousins, Sara A. O.}, title = {Climatic control of forest herb seed banks along a latitudinal gradient}, series = {Global ecology and biogeography : a journal of macroecology}, volume = {22}, journal = {Global ecology and biogeography : a journal of macroecology}, number = {10}, publisher = {Wiley-Blackwell}, address = {Hoboken}, issn = {1466-822X}, doi = {10.1111/geb.12068}, pages = {1106 -- 1117}, year = {2013}, abstract = {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.}, language = {en} } @article{WasofLenoirGalletMoronetal.2013, author = {Wasof, Safaa and Lenoir, Jonathan and Gallet-Moron, Emilie and Jamoneau, Aurelien and Brunet, J{\"o}rg and Cousins, Sara A. O. and De Frenne, Pieter and Diekmann, Martin and Hermy, Martin and Kolb, Annette and Liira, Jaan and Verheyen, Kris and Wulf, Monika and Decocq, Guillaume}, title = {Ecological niche shifts of understorey plants along a latitudinal gradient of temperate forests in north-western Europe}, series = {Global ecology and biogeography : a journal of macroecology}, volume = {22}, journal = {Global ecology and biogeography : a journal of macroecology}, number = {10}, publisher = {Wiley-Blackwell}, address = {Hoboken}, issn = {1466-822X}, doi = {10.1111/geb.12073}, pages = {1130 -- 1140}, year = {2013}, abstract = {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.}, language = {en} } @article{DeFrenneRodriguezSanchezCoomesetal.2013, author = {De Frenne, Pieter and Rodriguez-Sanchez, Francisco and Coomes, David Anthony and B{\"a}ten, Lander and Verstr{\"a}ten, Gorik and Vellend, Mark and Bernhardt-R{\"o}mermann, Markus and Brown, Carissa D. and Brunet, J{\"o}rg and Cornelis, Johnny and Decocq, Guillaume M. and Dierschke, Hartmut and Eriksson, Ove and Gilliam, Frank S. and Hedl, Radim and Heinken, Thilo and Hermy, Martin and Hommel, Patrick and Jenkins, Michael A. and Kelly, Daniel L. and Kirby, Keith J. and Mitchell, Fraser J. G. and Naaf, Tobias and Newman, Miles and Peterken, George and Petrik, Petr and Schultz, Jan and Sonnier, Gregory and Van Calster, Hans and Waller, Donald M. and Walther, Gian-Reto and White, Peter S. and Woods, Kerry D. and Wulf, Monika and Graae, Bente Jessen and Verheyen, Kris}, title = {Microclimate moderates plant responses to macroclimate warming}, series = {Proceedings of the National Academy of Sciences of the United States of America}, volume = {110}, journal = {Proceedings of the National Academy of Sciences of the United States of America}, number = {46}, publisher = {National Acad. of Sciences}, address = {Washington}, issn = {0027-8424}, doi = {10.1073/pnas.1311190110}, pages = {18561 -- 18565}, year = {2013}, abstract = {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.}, language = {en} } @article{deFrenneKolbVerheyenetal.2009, author = {de Frenne, Pieter and Kolb, Annette and Verheyen, Kris and Brunet, Johanne and Chabrerie, Olivier and Decocq, Guillaume and Diekmann, Martin and Eriksson, Ove and Heinken, Thilo and Hermy, Martin and J{\~o}gar, {\"U}lle and Stanton, Sara and Quataert, Paul and Zindel, Renate and Zobel, Martin and Graae, Bente Jessen}, title = {Unravelling the effects of temperature, latitude and local environment on the reproduction of forest herbs}, issn = {1466-822X}, doi = {10.1111/j.1466-8238.2009.00487.x}, year = {2009}, abstract = {Aim To investigate the effect of temperature, latitude and local environment on the reproductive traits of widespread perennial forest herbs to better understand the potential impacts of rising temperatures on their population dynamics and colonization capacities. Location Six regions along a latitudinal gradient from France to Sweden. Methods Within each region, we collected data from three to five populations of up to six species. For each species, several variables were recorded in each region (temperature, latitude) and population (local abiotic and biotic environmental variables), and seed production and germination were estimated. Resource investment in reproduction (RIR) was quantified as seed number ¥ seed mass, while germinable seed output (GSO) was expressed as seed number ¥ germination percentage.We performed linear regression and mixed effect models to investigate the effects of temperature (growing degree hours), latitude and local abiotic and biotic environment on RIR and GSO. Results Temperature and latitude explained most of the variation in RIR and GSO for early flowering species with a northerly distribution range edge (Anemone nemorosa, Paris quadrifolia and Oxalis acetosella). Reproduction of the more southerly distributed species (Brachypodium sylvaticum, Circaea lutetiana and Primula elatior), in contrast, was independent of temperature/latitude. In the late summer species, B. sylvaticum and C. lutetiana, variation in RIR and GSO was best explained by local environmental variables, while none of the investigated variables appeared to be related to reproduction in P. elatior. Main conclusions We showed that reproduction of only two early flowering, northerly distributed species was related to temperature. This suggests that the potential reproductive response of forest herbs to climate warming partly depends on their phenology and distribution, but also that the response is to some extent species dependent. These findings should be taken into account when predictions about future shifts in distribution range are made.}, language = {en} } @article{GraaeVerheyenKolbetal.2009, author = {Graae, Bente Jessen and Verheyen, Kris and Kolb, Annette and van der Veken, Sebastian and Heinken, Thilo and Chabrerie, Olivier and Diekmann, Martin and Valtinat, Karin and Zindel, Renate and Karlsson, Elisabeth and Str{\"o}m, Lotta and Decocq, Guillaume and Hermy, Martin and Baskin, Carol C.}, title = {Germination requirements and seed mass of slow- and fast-colonizing temperate forest herbs along a latitudinal gradient}, issn = {1195-6860}, doi = {10.2980/16-2-3234}, year = {2009}, abstract = {Predictions on displacement of suitable habitats due to climate change suggest that plant species with poor colonization ability may be unable to move fast enough to match forecasted climate-induced changes in habitat distribution. However, studies on early Holocene plant migration show fast migration of many plant species that are poor colonizers today. We hypothesize that warmer temperatures during the early Holocene yielded higher seed quality, contributing to explaining the fast migration. We studied how the 3 seed quality variables, seed mass, germinability, and requirements for break of seed dormancy, vary for seeds of 11 forest herb species with varying colonization capacity collected along a 1400-km latitudinal gradient. Within species, seed mass showed a positive correlation with latitude, whereas germinability was more positively correlated with temperature (growing degree hours obtained at time of seed collection). Only slow-colonizing species increased germinability with temperature, whereas only fast-colonizing species increased germinability with latitude. These interactions were only detectable when analyzing germinability of the seeds, even though this trait and seed mass were correlated. The requirement for dormancy break did not correlate with latitude or temperature. The results indicate that seed development of slow colonizers may be favoured by a warmer climate, which in turn may be important for their migration capacity.}, language = {en} } @article{VerheyenBaetenDeFrenneetal.2012, author = {Verheyen, Kris and Baeten, Lander and De Frenne, Pieter and Bernhardt-R{\"o}mermann, Markus and Brunet, Jorg and Cornelis, Johnny and Decocq, Guillaume and Dierschke, Hartmut and Eriksson, Ove and Hedl, Radim and Heinken, Thilo and Hermy, Martin and Hommel, Patrick and Kirby, Keith J. and Naaf, Tobias and Peterken, George and Petrik, Petr and Pfadenhauer, Joerg and Van Calster, Hans and Walther, Gian-Reto and Wulf, Monika and Verstraeten, Gorik}, title = {Driving factors behind the eutrophication signal in understorey plant communities of deciduous temperate forests}, series = {The journal of ecology}, volume = {100}, journal = {The journal of ecology}, number = {2}, publisher = {Wiley-Blackwell}, address = {Hoboken}, issn = {0022-0477}, doi = {10.1111/j.1365-2745.2011.01928.x}, pages = {352 -- 365}, year = {2012}, abstract = {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.}, language = {en} } @article{CaronDeFrenneBrunetetal.2015, author = {Caron, Maria Mercedes and De Frenne, Pieter and Brunet, J{\"o}rg and Chabrerie, Olivier and Cousins, Sara A. O. and Decocq, Guillaume and Diekmann, Martin and Graae, Bente Jessen and Heinken, Thilo and Kolb, Annette and Lenoir, Jonathan and Naaf, Tobias and Plue, Jan and Selvi, Federico and Wulf, Monika and Verheyen, Kris}, title = {Divergent regeneration responses of two closely related tree species to direct abiotic and indirect biotic effects of climate change}, series = {Forest ecology and management}, volume = {342}, journal = {Forest ecology and management}, publisher = {Elsevier}, address = {Amsterdam}, issn = {0378-1127}, doi = {10.1016/j.foreco.2015.01.003}, pages = {21 -- 29}, year = {2015}, abstract = {Changing temperature and precipitation can strongly influence plant reproduction. However, also biotic interactions might indirectly affect the reproduction and recruitment success of plants in the context of climate change. Information about the interactive effects of changes in abiotic and biotic factors is essential, but still largely lacking, to better understand the potential effects of a changing climate on plant populations. Here we analyze the regeneration from seeds of Acer platanoides and Acer pseudoplatanus, two currently secondary forest tree species from seven regions along a 2200 km-wide latitudinal gradient in Europe. We assessed the germination, seedling survival and growth during two years in a common garden experiment where temperature, precipitation and competition with the understory vegetation were manipulated. A. platanoides was more sensitive to changes in biotic conditions while A. pseudoplatanus was affected by both abiotic and biotic changes. In general, competition reduced (in A. platanoides) and warming enhanced (in A. pseudoplatanus) germination and survival, respectively. Reduced competition strongly increased the growth of A. platanoides seedlings. Seedling responses were independent of the conditions experienced by the mother tree during seed production and maturation. Our results indicate that, due to the negative effects of competition on the regeneration of A. platanoides, it is likely that under stronger competition (projected under future climatic conditions) this species will be negatively affected in terms of germination, survival and seedling biomass. Climate-change experiments including both abiotic and biotic factors constitute a key step forward to better understand the response of tree species' regeneration to climate change. (C) 2015 Elsevier B.V. All rights reserved.}, language = {en} } @article{DeFrenneGraaeBrunetetal.2012, author = {De Frenne, Pieter and Graae, Bente J. and Brunet, J{\"o}rg and Shevtsova, Anna and De Schrijver, An and Chabrerie, Olivier and Cousins, Sara A. O. and Decocq, Guillaume and Diekmann, Martin and Hermy, Martin and Heinken, Thilo and Kolb, Annette and Nilsson, Christer and Stanton, Sharon and Verheyen, Kris}, title = {The response of forest plant regeneration to temperature variation along a latitudinal gradient}, series = {Annals of botany}, volume = {109}, journal = {Annals of botany}, number = {5}, publisher = {Oxford Univ. Press}, address = {Oxford}, issn = {0305-7364}, doi = {10.1093/aob/mcs015}, pages = {1037 -- 1046}, year = {2012}, abstract = {The response of forest herb regeneration from seed to temperature variations across latitudes was experimentally assessed in order to forecast the likely response of understorey community dynamics to climate warming. Seeds of two characteristic forest plants (Anemone nemorosa and Milium effusum) were collected in natural populations along a latitudinal gradient from northern France to northern Sweden and exposed to three temperature regimes in growth chambers (first experiment). To test the importance of local adaptation, reciprocal transplants were also made of adult individuals that originated from the same populations in three common gardens located in southern, central and northern sites along the same gradient, and the resulting seeds were germinated (second experiment). Seedling establishment was quantified by measuring the timing and percentage of seedling emergence, and seedling biomass in both experiments. Spring warming increased emergence rates and seedling growth in the early-flowering forb A. nemorosa. Seedlings of the summer-flowering grass M. effusum originating from northern populations responded more strongly in terms of biomass growth to temperature than southern populations. The above-ground biomass of the seedlings of both species decreased with increasing latitude of origin, irrespective of whether seeds were collected from natural populations or from the common gardens. The emergence percentage decreased with increasing home-away distance in seeds from the transplant experiment, suggesting that the maternal plants were locally adapted. Decreasing seedling emergence and growth were found from the centre to the northern edge of the distribution range for both species. Stronger responses to temperature variation in seedling growth of the grass M. effusum in the north may offer a way to cope with environmental change. The results further suggest that climate warming might differentially affect seedling establishment of understorey plants across their distribution range and thus alter future understorey plant dynamics.}, language = {en} } @article{BaetenWartonVanCalsteretal.2014, author = {Baeten, Lander and Warton, David I. and Van Calster, Hans and De Frenne, Pieter and Verstraeten, Gorik and Bonte, Dries and Bernhardt-R{\"o}mermann, Markus and Cornelis, Johnny and Decocq, Guillaume and Eriksson, Ove and Hedl, Radim and Heinken, Thilo and Hermy, Martin and Hommel, Patrick and Kirby, Keith J. and Naaf, Tobias and Petrik, Petr and Walther, Gian-Reto and Wulf, Monica and Verheyen, Kris}, title = {A model-based approach to studying changes in compositional heterogeneity}, series = {Methods in ecology and evolution : an official journal of the British Ecological Society}, volume = {5}, journal = {Methods in ecology and evolution : an official journal of the British Ecological Society}, number = {2}, publisher = {Wiley-Blackwell}, address = {Hoboken}, issn = {2041-210X}, pages = {156 -- 164}, year = {2014}, language = {en} } @article{DeFrenneKolbGraaeetal.2011, author = {De Frenne, P. and Kolb, Annette and Graae, Benete Jessen and Decocq, Guillaume and Baltora, S. and De Schrijver, A. and Brunet, J. and Chabrerie, Olivier and Cousins, Sara A. O. and Dhondt, Rob and Diekmann, Martin and Gruwez, R. and Heinken, Thilo and Hermy, Martin and Liira, J. and Saguez, R. and Shevtsova, Anna and Baskin, Carol C. and Verheyen, Kris}, title = {A latitudinal gradient in seed nutrients of the forest herb Anemone nemorosa}, series = {Plant biology}, volume = {13}, journal = {Plant biology}, number = {3}, publisher = {Wiley-Blackwell}, address = {Malden}, issn = {1435-8603}, doi = {10.1111/j.1438-8677.2010.00404.x}, pages = {493 -- 501}, year = {2011}, abstract = {The nutrient concentration in seeds determines many aspects of potential success of the sexual reproductive phase of plants, including the seed predation probability, efficiency of seed dispersal and seedling performance. Despite considerable research interest in latitudinal gradients of foliar nutrients, a similar gradient for seeds remains unexplored. We investigated a potential latitudinal gradient in seed nutrient concentrations within the widespread European understorey forest herb Anemone nemorosa L. We sampled seeds of A. nemorosa in 15 populations along a 1900-km long latitudinal gradient at three to seven seed collection dates post-anthesis and investigated the relative effects of growing degree-hours > 5 degrees C, soil characteristics and latitude on seed nutrient concentrations. Seed nitrogen, nitrogen:phosphorus ratio and calcium concentration decreased towards northern latitudes, while carbon:nitrogen ratios increased. When taking differences in growing degree-hours and measured soil characteristics into account and only considering the most mature seeds, the latitudinal decline remained particularly significant for seed nitrogen concentration. We argue that the decline in seed nitrogen concentration can be attributed to northward decreasing seed provisioning due to lower soil nitrogen availability or greater investment in clonal reproduction. This pattern may have large implications for the reproductive performance of this forest herb as the degree of seed provisioning ultimately co-determines seedling survival and reproductive success.}, language = {en} } @article{DeFrenneGraaeKolbetal.2011, author = {De Frenne, Pieter and Graae, Bente J. and Kolb, Annette and Shevtsova, Anna and Baeten, Lander and Brunet, J{\"o}rg and Chabrerie, Olivier and Cousins, Sara A. O. and Decocq, Guillaume and Dhondt, Rob and Diekmann, Martin and Gruwez, Robert and Heinken, Thilo and Hermy, Martin and Oster, Mathias and Saguez, Robert and Stanton, Sharon and Tack, Wesley and Vanhellemont, Margot and Verheyen, Kris}, title = {An intraspecific application of the leaf-height-seed ecology strategy scheme to forest herbs along a latitudinal gradient}, series = {Ecography : pattern and diversity in ecology ; research papers forum}, volume = {34}, journal = {Ecography : pattern and diversity in ecology ; research papers forum}, number = {1}, publisher = {Wiley-Blackwell}, address = {Malden}, issn = {0906-7590}, doi = {10.1111/j.1600-0587.2010.06399.x}, pages = {132 -- 140}, year = {2011}, abstract = {We measured LHS traits in 41 Anemone nemorosa and 44 Milium effusum populations along a 1900-2300 km latitudinal gradient from N France to N Sweden. We then applied multilevel models to identify the effects of regional (temperature, latitude) and local (soil fertility and acidity, overstorey canopy cover) environmental factors on LHS traits. Both species displayed a significant 4\% increase in plant height with every degree northward shift (almost a two-fold plant height difference between the southernmost and northernmost populations). Neither seed mass nor SLA showed a significant latitudinal cline. Temperature had a large effect on the three LHS traits of Anemone. Latitude, canopy cover and soil nutrients were related to the SLA and plant height of Milium. None of the investigated variables appeared to be related to the seed mass of Milium. The variation in LHS traits indicates that the ecological strategy determined by the position of each population in this three-factor triangle is not constant along the latitudinal gradient. The significant increase in plant height suggests greater competitive abilities for both species in the northernmost populations. We also found that the studied environmental factors affected the LHS traits of the two species on various scales: spring-flowering Anemone was affected more by temperature, whereas early-summer flowering Milium was affected more by local and other latitude-related factors. Finally, previously reported cross-species correlations between LHS traits and latitude were generally unsupported by our within-species approach.}, language = {en} } @article{DeFrenneBaetenGraaeetal.2011, author = {De Frenne, Pieter and Baeten, Lander and Graae, Bente J. and Brunet, Jorg and Wulf, Monika and Orczewska, Anna and Kolb, Annette and Jansen, Ivy and Jamoneau, Aurelien and Jacquemyn, Hans and Hermy, Martin and Diekmann, Martin and De Schrijver, An and De Sanctis, Michele and Decocq, Guillaume and Cousins, Sara A. O. and Verheyen, Kris}, title = {Interregional variation in the floristic recovery of post-agricultural forests}, series = {The journal of ecology}, volume = {99}, journal = {The journal of ecology}, number = {2}, publisher = {Wiley-Blackwell}, address = {Malden}, issn = {0022-0477}, doi = {10.1111/j.1365-2745.2010.01768.x}, pages = {600 -- 609}, year = {2011}, abstract = {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.}, language = {en} } @article{BernhardtRoemermannBaetenCravenetal.2015, author = {Bernhardt-R{\"o}mermann, Markus and Baeten, Lander and Craven, Dylan and De Frenne, Pieter and Hedl, Radim and Lenoir, Jonathan and Bert, Didier and Brunet, Jorg and Chudomelova, Marketa and Decocq, Guillaume and Dierschke, Hartmut and Dirnboeck, Thomas and D{\"o}rfler, Inken and Heinken, Thilo and Hermy, Martin and Hommel, Patrick and Jaroszewicz, Bogdan and Keczynski, Andrzej and Kelly, Daniel L. and Kirby, Keith J. and Kopecky, Martin and Macek, Martin and Malis, Frantisek and Mirtl, Michael and Mitchell, Fraser J. G. and Naaf, Tobias and Newman, Miles and Peterken, George and Petrik, Petr and Schmidt, Wolfgang and Standovar, Tibor and Toth, Zoltan and Van Calster, Hans and Verstraeten, Gorik and Vladovic, Jozef and Vild, Ondrej and Wulf, Monika and Verheyen, Kris}, title = {Drivers of temporal changes in temperate forest plant diversity vary across spatial scales}, series = {Global change biology}, volume = {21}, journal = {Global change biology}, number = {10}, publisher = {Wiley-Blackwell}, address = {Hoboken}, issn = {1354-1013}, doi = {10.1111/gcb.12993}, pages = {3726 -- 3737}, year = {2015}, abstract = {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.}, language = {en} }