TY - JOUR A1 - Graae, Bente Jessen A1 - Verheyen, Kris A1 - Kolb, Annette A1 - van der Veken, Sebastian A1 - Heinken, Thilo A1 - Chabrerie, Olivier A1 - Diekmann, Martin A1 - Valtinat, Karin A1 - Zindel, Renate A1 - Karlsson, Elisabeth A1 - Ström, Lotta A1 - Decocq, Guillaume A1 - Hermy, Martin A1 - Baskin, Carol C. T1 - Germination requirements and seed mass of slow- and fast-colonizing temperate forest herbs along a latitudinal gradient N2 - 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. Y1 - 2009 UR - http://www.bioone.org/loi/ecos U6 - https://doi.org/10.2980/16-2-3234 SN - 1195-6860 ER - TY - JOUR A1 - Winkler, Eckart A1 - Heinken, Thilo T1 - Spread of an ant-dispersed annual herb : an individual-based simulation study on population development of Melampyrum pratense L. N2 - The paper presents a simulation and parameter-estimation approach for evaluating stochastic patterns of population growth and spread of an annual forest herb, Melampyrum pratense (Orobanchaceae). The survival of a species during large-scale changes in land use and climate will depend, to a considerable extent, on its dispersal and colonisation abilities. Predictions on species migration need a combination of field studies and modelling efforts. Our study on the ability of M. pratense to disperse into so far unoccupied areas is based on experiments in secondary woodland in NE Germany. Experiments started in 1997 at three sites where the species was not yet present, with 300 seeds sown within 1m2. Population development was then recorded until 2001 by mapping of individuals with a resolution of 5 cm. Additional observations considered density dependence of seed production. We designed a spatially explicit individual-based computer simulation model to explain the spatial patterns of population development and to predict future population spread. Besides primary drop of seeds (barochory) it assumed secondary seed transport by ants (myrmecochory) with an exponentially decreasing dispersal tail. An important feature of population-pattern explanation was the simultaneous estimation of both population-growth and dispersal parameters from consistent spatio-temporal data sets. As the simulation model produced stochastic time series and random spatially discrete distributions of individuals we estimated parameters by minimising the expectation of weighted sum of squares. These sums of squares criteria considered population sizes, radial population distributions around the area of origin and distributions of individuals within squares of 25cm×25 cm, the range of density action. Optimal parameter values, together with the precision of the estimates, were obtained from calculating sum of squares in regular grids of parameter values. Our modelling results showed that transport of fractions of seeds by ants over distances of 1-2m was indispensable for explaining the observed population spread that led to distances of at most 8mfrom population origin within 3 years. Projections of population development over four additional years gave a diffusion-like increase of population area without any "outposts". This prediction generated by the simulation model gave a hypothesis which should be revised by additional field observations. Some structural deviations between observations and model output already indicated that for full understanding of population spread the set of dispersal mechanisms assumed in the model may have to be extended by additional features of plant-animal mutualism. Y1 - 2007 UR - http://www.sciencedirect.com/science/journal/03043800 SN - 0304-3800 ER - TY - JOUR A1 - Liang, Wei A1 - Heinrich, Ingo A1 - Simard, Sonia A1 - Helle, Gerhard A1 - Linan, Isabel Dorado A1 - Heinken, Thilo T1 - Climate signals derived from cell anatomy of Scots pine in NE Germany JF - Tree physiology N2 - Tree-ring chronologies of Pinus sylvestris L. from latitudinal and altitudinal limits of the species distribution have been widely used for climate reconstructions, but there are many sites within the temperate climate zone, as is the case in northeastern Germany, at which there is little evidence of a clear climate signal in the chronologies. In this study, we developed long chronologies of several cell structure variables (e. g., average lumen area and cell wall thickness) from P. sylvestris growing in northeastern Germany and investigated the influence of climate on ring widths and cell structure variables. We found significant correlations between cell structure variables and temperature, and between tree-ring width and relative humidity and vapor pressure, respectively, enabling the development of robust reconstructions from temperate sites that have not yet been realized. Moreover, it has been shown that it may not be necessary to detrend chronologies of cell structure variables and thus low-frequency climate signals may be retrieved from longer cell structure chronologies. The relatively extensive resource of archaeological material of P. sylvestris covering approximately the last millennium may now be useful for climate reconstructions in northeastern Germany and other sites in the temperate climate zone. KW - cell structure KW - dendroclimatology KW - Pinus sylvestris KW - quantitative wood anatomy KW - tree rings Y1 - 2013 U6 - https://doi.org/10.1093/treephys/tpt059 SN - 0829-318X SN - 1758-4469 VL - 33 IS - 8 SP - 833 EP - 844 PB - Oxford Univ. Press CY - Oxford ER - TY - JOUR A1 - Verheyen, Kris A1 - Baeten, Lander A1 - De Frenne, Pieter A1 - Bernhardt-Römermann, Markus A1 - Brunet, Jorg A1 - Cornelis, Johnny A1 - Decocq, Guillaume A1 - Dierschke, Hartmut A1 - Eriksson, Ove A1 - Hedl, Radim A1 - Heinken, Thilo A1 - Hermy, Martin A1 - Hommel, Patrick A1 - Kirby, Keith J. A1 - Naaf, Tobias A1 - Peterken, George A1 - Petrik, Petr A1 - Pfadenhauer, Joerg A1 - Van Calster, Hans A1 - Walther, Gian-Reto A1 - Wulf, Monika A1 - Verstraeten, Gorik T1 - Driving factors behind the eutrophication signal in understorey plant communities of deciduous temperate forests JF - The journal of ecology N2 - 1. Atmospheric nitrogen (N) deposition is expected to change forest understorey plant community composition and diversity, but results of experimental addition studies and observational studies are not yet conclusive. A shortcoming of observational studies, which are generally based on resurveys or sampling along large deposition gradients, is the occurrence of temporal or spatial confounding factors. 2. We were able to assess the contribution of N deposition versus other ecological drivers on forest understorey plant communities by combining a temporal and spatial approach. Data from 1205 (semi-)permanent vegetation plots taken from 23 rigorously selected understorey resurvey studies along a large deposition gradient across deciduous temperate forest in Europe were compiled and related to various local and regional driving factors, including the rate of atmospheric N deposition, the change in large herbivore densities and the change in canopy cover and composition. 3. Although no directional change in species richness occurred, there was considerable floristic turnover in the understorey plant community and a shift in species composition towards more shade-tolerant and nutrient-demanding species. However, atmospheric N deposition was not important in explaining the observed eutrophication signal. This signal seemed mainly related to a shift towards a denser canopy cover and a changed canopy species composition with a higher share of species with more easily decomposed litter. 4. Synthesis. Our multi-site approach clearly demonstrates that one should be cautious when drawing conclusions about the impact of atmospheric N deposition based on the interpretation of plant community shifts in single sites or regions due to other, concurrent, ecological changes. Even though the effects of chronically increased N deposition on the forest plant communities are apparently obscured by the effects of canopy changes, the accumulated N might still have a significant impact. However, more research is needed to assess whether this N time bomb will indeed explode when canopies will open up again. KW - atmospheric deposition KW - determinants of plant community diversity and structure KW - Ellenberg indicator values KW - forest herbs KW - forest management KW - large herbivores KW - north-western Europe KW - resurveys KW - (semi-)permanent plots Y1 - 2012 U6 - https://doi.org/10.1111/j.1365-2745.2011.01928.x SN - 0022-0477 VL - 100 IS - 2 SP - 352 EP - 365 PB - Wiley-Blackwell CY - Hoboken ER - TY - JOUR A1 - Schulze, Kiowa Alraune A1 - Buchwald, Rainer A1 - Heinken, Thilo T1 - Epizoochory via the hooves - the European bison (Bison bonasus L.) as a dispersal agent of seeds in an open-forest-mosaic JF - Tuexenia : Mitteilungen der Floristisch-Soziologischen Arbeitsgemeinschaft N2 - Large herbivores are significant vectors for the long-distance dispersal of seeds in various habitats, both attached to animals (epizoochory) and via gut passage (endozoochory). The majority of studies on epizoochory have examined dispersal in the fur of domesticated ungulates. Studies on wild ungulates are important to understand dispersal processes in many habitats, but rare due to methodological constraints. We studied epizoochory of seeds by European bison in an open-forest-mosaic (nutrient-poor grassland and heathland, mixed forest) in NW Germany, where bison had been introduced for the purpose of nature conservation. At the study site it was possible to apply a method by which hoof material of free-ranging bison was non-invasively collected. We identified a total of 1082 seeds from 32 plant species in the hoof material. The three most abundant species were Polygonum aviculare, Agrostis capillaris and Betula spp. Seed species originated from various habitat types of the study area, while the majority of seeds derived from trampled areas. Compared to the non-dispersed plant species of the study area, dispersed plant species had a higher seed longevity index, suggesting that many seeds were picked up from the soil seed bank. Epizoochory ranking indices of dispersed seed species, classifying the importance of epizoochory, revealed that transport in the fur may be of minor importance for many species, i.e. epizoochory by the hooves turned out to be negatively correlated to epizoochory in the fur. We conclude that European bison disperses a considerable number of seed species through trampling. Further research should consider epizoochory via the hooves and include integrative approaches to understand the different dispersal mechanisms by ungulates and their long-term synergetic effect on plant communities. KW - diaspore KW - external dispersal KW - large ungulate KW - secondary seed dispersal KW - trampling Y1 - 2014 SN - 0722-494X IS - 34 SP - 131 EP - 144 PB - Floristisch-Soziologische Arbeitsgemeinschaft CY - Göttingen ER - TY - JOUR A1 - Caron, Maria Mercedes A1 - De Frenne, Pieter A1 - Brunet, Jörg A1 - Chabrerie, Olivier A1 - Cousins, Sara A. O. A1 - Decocq, Guillaume A1 - Diekmann, Martin A1 - Graae, Bente Jessen A1 - Heinken, Thilo A1 - Kolb, Annette A1 - Lenoir, Jonathan A1 - Naaf, Tobias A1 - Plue, Jan A1 - Selvi, Federico A1 - Wulf, Monika A1 - Verheyen, Kris T1 - Divergent regeneration responses of two closely related tree species to direct abiotic and indirect biotic effects of climate change JF - Forest ecology and management N2 - 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. KW - Acer KW - Regeneration KW - Latitudinal gradient KW - Temperature KW - Precipitation KW - Competition Y1 - 2015 U6 - https://doi.org/10.1016/j.foreco.2015.01.003 SN - 0378-1127 SN - 1872-7042 VL - 342 SP - 21 EP - 29 PB - Elsevier CY - Amsterdam ER - TY - JOUR A1 - De Frenne, Pieter A1 - Graae, Bente J. A1 - Brunet, Jörg A1 - Shevtsova, Anna A1 - De Schrijver, An A1 - Chabrerie, Olivier A1 - Cousins, Sara A. O. A1 - Decocq, Guillaume A1 - Diekmann, Martin A1 - Hermy, Martin A1 - Heinken, Thilo A1 - Kolb, Annette A1 - Nilsson, Christer A1 - Stanton, Sharon A1 - Verheyen, Kris T1 - The response of forest plant regeneration to temperature variation along a latitudinal gradient JF - Annals of botany N2 - 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. KW - Anemone nemorosa KW - climate change KW - common garden KW - growth chambers KW - latitudinal gradient KW - local adaptation KW - Milium effusum KW - plant regeneration KW - range edges KW - recruitment KW - seedling establishment KW - temperature Y1 - 2012 U6 - https://doi.org/10.1093/aob/mcs015 SN - 0305-7364 VL - 109 IS - 5 SP - 1037 EP - 1046 PB - Oxford Univ. Press CY - Oxford ER - TY - JOUR A1 - Baeten, Lander A1 - Warton, David I. A1 - Van Calster, Hans A1 - De Frenne, Pieter A1 - Verstraeten, Gorik A1 - Bonte, Dries A1 - Bernhardt-Römermann, Markus A1 - Cornelis, Johnny A1 - Decocq, Guillaume A1 - Eriksson, Ove A1 - Hedl, Radim A1 - Heinken, Thilo A1 - Hermy, Martin A1 - Hommel, Patrick A1 - Kirby, Keith J. A1 - Naaf, Tobias A1 - Petrik, Petr A1 - Walther, Gian-Reto A1 - Wulf, Monica A1 - Verheyen, Kris T1 - A model-based approach to studying changes in compositional heterogeneity JF - Methods in ecology and evolution : an official journal of the British Ecological Society Y1 - 2014 SN - 2041-210X SN - 2041-2096 VL - 5 IS - 2 SP - 156 EP - 164 PB - Wiley-Blackwell CY - Hoboken ER - TY - JOUR A1 - De Frenne, P. A1 - Kolb, Annette A1 - Graae, Benete Jessen A1 - Decocq, Guillaume A1 - Baltora, S. A1 - De Schrijver, A. A1 - Brunet, J. A1 - Chabrerie, Olivier A1 - Cousins, Sara A. O. A1 - Dhondt, Rob A1 - Diekmann, Martin A1 - Gruwez, R. A1 - Heinken, Thilo A1 - Hermy, Martin A1 - Liira, J. A1 - Saguez, R. A1 - Shevtsova, Anna A1 - Baskin, Carol C. A1 - Verheyen, Kris T1 - A latitudinal gradient in seed nutrients of the forest herb Anemone nemorosa JF - Plant biology N2 - 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. KW - Collection date KW - latitude KW - nutrient stoichiometry KW - seed nitrogen KW - seed predation KW - seed provisioning KW - sexual reproduction KW - wood anemone Y1 - 2011 U6 - https://doi.org/10.1111/j.1438-8677.2010.00404.x SN - 1435-8603 VL - 13 IS - 3 SP - 493 EP - 501 PB - Wiley-Blackwell CY - Malden ER - TY - JOUR A1 - Albert, Aurelie A1 - Auffret, Alistair G. A1 - Cosyns, Eric A1 - Cousins, Sara A. O. A1 - Eichberg, Carsten A1 - Eycott, Amy E. A1 - Heinken, Thilo A1 - Hoffmann, Maurice A1 - Jaroszewicz, Bogdan A1 - Malo, Juan E. A1 - Marell, Anders A1 - Mouissie, Maarten A1 - Pakeman, Robin J. A1 - Picard, Melanie A1 - Plue, Jan A1 - Poschlod, Peter A1 - Provoost, Sam A1 - Schulze, Kiowa Alraune A1 - Baltzinger, Christophe T1 - Seed dispersal by ungulates as an ecological filter: a trait-based meta-analysis JF - Oikos N2 - Plant communities are often dispersal-limited and zoochory can be an efficient mechanism for plants to colonize new patches of potentially suitable habitat. We predicted that seed dispersal by ungulates acts as an ecological filter - which differentially affects individuals according to their characteristics and shapes species assemblages - and that the filter varies according to the dispersal mechanism (endozoochory, fur-epizoochory and hoof-epizoochory). We conducted two-step individual participant data meta-analyses of 52 studies on plant dispersal by ungulates in fragmented landscapes, comparing eight plant traits and two habitat indicators between dispersed and non-dispersed plants. We found that ungulates dispersed at least 44% of the available plant species. Moreover, some plant traits and habitat indicators increased the likelihood for plant of being dispersed. Persistent or nitrophilous plant species from open habitats or bearing dry or elongated diaspores were more likely to be dispersed by ungulates, whatever the dispersal mechanism. In addition, endozoochory was more likely for diaspores bearing elongated appendages whereas epizoochory was more likely for diaspores released relatively high in vegetation. Hoof-epizoochory was more likely for light diaspores without hooked appendages. Fur-epizoochory was more likely for diaspores with appendages, particularly elongated or hooked ones. We thus observed a gradient of filtering effect among the three dispersal mechanisms. Endozoochory had an effect of rather weak intensity (impacting six plant characteristics with variations between ungulate-dispersed and non-dispersed plant species mostly below 25%), whereas hoof-epizoochory had a stronger effect (eight characteristics included five ones with above 75% variation), and fur-epizoochory an even stronger one (nine characteristics included six ones with above 75% variation). Our results demonstrate that seed dispersal by ungulates is an ecological filter whose intensity varies according to the dispersal mechanism considered. Ungulates can thus play a key role in plant community dynamics and have implications for plant spatial distribution patterns at multiple scales. Y1 - 2015 U6 - https://doi.org/10.1111/oik.02512 SN - 0030-1299 SN - 1600-0706 VL - 124 IS - 9 SP - 1109 EP - 1120 PB - Wiley-Blackwell CY - Hoboken ER -