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 - Diekmann, Martin A1 - Andres, Christian A1 - Becker, Thomas A1 - Bennie, Jonathan A1 - Blueml, Volker A1 - Bullock, James M. A1 - Culmsee, Heike A1 - Fanigliulo, Miriam A1 - Hahn, Annett A1 - Heinken, Thilo A1 - Leuschner, Christoph A1 - Luka, Stefanie A1 - Meissner, Justus A1 - Müller, Josef A1 - Newton, Adrian A1 - Peppler-Lisbach, Cord A1 - Rosenthal, Gert A1 - van den Berg, Leon J. L. A1 - Vergeer, Philippine A1 - Wesche, Karsten T1 - Patterns of long-term vegetation change vary between different types of semi-natural grasslands in Western and Central Europe JF - Journal of vegetation science N2 - Questions Has plant species richness in semi-natural grasslands changed over recent decades? Do the temporal trends of habitat specialists differ from those of habitat generalists? Has there been a homogenization of the grassland vegetation? Location Different regions in Germany and the UK. Methods We conducted a formal meta-analysis of re-survey vegetation studies of semi-natural grasslands. In total, 23 data sets were compiled, spanning up to 75 years between the surveys, including 13 data sets from wet grasslands, six from dry grasslands and four from other grassland types. Edaphic conditions were assessed using mean Ellenberg indicator values for soil moisture, nitrogen and pH. Changes in species richness and environmental variables were evaluated using response ratios. Results In most wet grasslands, total species richness declined over time, while habitat specialists almost completely vanished. The number of species losses increased with increasing time between the surveys and were associated with a strong decrease in soil moisture and higher soil nutrient contents. Wet grasslands in nature reserves showed no such changes or even opposite trends. In dry grasslands and other grassland types, total species richness did not consistently change, but the number or proportions of habitat specialists declined. There were also considerable changes in species composition, especially in wet grasslands that often have been converted into intensively managed, highly productive meadows or pastures. We did not find a general homogenization of the vegetation in any of the grassland types. Conclusions The results document the widespread deterioration of semi-natural grasslands, especially of those types that can easily be transformed to high production grasslands. The main causes for the loss of grassland specialists are changed management in combination with increased fertilization and nitrogen deposition. Dry grasslands are most resistant to change, but also show a long-term trend towards an increase in more mesotrophic species. KW - dry grasslands KW - fragmentation KW - homogenization KW - management KW - meta-analysis KW - nitrogen deposition KW - quasi-permanent plot KW - re-survey KW - species richness KW - wet grasslands Y1 - 2019 U6 - https://doi.org/10.1111/jvs.12727 SN - 1100-9233 SN - 1654-1103 VL - 30 IS - 2 SP - 187 EP - 202 PB - Wiley CY - Hoboken ER - TY - JOUR A1 - Kurze, Susanne A1 - Heinken, Thilo A1 - Fartmann, Thomas T1 - Nitrogen enrichment in host plants increases the mortality of common Lepidoptera species JF - Oecologia N2 - The recent decline of Lepidoptera species strongly correlates with the increasing intensification of agriculture in Western and Central Europe. However, the effects of changed host-plant quality through agricultural fertilization on this insect group remain largely unexplored. For this reason, we tested the response of six common butterfly and moth species to host-plant fertilization using fertilizer quantities usually applied in agriculture. The larvae of the study species Coenonympha pamphilus, Lycaena phlaeas, Lycaena tityrus, Pararge aegeria, Rivula sericealis and Timandra comae were distributed according to a split-brood design to three host-plant treatments comprising one control treatment without fertilization and two fertilization treatments with an input of 150 and 300kgNha(-1)year(-1), respectively. In L.tityrus, we used two additional fertilization treatments with an input of 30 and 90kgNha(-1)year(-1), respectively. Fertilization increased the nitrogen concentration of both host-plant species, Rumex acetosella and Poa pratensis, and decreased the survival of larvae in all six Lepidoptera species by at least one-third, without clear differences between sorrel- and grass-feeding species. The declining survival rate in all species contradicts the well-accepted nitrogen-limitation hypothesis, which predicts a positive response in species performance to dietary nitrogen content. In contrast, this study presents the first evidence that current fertilization quantities in agriculture exceed the physiological tolerance of common Lepidoptera species. Our results suggest that (1) the negative effect of plant fertilization on Lepidoptera has previously been underestimated and (2) that it contributes to the range-wide decline of Lepidoptera. KW - Agricultural fertilization KW - Global change KW - Host-plant quality KW - Nitrogen-limitation hypothesis KW - Rearing experiment Y1 - 2018 U6 - https://doi.org/10.1007/s00442-018-4266-4 SN - 0029-8549 SN - 1432-1939 VL - 188 IS - 4 SP - 1227 EP - 1237 PB - Springer CY - New York ER - TY - JOUR A1 - Schöpke, Benito A1 - Heinze, Johannes A1 - Pätzig, Marlene A1 - Heinken, Thilo T1 - Do dispersal traits of wetland plant species explain tolerance against isolation effects in naturally fragmented habitats? JF - Plant ecology : an international journal N2 - The effects of habitat fragmentation and isolation on plant species richness have been verified for a wide range of anthropogenically fragmented habitats, but there is currently little information about their effects in naturally small and isolated habitats. We tested whether habitat area, heterogeneity, and isolation affect the richness of wetland vascular plant species in kettle holes, i.e., small glacially created wetlands, in an agricultural landscape of 1 km(2) in NE Germany. We compared fragmentation effects with those of forest fragments in the same landscape window. Since wetland and forest species might differ in their tolerance to isolation, and because isolation effects on plant species may be trait dependent, we asked which key life history traits might foster differences in isolation tolerance between wetland and forest plants. We recorded the flora and vegetation types in 83 isolated sites that contained 81 kettle holes and 25 forest fragments. Overall, the number of wetland species increased with increasing area and heterogeneity, i.e., the number of vegetation types, while area was not a surrogate for heterogeneity in these naturally fragmented systems. Isolation did not influence the number of wetland species but decreased the number of forest species. We also found that seeds of wetland species were on average lighter, more persistent and better adapted to epizoochory, e.g., by waterfowl, than seeds of forest species. Therefore, we suggest that wetland species are more tolerant to isolation than forest species due to their higher dispersal potential in space and time, which may counterbalance the negative effects of isolation. KW - Forest species KW - Habitat fragmentation KW - Isolation KW - Kettle holes KW - Life history traits KW - Wetland species Y1 - 2019 U6 - https://doi.org/10.1007/s11258-019-00955-8 SN - 1385-0237 SN - 1573-5052 VL - 220 IS - 9 SP - 801 EP - 815 PB - Springer CY - Dordrecht ER - TY - JOUR A1 - Kurze, Susanne A1 - Heinken, Thilo A1 - Fartmann, Thomas T1 - Nitrogen enrichment of host plants has mostly beneficial effects on the life-history traits of nettle-feeding butterflies JF - Acta oecologica : international journal of ecology N2 - Butterflies rank among the most threatened animal groups throughout Europe. However, current population trends differ among species. The nettle-feeding butterflies Aglais io and Aglais urticae cope successfully with the anthropogenic land-use change. Both species are assumed to be pre-adapted to higher nitrogen contents in their host plant, stinging nettle (Urtica dioica). However, it is currently unknown, whether this pre-adaptation enables both Aglais species to cope successfully or even to benefit from the excessive nitrogen availabilities in nettles growing in modern farmlands. For this reason, this study focused on the response of both Aglais species to unfertilized nettles compared to nettles receiving 150 or 300 kg N ha(-1) yr(-1) (i.e., common fertilizer quantities of modern-day agriculture). Fertilized nettles were characterized by higher nitrogen concentrations and lower C:N ratios compared to the control group. In both Aglais species, the individuals feeding on fertilized nettles had higher survival rates, shorter larval periods and heavier pupae and, in A. urticae also longer forewings. All these trait shifts are beneficial for the individuals, lowering their risk to die before reproduction and increasing their reproductive potential. These responses agree with the well-accepted nitrogen-limitation hypothesis predicting a positive relationship between the nitrogen content of the diet and the performance of herbivorous insects. Furthermore, our findings suggest that the increasing abundance of both Aglais species may result not only from the increasing spread of nettles into the farmland but also from changes in their quality due to the eutrophication of the landscape during recent decades. KW - Aglais KW - Fertilization KW - Host-plant quality KW - Landscape eutrophication KW - Nitrogen-limitation hypothesis KW - Rearing experiment Y1 - 2017 U6 - https://doi.org/10.1016/j.actao.2017.11.005 SN - 1146-609X SN - 1873-6238 VL - 85 SP - 157 EP - 164 PB - Elsevier CY - Amsterdam ER - TY - JOUR A1 - De Frenne, Pieter A1 - Blondeel, H. A1 - Brunet, J. A1 - Caron, M. M. A1 - Chabrerie, O. A1 - Cougnon, M. A1 - Cousins, S. A. O. A1 - Decocq, G. A1 - Diekmann, M. A1 - Graae, B. J. A1 - Hanley, M. E. A1 - Heinken, Thilo A1 - Hermy, M. A1 - Kolb, A. A1 - Lenoir, J. A1 - Liira, J. A1 - Orczewska, A. A1 - Shevtsova, A. A1 - Vanneste, T. A1 - Verheyen, K. T1 - Atmospheric nitrogen deposition on petals enhances seed quality of the forest herb Anemone nemorosa JF - Plant biology N2 - Elevated atmospheric input of nitrogen (N) is currently affecting plant biodiversity and ecosystem functioning. The growth and survival of numerous plant species is known to respond strongly to N fertilisation. Yet, few studies have assessed the effects of N deposition on seed quality and reproductive performance, which is an important life-history stage of plants. Here we address this knowledge gap by assessing the effects of atmospheric N deposition on seed quality of the ancient forest herb Anemone nemorosa using two complementary approaches. By taking advantage of the wide spatiotemporal variation in N deposition rates in pan-European temperate and boreal forests over 2years, we detected positive effects of N deposition on the N concentration (percentage N per unit seed mass, increased from 2.8% to 4.1%) and N content (total N mass per seed more than doubled) of A.nemorosa seeds. In a complementary experiment, we applied ammonium nitrate to aboveground plant tissues and the soil surface to determine whether dissolved N sources in precipitation could be incorporated into seeds. Although the addition of N to leaves and the soil surface had no effect, a concentrated N solution applied to petals during anthesis resulted in increased seed mass, seed N concentration and N content. Our results demonstrate that N deposition on the petals enhances bioaccumulation of N in the seeds of A.nemorosa. Enhanced atmospheric inputs of N can thus not only affect growth and population dynamics via root or canopy uptake, but can also influence seed quality and reproduction via intake through the inflorescences. KW - Latitudinal gradient KW - nitrogen deposition KW - nutrient stoichiometry KW - seed provisioning KW - seed quality KW - sexual reproduction KW - wood anemone Y1 - 2018 U6 - https://doi.org/10.1111/plb.12688 SN - 1435-8603 SN - 1438-8677 VL - 20 IS - 3 SP - 619 EP - 626 PB - Wiley CY - Hoboken ER - TY - JOUR A1 - Heinken, Thilo A1 - Schmidt, Marcus A1 - von Oheimb, Goddert A1 - Kriebitzsch, Wolf-Ulrich A1 - Ellenberg, Hermann T1 - Soil seed banks near rubbing trees indicate dispersal of plant species into forests by wild boar N2 - Current knowledge about processes that generate long-distance dispersal of plants is still limited despite its importance for persistence of populations and colonization of new potential habitats. Today wild Large mammals are presumed to be important vectors for long-distance transport of diaspores within and between European temperate forest patches, and in particular wild boars recently came into focus. Here we use a specific habit of wild boar, i.e. wallowing in mud and subsequent rubbing against trees, to evaluate epizoochorous dispersal of vascular plant diaspores. We present soil seed bank data from 27 rubbing trees versus 27 control trees from seven forest areas in Germany. The mean number of viable seeds and the plant species number were higher in soil samples near rubbing trees compared with control trees. Ten of the 20 most frequent species were more frequent, and many species exclusively appeared in the soil samples near rubbing trees. The large number of plant species and seeds - more than 1000 per tree - in the soils near rubbing trees is difficult to explain unless the majority were dispersed by wild boar. Hooked and bristly diaspores, i.e. those adapted to epizoochory, were more frequent; however, many species with unspecialized diaspores occurred exclusively near rubbing trees. As opposed to plant species closely tied to forests species which occur both in forest and open vegetation and non-forest species were more frequent near rubbing trees compared with controls. These findings are consistent with previous studies on diaspore loads in the coats and hooves of shot wild boars. However, our method allows to identify the transport of diaspores from the open landscape into forest stands, where they might especially emerge after disturbance, and a clustered distribution of epizoochorically dispersed seeds. Moreover, accumulation of seeds of wetness indicators near rubbing trees demonstrates directed dispersal of plant species inhabiting wet places among remote wallows. Y1 - 2006 UR - http://www.sciencedirect.com/science/journal/14391791 U6 - https://doi.org/10.1016/j.baae.2005.04.006 SN - 1439-1791 ER - TY - JOUR A1 - Heinken, Thilo A1 - Zippel, Elke T1 - Natural re-colonization of experimental gaps by terricolous bryophytes in Central European pine forests N2 - In northeastern German pine forests we studied re-colonization patterns of experimental gaps by four dominant bryophyte species (Dicranum scoparium, Hypnum jutlandicum, Pleurozium schreberi and Scleropodium purum) over three years. Both vegetation and litter layer were removed on 1 m(2) plots within +/- pure colonies of the experimental species, while the humus layer was left intact. All plots were vegetatively re-colonized by the species which was dominant before gap creation. Three mechanisms of re-colonization occurred and interacted: (1) advance of surrounding shoots from the edge into the gaps by clonal growth, (2) dispersal of detached single shoots as well as larger clumps of multiple shoots into the plots, resulting in new colonies by continuing growth, and (3) regeneration from a soil diaspore bank consisting of seemingly dead stem fragments in the humus layer of the gaps. Scleropodium purum, which occurs at locations with good water and nutrient supply, displayed the most rapid growth. Here, some plots were completely recovered after three years. Despite lower rates of advance from the edge, colonization of Hypnum jutlandicum was faster than and of Dicranum scoparium as fast than that of Pleurozium schreberi because of a larger diaspore bank. Thus, each bryophyte species was characterized by a different habitat occupation strategy. The different clonal colonization strategies account for the high competitive capacity and regeneration potential of the investigated bryophyte species in pine forests despite of the lack of generative reproduction. Experimental disturbance resulted in a temporary increase of bryophyte diversity, because short-lived Colonists with a low competitive capacity colonized the gaps, before they will be overgrown by the dominant Perennials Y1 - 2004 SN - 0029-5035 ER - TY - JOUR A1 - Plue, Jan A1 - De Frenne, Pieter A1 - Acharya, Kamal A1 - Brunet, Jörg 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 - Verheyen, Kris A1 - Wulf, Monika A1 - Cousins, Sara A. O. T1 - Where does the community start, and where does it end? BT - including the seed bank to reassess forest herb layer responses to the environment JF - Journal of vegetation science N2 - 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. KW - Above-ground KW - Below-ground KW - Canopy KW - Disturbance KW - Diversity KW - Light availability KW - NWEurope KW - Plant community KW - Species co-existence KW - Storage effect Y1 - 2017 U6 - https://doi.org/10.1111/jvs.12493 SN - 1100-9233 SN - 1654-1103 VL - 28 IS - 2 SP - 424 EP - 435 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 - Perring, Michael P. A1 - Bernhardt-Roemermann, Markus A1 - Baeten, Lander A1 - Midolo, Gabriele A1 - Blondeel, Haben A1 - Depauw, Leen A1 - Landuyt, Dries A1 - Maes, Sybryn L. A1 - De Lombaerde, Emiel A1 - Caron, Maria Mercedes A1 - Vellend, Mark A1 - Brunet, Joerg A1 - Chudomelova, Marketa A1 - Decocq, Guillaume A1 - Diekmann, Martin A1 - Dirnboeck, Thomas A1 - Doerfler, Inken A1 - Durak, Tomasz A1 - De Frenne, Pieter A1 - Gilliam, Frank S. A1 - Hedl, Radim A1 - Heinken, Thilo A1 - Hommel, Patrick A1 - Jaroszewicz, Bogdan A1 - Kirby, Keith J. A1 - Kopecky, Martin A1 - Lenoir, Jonathan A1 - Li, Daijiang A1 - Malis, Frantisek A1 - Mitchell, Fraser J. G. A1 - Naaf, Tobias A1 - Newman, Miles A1 - Petrik, Petr A1 - Reczynska, Kamila A1 - Schmidt, Wolfgang A1 - Standovar, Tibor A1 - Swierkosz, Krzysztof A1 - Van Calster, Hans A1 - Vild, Ondrej A1 - Wagner, Eva Rosa A1 - Wulf, Monika A1 - Verheyen, Kris T1 - Global environmental change effects on plant community composition trajectories depend upon management legacies JF - Global change biology N2 - 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. KW - biodiversity change KW - climate change KW - disturbance regime KW - forestREplot KW - herbaceous layer KW - management intensity KW - nitrogen deposition KW - plant functional traits KW - time lag KW - vegetation resurvey Y1 - 2017 U6 - https://doi.org/10.1111/gcb.14030 SN - 1354-1013 SN - 1365-2486 VL - 24 IS - 4 SP - 1722 EP - 1740 PB - Wiley CY - Hoboken ER - TY - JOUR A1 - Heim, Ramona J. A1 - Hölzel, Norbert A1 - Heinken, Thilo A1 - Kamp, Johannes A1 - Thomas, Alexander A1 - Darman, Galina F. A1 - Smirenski, Sergei M. A1 - Heim, Wieland T1 - Post-burn and long-term fire effects on plants and birds in floodplain wetlands of the Russian Far East JF - Biodiversity and conservation N2 - Wildfires affect biodiversity at multiple levels. While vegetation is directly changed by fire events, animals are often indirectly affected through changes in habitat and food availability. Globally, fire frequency and the extent of fires are predicted to increase in the future. The impact of fire on the biodiversity of temperate wetlands has gained little attention so far. We compared species richness and abundance of plants and birds in burnt and unburnt areas in the Amur floodplain/Russian Far East in the year of fire and 1 year after. We also analysed vegetation recovery in relation to time since fire over a period of 18 years. Plant species richness was higher in burnt compared to unburnt plots in the year of the fire, but not in the year after. This suggests that fire has a positive short-term effect on plant diversity. Bird species richness and abundance were lower on burnt compared to unburnt plots in the year of the fire, but not in the year after. Over a period of 18 years, high fire frequency led to an increase in herb cover and a decrease in grass cover. We show that the effects on biodiversity are taxon- and species-specific. Fire management strategies in temperate wetlands should consider fire frequency as a key driving force of vegetation structure, with carry-over effects on higher trophic levels. Designing fire refuges, i.e., areas that do not burn annually, might locally be necessary to maintain high species richness. KW - Disturbance KW - Bird species richness KW - Vegetation structure KW - Fire frequency KW - Amur River Y1 - 2019 U6 - https://doi.org/10.1007/s10531-019-01746-3 SN - 0960-3115 SN - 1572-9710 VL - 28 IS - 6 SP - 1611 EP - 1628 PB - Springer CY - Dordrecht ER - TY - JOUR A1 - Maes, Sybryn L. A1 - Blondeel, Haben A1 - Perring, Michael P. A1 - Depauw, Leen A1 - Brumelis, Guntis A1 - Brunet, Jörg A1 - Decocq, Guillaume A1 - den Ouden, Jan A1 - Haerdtle, Werner A1 - Hedl, Radim A1 - Heinken, Thilo A1 - Heinrichs, Steffi A1 - Jaroszewicz, Bogdan A1 - Kirby, Keith J. A1 - Kopecky, Martin A1 - Malis, Frantisek A1 - Wulf, Monika A1 - Verheyen, Kris T1 - Litter quality, land-use history, and nitrogen deposition effects on topsoil conditions across European temperate deciduous forests JF - Forest ecology and management N2 - 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. KW - Soil fertility KW - Ancient forest KW - Post-agricultural forest KW - Coppice KW - High forest KW - pH KW - Phosphorus KW - Base cations KW - Nutrient cycling KW - Decomposition Y1 - 2019 U6 - https://doi.org/10.1016/j.foreco.2018.10.056 SN - 0378-1127 SN - 1872-7042 VL - 433 SP - 405 EP - 418 PB - Elsevier CY - Amsterdam 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 - TY - JOUR A1 - Heinken, Thilo A1 - Weber, Ewald T1 - Consequences of habitat fragmentation for plant species do we know enough? JF - Perspectives in plant ecology, evolution and systematics N2 - Habitat fragmentation is one of the most important causes for the decline of plant species. However, plants differing in phylogeny, habitat requirements and biology are likely to respond differently to habitat fragmentation. We ask whether case studies on the effects of habitat fragmentation conducted so far allow generalizations about its effects on the fitness and genetic diversity of populations of endangered plant species. We compared the characteristics of plant species endangered in Germany whose sensitivity to habitat fragmentation had been studied with those of the endangered species that had not been studied. We found strong discrepancies between the two groups with regard to their taxonomy and traits relevant to their sensitivity to habitat fragmentation. Monocots, graminoids, clonal, abiotically pollinated and self compatible species were underrepresented among the studied species, and most study species were from a few habitat types, in particular grasslands. We conclude that our current knowledge of the effects of habitat fragmentation on plant populations is not sufficient to provide widely applicable guidelines for species management. The selection of species studied so far has been biased toward species from certain habitats and species exhibiting traits that probably make them vulnerable to habitat fragmentation. Future studies should include community-wide approaches in different habitats, e.g. re-visitation studies in which the species pool is assessed at different time intervals, and population-biological studies of species from a wide range of habitats, and of different life forms and growth strategies. A more representative picture of the effects of habitat fragmentation would allow a better assessment of threats and more specific recommendations for optimally managing populations of endangered plants. KW - CSR-strategies KW - Naturally rare species KW - Plant conservation KW - Recently rare species KW - Species traits KW - Taxonomic position Y1 - 2013 U6 - https://doi.org/10.1016/j.ppees.2013.05.003 SN - 1433-8319 VL - 15 IS - 4 SP - 205 EP - 216 PB - Elsevier CY - Jena ER - TY - JOUR A1 - de Frenne, Pieter A1 - Graae, Bente Jessen A1 - Kolb, Annette A1 - Brunet, Jörg A1 - Chabrerie, Olivier A1 - Cousins, Sara A. O. A1 - Decocq, Guillaume A1 - Dhondt, Rob A1 - Diekmann, Martin A1 - Eriksson, Olof A1 - Heinken, Thilo A1 - Hermy, Martin A1 - Jögar, uelle A1 - Saguez, Robert A1 - Shevtsova, Anna A1 - Stanton, Sharon A1 - Zindel, Renate A1 - Zobel, Martin A1 - Verheyen, Kris T1 - Significant effects of temperature on the reproductive output of the forest herb Anemone nemorosa L. N2 - 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. Y1 - 2010 UR - http://www.sciencedirect.com/science/journal/03781127 U6 - https://doi.org/10.1016/j.foreco.2009.04.038 SN - 0378-1127 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 - Heinken, Thilo A1 - Rohner, Maria-Sofie A1 - Hoppert, Michael T1 - Red wood ants (Formica rufa group) disperse bryophyte and lichen fragments on a local scale N2 - Gametophyte and thallus fragments, respectively, may be an important or even the only mode of reproduction for many bryophytes and lichens species. Until now especially birds and mammals have been identifi ed as potential animal dispersal vectors of fragments. This study investigates the dispersal of bryophyte and lichen fragments by red wood ants which build large nest mounds from plant material and are abundant in European coniferous forests. We sampled nest material from 25 nest mounds in fi ve different pine and spruce forest types in Germany and found numerous fragments of 20 bryophyte and ten lichen species. As they occurred on almost all studied mounds and often in large numbers we conclude that collecting cryptogam fragments as nest material is a characteristic feature for the Formica rufa group in coniferous forests. Species number and composition of fragments on mounds coincided with the epigeic vegetation around ant nests to a large extent: Almost all collected species were present in the vegetation, and dominant fragment species occurred in large amounts in the vicinity of ant nests. Lichen fragments were larger than bryophyte fragments. Certain life forms (weft-forming bryophytes, reindeer lichens) were accumulated on mounds, while others (tall turfs, cup-type Cladonia species) were discriminated, refl ecting fragmentation features of species. Collected fragments may regenerate to mature plants if nest mounds are abandoned, and especially if they are lost during transport over several metres. We conclude that dispersal of fragments by red wood ants contributes to maintain epigeic bryophyte and lichen diversity of coniferous forests by supporting colonisation after disturbances, which occur on different spatial and temporal scales. Y1 - 2007 SN - 0078-2238 ER - TY - JOUR A1 - Schwarzer, Christian A1 - Heinken, Thilo A1 - Luthardt, Vera A1 - Joshi, Jasmin Radha T1 - Latitudinal shifts in species interactions interfere with resistance of southern but not of northern bog-plant communities to experimental climate change JF - The journal of ecology N2 - The persistence of species under changed climatic conditions depends on adaptations and plastic responses to these conditions and on interactions with their local plant community resulting in direct and indirect effects of changed climatic conditions. Populations at species' range margins may be especially crucial in containing a gene pool comprising adaptations to extreme climatic conditions. Many species of northern European bog ecosystems reach their southern lowland range limit in central Europe. In a common-garden experiment, we experimentally assessed the impact of projected climatic changes on five bog-plant species (including peat moss Sphagnum magellanicum) sampled along a latitudinal gradient of 1400km from Scandinavia to the marginal lowland populations in Germany. Populations were cultivated in monocultures and in experimental communities composed of all five species from their local community, and exposed to five combinations of three climate treatments (warming, fluctuating water-tables, fertilization) in a southern common garden. Whereas most monocultures showed a decreasing biomass production from southern to northern origins under southern environmental conditions, in the experimental mixed-species communities, an increasing biomass production towards northern communities was observed together with a shift in interspecific interactions along the latitudinal gradient. While negative dominance effects prevailed in southern communities, higher net biodiversity effects were observed in northern subarctic communities. The combined effects of climate treatments increased biomass production in monocultures of most origins. In communities, however, overall the treatments did not result in significantly changed biomass production. Among individual treatments, water-table fluctuations caused a significant decrease in biomass production, but only in southern communities, indicating higher vulnerability to changed climatic conditions. Here, negative effects of climate treatments on graminoids were not compensated by the slightly increased growth of peat moss that benefited from interspecific interactions only in northern communities.Synthesis. We conclude that shifting interactions within multispecies communities caused pronounced responses to changed climatic conditions in wetland communities of temperate southern marginal, but not of northern subarctic origin. Therefore, future models investigating the impacts of climate change on plant communities should consider geographical variation in species interactions an important factor influencing community responses to changed climatic conditions. KW - additive partitioning of biodiversity effects KW - biodiversity KW - ecosystem services KW - ecosystem stability KW - intraspecific divergence KW - multifactorial environmental change KW - nitrogen deposition KW - northern peatlands KW - Sphagnum magellanicum KW - wetland ecosystems Y1 - 2013 U6 - https://doi.org/10.1111/1365-2745.12158 SN - 0022-0477 SN - 1365-2745 VL - 101 IS - 6 SP - 1484 EP - 1497 PB - Wiley-Blackwell CY - Hoboken ER - TY - JOUR A1 - De Frenne, Pieter A1 - Rodriguez-Sanchez, Francisco A1 - Coomes, David Anthony A1 - Bäten, Lander A1 - Versträten, Gorik A1 - Vellend, Mark A1 - Bernhardt-Römermann, Markus A1 - Brown, Carissa D. A1 - Brunet, Jörg A1 - Cornelis, Johnny A1 - Decocq, Guillaume M. A1 - Dierschke, Hartmut A1 - Eriksson, Ove A1 - Gilliam, Frank S. A1 - Hedl, Radim A1 - Heinken, Thilo A1 - Hermy, Martin A1 - Hommel, Patrick A1 - Jenkins, Michael A. A1 - Kelly, Daniel L. A1 - Kirby, Keith J. A1 - Mitchell, Fraser J. G. A1 - Naaf, Tobias A1 - Newman, Miles A1 - Peterken, George A1 - Petrik, Petr A1 - Schultz, Jan A1 - Sonnier, Gregory A1 - Van Calster, Hans A1 - Waller, Donald M. A1 - Walther, Gian-Reto A1 - White, Peter S. A1 - Woods, Kerry D. A1 - Wulf, Monika A1 - Graae, Bente Jessen A1 - Verheyen, Kris T1 - Microclimate moderates plant responses to macroclimate warming JF - Proceedings of the National Academy of Sciences of the United States of America N2 - 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. KW - climate change KW - forest management KW - understory KW - climatic debt KW - range shifts Y1 - 2013 U6 - https://doi.org/10.1073/pnas.1311190110 SN - 0027-8424 VL - 110 IS - 46 SP - 18561 EP - 18565 PB - National Acad. of Sciences CY - Washington ER -