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 - Schmidt, M. A1 - von Oheimb, Goddert A1 - Kriebitzsch, Wolf-Ulrich A1 - Ellenberg, H. T1 - Ausbreitung von Pflanzen durch Schalenwild Y1 - 2005 SN - 0936-1294 - 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 -