@article{PlueDeFrenneAcharyaetal.2017, author = {Plue, Jan and De Frenne, Pieter and Acharya, Kamal and Brunet, J{\"o}rg and Chabrerie, Olivier and Decocq, Guillaume and Diekmann, Martin and Graae, Bente J. and Heinken, Thilo and Hermy, Martin and Kolb, Annette and Lemke, Isgard and Liira, Jaan and Naaf, Tobias and Verheyen, Kris and Wulf, Monika and Cousins, Sara A. O.}, title = {Where does the community start, and where does it end?}, series = {Journal of vegetation science}, volume = {28}, journal = {Journal of vegetation science}, number = {2}, publisher = {Wiley}, address = {Hoboken}, issn = {1100-9233}, doi = {10.1111/jvs.12493}, pages = {424 -- 435}, year = {2017}, abstract = {QuestionBelow-ground processes are key determinants of above-ground plant population and community dynamics. Still, our understanding of how environmental drivers shape plant communities is mostly based on above-ground diversity patterns, bypassing below-ground plant diversity stored in seed banks. As seed banks may shape above-ground plant communities, we question whether concurrently analysing the above- and below-ground species assemblages may potentially enhance our understanding of community responses to environmental variation. LocationTemperate deciduous forests along a 2000km latitudinal gradient in NW Europe. MethodsHerb layer, seed bank and local environmental data including soil pH, canopy cover, forest cover continuity and time since last canopy disturbance were collected in 129 temperate deciduous forest plots. We quantified herb layer and seed bank diversity per plot and evaluated how environmental variation structured community diversity in the herb layer, seed bank and the combined herb layer-seed bank community. ResultsSeed banks consistently held more plant species than the herb layer. How local plot diversity was partitioned across the herb layer and seed bank was mediated by environmental variation in drivers serving as proxies of light availability. The herb layer and seed bank contained an ever smaller and ever larger share of local diversity, respectively, as both canopy cover and time since last canopy disturbance decreased. Species richness and -diversity of the combined herb layer-seed bank community responded distinctly differently compared to the separate assemblages in response to environmental variation in, e.g. forest cover continuity and canopy cover. ConclusionsThe seed bank is a below-ground diversity reservoir of the herbaceous forest community, which interacts with the herb layer, although constrained by environmental variation in e.g. light availability. The herb layer and seed bank co-exist as a single community by means of the so-called storage effect, resulting in distinct responses to environmental variation not necessarily recorded in the individual herb layer or seed bank assemblages. Thus, concurrently analysing above- and below-ground diversity will improve our ecological understanding of how understorey plant communities respond to environmental variation.}, language = {en} } @article{ReineckeKlemmHeinken2014, author = {Reinecke, Jennifer and Klemm, Gunther and Heinken, Thilo}, title = {Vegetation change and homogenization of species composition in temperate nutrient deficient scots pine forests after 45 yr}, series = {Journal of vegetation science}, volume = {25}, journal = {Journal of vegetation science}, number = {1}, publisher = {Wiley-Blackwell}, address = {Hoboken}, issn = {1100-9233}, doi = {10.1111/jvs.12069}, pages = {113 -- 121}, year = {2014}, abstract = {QuestionDoes eutrophication drive vegetation change in pine forests on nutrient deficient sites and thus lead to the homogenization of understorey species composition? LocationForest area (1600ha) in the Lower Spreewald, Brandenburg, Germany. MethodsResurvey of 77 semi-permanent plots after 45yr, including vascular plants, bryophytes and ground lichens. We applied multidimensional ordination of species composition, dissimilarity indices, mean Ellenberg indicator values and the concept of winner/loser species to identify vegetation change between years. Differential responses along a gradient of nutrient availability were analysed on the basis of initial vegetation type, reflecting topsoil N availability of plots. ResultsSpecies composition changed strongly and overall shifted towards higher N and slightly lower light availability. Differences in vegetation change were related to initial vegetation type, with strongest compositional changes in the oligotrophic forest type, but strongest increase of nitrophilous species in the mesotrophic forest type. Despite an overall increase in species number, species composition was homogenized between study years due to the loss of species (mainly ground lichens) on the most oligotrophic sites. ConclusionsThe response to N enrichment is confounded by canopy closure on the N-richest sites and probably by water limitation on N-poorest sites. The relative importance of atmospheric N deposition in the eutrophication effect is difficult to disentangle from natural humus accumulation after historical litter raking. However, the profound differences in species composition between study years across all forest types suggest that atmospheric N deposition contributes to the eutrophication, which drives understorey vegetation change and biotic homogenization in Central European Scots pine forests on nutrient deficient sites.}, language = {en} } @article{deFrenneKolbVerheyenetal.2009, author = {de Frenne, Pieter and Kolb, Annette and Verheyen, Kris and Brunet, Johanne and Chabrerie, Olivier and Decocq, Guillaume and Diekmann, Martin and Eriksson, Ove and Heinken, Thilo and Hermy, Martin and J{\~o}gar, {\"U}lle and Stanton, Sara and Quataert, Paul and Zindel, Renate and Zobel, Martin and Graae, Bente Jessen}, title = {Unravelling the effects of temperature, latitude and local environment on the reproduction of forest herbs}, issn = {1466-822X}, doi = {10.1111/j.1466-8238.2009.00487.x}, year = {2009}, abstract = {Aim To investigate the effect of temperature, latitude and local environment on the reproductive traits of widespread perennial forest herbs to better understand the potential impacts of rising temperatures on their population dynamics and colonization capacities. Location Six regions along a latitudinal gradient from France to Sweden. Methods Within each region, we collected data from three to five populations of up to six species. For each species, several variables were recorded in each region (temperature, latitude) and population (local abiotic and biotic environmental variables), and seed production and germination were estimated. Resource investment in reproduction (RIR) was quantified as seed number ¥ seed mass, while germinable seed output (GSO) was expressed as seed number ¥ germination percentage.We performed linear regression and mixed effect models to investigate the effects of temperature (growing degree hours), latitude and local abiotic and biotic environment on RIR and GSO. Results Temperature and latitude explained most of the variation in RIR and GSO for early flowering species with a northerly distribution range edge (Anemone nemorosa, Paris quadrifolia and Oxalis acetosella). Reproduction of the more southerly distributed species (Brachypodium sylvaticum, Circaea lutetiana and Primula elatior), in contrast, was independent of temperature/latitude. In the late summer species, B. sylvaticum and C. lutetiana, variation in RIR and GSO was best explained by local environmental variables, while none of the investigated variables appeared to be related to reproduction in P. elatior. Main conclusions We showed that reproduction of only two early flowering, northerly distributed species was related to temperature. This suggests that the potential reproductive response of forest herbs to climate warming partly depends on their phenology and distribution, but also that the response is to some extent species dependent. These findings should be taken into account when predictions about future shifts in distribution range are made.}, language = {en} } @article{DeFrenneGraaeBrunetetal.2012, author = {De Frenne, Pieter and Graae, Bente J. and Brunet, J{\"o}rg and Shevtsova, Anna and De Schrijver, An and Chabrerie, Olivier and Cousins, Sara A. O. and Decocq, Guillaume and Diekmann, Martin and Hermy, Martin and Heinken, Thilo and Kolb, Annette and Nilsson, Christer and Stanton, Sharon and Verheyen, Kris}, title = {The response of forest plant regeneration to temperature variation along a latitudinal gradient}, series = {Annals of botany}, volume = {109}, journal = {Annals of botany}, number = {5}, publisher = {Oxford Univ. Press}, address = {Oxford}, issn = {0305-7364}, doi = {10.1093/aob/mcs015}, pages = {1037 -- 1046}, year = {2012}, abstract = {The response of forest herb regeneration from seed to temperature variations across latitudes was experimentally assessed in order to forecast the likely response of understorey community dynamics to climate warming. Seeds of two characteristic forest plants (Anemone nemorosa and Milium effusum) were collected in natural populations along a latitudinal gradient from northern France to northern Sweden and exposed to three temperature regimes in growth chambers (first experiment). To test the importance of local adaptation, reciprocal transplants were also made of adult individuals that originated from the same populations in three common gardens located in southern, central and northern sites along the same gradient, and the resulting seeds were germinated (second experiment). Seedling establishment was quantified by measuring the timing and percentage of seedling emergence, and seedling biomass in both experiments. Spring warming increased emergence rates and seedling growth in the early-flowering forb A. nemorosa. Seedlings of the summer-flowering grass M. effusum originating from northern populations responded more strongly in terms of biomass growth to temperature than southern populations. The above-ground biomass of the seedlings of both species decreased with increasing latitude of origin, irrespective of whether seeds were collected from natural populations or from the common gardens. The emergence percentage decreased with increasing home-away distance in seeds from the transplant experiment, suggesting that the maternal plants were locally adapted. Decreasing seedling emergence and growth were found from the centre to the northern edge of the distribution range for both species. Stronger responses to temperature variation in seedling growth of the grass M. effusum in the north may offer a way to cope with environmental change. The results further suggest that climate warming might differentially affect seedling establishment of understorey plants across their distribution range and thus alter future understorey plant dynamics.}, language = {en} } @article{DeFrenneBrunetShevtsovaetal.2011, author = {De Frenne, Pieter and Brunet, Jorg and Shevtsova, Anna and Kolb, Annette and Graae, Bente J. and Chabrerie, Olivier and Cousins, Sara Ao and Decocq, Guillaume and De Schrijver, An and Diekmann, Martin and Gruwez, Robert and Heinken, Thilo and Hermy, Martin and Nilsson, Christer and Stanton, Sharon and Tack, Wesley and Willaert, Justin and Verheyen, Kris}, title = {Temperature effects on forest herbs assessed by warming and transplant experiments along a latitudinal gradient}, series = {Global change biology}, volume = {17}, journal = {Global change biology}, number = {10}, publisher = {Wiley-Blackwell}, address = {Hoboken}, issn = {1354-1013}, doi = {10.1111/j.1365-2486.2011.02449.x}, pages = {3240 -- 3253}, year = {2011}, abstract = {Slow-colonizing forest understorey plants are probably not able to rapidly adjust their distribution range following large-scale climate change. Therefore, the acclimation potential to climate change within their actual occupied habitats will likely be key for their short-and long-term persistence. We combined transplant experiments along a latitudinal gradient with open-top chambers to assess the effects of temperature on phenology, growth and reproductive performance of multiple populations of slow-colonizing understorey plants, using the spring flowering geophytic forb Anemone nemorosa and the early summer flowering grass Milium effusum as study species. In both species, emergence time and start of flowering clearly advanced with increasing temperatures. Vegetative growth (plant height, aboveground biomass) and reproductive success (seed mass, seed germination and germinable seed output) of A. nemorosa benefited from higher temperatures. Climate warming may thus increase future competitive ability and colonization rates of this species. Apart from the effects on phenology, growth and reproductive performance of M. effusum generally decreased when transplanted southwards (e. g., plant size and number of individuals decreased towards the south) and was probably more limited by light availability in the south. Specific leaf area of both species increased when transplanted southwards, but decreased with open-top chamber installation in A. nemorosa. In general, individuals of both species transplanted at the home site performed best, suggesting local adaptation. We conclude that contrasting understorey plants may display divergent plasticity in response to changing temperatures which may alter future understorey community dynamics.}, language = {en} } @article{WinklerHeinken2007, author = {Winkler, Eckart and Heinken, Thilo}, title = {Spread of an ant-dispersed annual herb : an individual-based simulation study on population development of Melampyrum pratense L.}, issn = {0304-3800}, year = {2007}, abstract = {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.}, language = {en} } @article{HeinkenSchmidtvonOheimbetal.2006, author = {Heinken, Thilo and Schmidt, Marcus and von Oheimb, Goddert and Kriebitzsch, Wolf-Ulrich and Ellenberg, Hermann}, title = {Soil seed banks near rubbing trees indicate dispersal of plant species into forests by wild boar}, issn = {1439-1791}, doi = {10.1016/j.baae.2005.04.006}, year = {2006}, abstract = {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.}, language = {en} } @article{deFrenneGraaeKolbetal.2010, author = {de Frenne, Pieter and Graae, Bente Jessen and Kolb, Annette and Brunet, J{\"o}rg and Chabrerie, Olivier and Cousins, Sara A. O. and Decocq, Guillaume and Dhondt, Rob and Diekmann, Martin and Eriksson, Olof and Heinken, Thilo and Hermy, Martin and J{\"o}gar, uelle and Saguez, Robert and Shevtsova, Anna and Stanton, Sharon and Zindel, Renate and Zobel, Martin and Verheyen, Kris}, title = {Significant effects of temperature on the reproductive output of the forest herb Anemone nemorosa L.}, issn = {0378-1127}, doi = {10.1016/j.foreco.2009.04.038}, year = {2010}, abstract = {Climate warming is already influencing plant migration in different parts of the world. Numerous models have been developed to forecast future plant distributions. Few studies, however, have investigated the potential effect of warming on the reproductive output of plants. Understorey forest herbs in particular, have received little attention in the debate on climate change impacts. This study focuses on the effect of temperature on sexual reproductive output (number of seeds, seed mass, germination percentage and seedling mass) of Anemone nemorosa L., a model species for slow colonizing herbaceous forest plants. We sampled seeds of A. nemorosa in populations along a 2400 km latitudinal gradient from northern France to northern Sweden during three growing seasons (2005,2006 and 2008). This study design allowed us to isolate the effects of accumulated temperature (Growing Degree Hours; GDH) from latitude and the local abiotic and biotic environment. Germination and seed sowing trials were performed in incubators, a greenhouse and under field conditions in a forest. Finally, we disentangled correlations between the different reproductive traits of A. nemorosa along the latitudinal gradient. We found a clear positive relationship between accumulated temperature and seed and seedling traits: reproductive output of A. nemorosa improved with increasing GDH along the latitudinal gradient. Seed mass and seedling mass, for instance, increased by 9.7\% and 10.4\%, respectively, for every 1000 degrees C h increase in GDH. We also derived strong correlations between several seed and seedling traits both under field conditions and in incubators. Our results indicate that seed mass, incubator-based germination percentage (Germ\%(Inc)) and the output of germinable seeds (product of number of seeds and Germ\%(Inc) divided by 100) from plants grown along a latitudinal gradient (i.e. at different temperature regimes) provide valuable proxies to parameterize key population processes in models. We conclude that (1) climate warming may have a pronounced positive impact on sexual reproduction of A. nemorosa and (2) climate models forecasting plant distributions would benefit from including the temperature sensitivity of key seed traits and population processes.}, language = {en} } @article{AlbertAuffretCosynsetal.2015, author = {Albert, Aurelie and Auffret, Alistair G. and Cosyns, Eric and Cousins, Sara A. O. and Eichberg, Carsten and Eycott, Amy E. and Heinken, Thilo and Hoffmann, Maurice and Jaroszewicz, Bogdan and Malo, Juan E. and Marell, Anders and Mouissie, Maarten and Pakeman, Robin J. and Picard, Melanie and Plue, Jan and Poschlod, Peter and Provoost, Sam and Schulze, Kiowa Alraune and Baltzinger, Christophe}, title = {Seed dispersal by ungulates as an ecological filter: a trait-based meta-analysis}, series = {Oikos}, volume = {124}, journal = {Oikos}, number = {9}, publisher = {Wiley-Blackwell}, address = {Hoboken}, issn = {0030-1299}, doi = {10.1111/oik.02512}, pages = {1109 -- 1120}, year = {2015}, abstract = {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.}, language = {en} } @article{DeLombaerdeVerheyenPerringetal.2018, author = {De Lombaerde, Emiel and Verheyen, Kris and Perring, Michael P. and Bernhardt-Roemermann, Markus and Van Calster, Hans and Brunet, Jorg and Chudomelova, Marketa and Decocq, Guillaume and Diekmann, Martin and Durak, Tomasz and Hedl, Radim and Heinken, Thilo and Hommel, Patrick and Jaroszewicz, Bogdan and Kopecky, Martin and Lenoir, Jonathan and Macek, Martin and M{\´a}liš, František and Mitchell, Fraser J. G. and Naaf, Tobias and Newman, Miles and Petř{\´i}k, Petr and Reczyńska, Kamila and Schmidt, Wolfgang and Swierkosz, Krzysztof and Vild, Ondrej and Wulf, Monika and Baetena, Lander}, title = {Responses of competitive understorey species to spatial environmental gradients inaccurately explain temporal changes}, series = {Basic and applied ecology : Journal of the Gesellschaft f{\"u}r {\"O}kologie}, volume = {30}, journal = {Basic and applied ecology : Journal of the Gesellschaft f{\"u}r {\"O}kologie}, publisher = {Elsevier GMBH}, address = {M{\"u}nchen}, issn = {1439-1791}, doi = {10.1016/j.baae.2018.05.013}, pages = {52 -- 64}, year = {2018}, abstract = {Understorey plant communities play a key role in the functioning of forest ecosystems. Under favourable environmental conditions, competitive understorey species may develop high abundances and influence important ecosystem processes such as tree regeneration. Thus, understanding and predicting the response of competitive understorey species as a function of changing environmental conditions is important for forest managers. In the absence of sufficient temporal data to quantify actual vegetation changes, space-for-time (SFT) substitution is often used, i.e. studies that use environmental gradients across space to infer vegetation responses to environmental change over time. Here we assess the validity of such SFT approaches and analysed 36 resurvey studies from ancient forests with low levels of recent disturbances across temperate Europe to assess how six competitive understorey plant species respond to gradients of overstorey cover, soil conditions, atmospheric N deposition and climatic conditions over space and time. The combination of historical and contemporary surveys allows (i) to test if observed contemporary patterns across space are consistent at the time of the historical survey, and, crucially, (ii) to assess whether changes in abundance over time given recorded environmental change match expectations from patterns recorded along environmental gradients in space. We found consistent spatial relationships at the two periods: local variation in soil variables and overstorey cover were the best predictors of individual species' cover while interregional variation in coarse-scale variables, i.e. N deposition and climate, was less important. However, we found that our SFT approach could not accurately explain the large variation in abundance changes over time. We thus recommend to be cautious when using SFT substitution to infer species responses to temporal changes.}, language = {en} } @article{HeinkenRohnerHoppert2007, author = {Heinken, Thilo and Rohner, Maria-Sofie and Hoppert, Michael}, title = {Red wood ants (Formica rufa group) disperse bryophyte and lichen fragments on a local scale}, issn = {0078-2238}, year = {2007}, abstract = {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.}, language = {en} } @article{HeimHoelzelHeinkenetal.2019, author = {Heim, Ramona J. and H{\"o}lzel, Norbert and Heinken, Thilo and Kamp, Johannes and Thomas, Alexander and Darman, Galina F. and Smirenski, Sergei M. and Heim, Wieland}, title = {Post-burn and long-term fire effects on plants and birds in floodplain wetlands of the Russian Far East}, series = {Biodiversity and conservation}, volume = {28}, journal = {Biodiversity and conservation}, number = {6}, publisher = {Springer}, address = {Dordrecht}, issn = {0960-3115}, doi = {10.1007/s10531-019-01746-3}, pages = {1611 -- 1628}, year = {2019}, abstract = {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.}, language = {en} } @article{LemkeKolbGraaeetal.2015, author = {Lemke, Isgard H. and Kolb, Annette and Graae, Bente J. and De Frenne, Pieter and Acharya, Kamal P. and Blandino, Cristina and Brunet, Jorg and Chabrerie, Olivier and Cousins, Sara A. O. and Decocq, Guillaume and Heinken, Thilo and Hermy, Martin and Liira, Jaan and Schmucki, Reto and Shevtsova, Anna and Verheyen, Kris and Diekmann, Martin}, title = {Patterns of phenotypic trait variation in two temperate forest herbs along a broad climatic gradient}, series = {Plant ecology : an international journal}, volume = {216}, journal = {Plant ecology : an international journal}, number = {11}, publisher = {Springer}, address = {Dordrecht}, issn = {1385-0237}, doi = {10.1007/s11258-015-0534-0}, pages = {1523 -- 1536}, year = {2015}, abstract = {Phenotypic trait variation plays a major role in the response of plants to global environmental change, particularly in species with low migration capabilities and recruitment success. However, little is known about the variation of functional traits within populations and about differences in this variation on larger spatial scales. In a first approach, we therefore related trait expression to climate and local environmental conditions, studying two temperate forest herbs, Milium effusum and Stachys sylvatica, along a similar to 1800-2500 km latitudinal gradient. Within each of 9-10 regions in six European countries, we collected data from six populations of each species and recorded several variables in each region (temperature, precipitation) and population (light availability, soil parameters). For each plant, we measured height, leaf area, specific leaf area, seed mass and the number of seeds and examined environmental effects on within-population trait variation as well as on trait means. Most importantly, trait variation differed both between and within populations. Species, however, differed in their response. Intrapopulation variation in Milium was consistently positively affected by higher mean temperatures and precipitation as well as by more fertile local soil conditions, suggesting that more productive conditions may select for larger phenotypic variation. In Stachys, particularly light availability positively influenced trait variation, whereas local soil conditions had no consistent effects. Generally, our study emphasises that intra-population variation may differ considerably across larger scales-due to phenotypic plasticity and/or underlying genetic diversity-possibly affecting species response to global environmental change.}, language = {en} } @article{DiekmannAndresBeckeretal.2019, author = {Diekmann, Martin and Andres, Christian and Becker, Thomas and Bennie, Jonathan and Blueml, Volker and Bullock, James M. and Culmsee, Heike and Fanigliulo, Miriam and Hahn, Annett and Heinken, Thilo and Leuschner, Christoph and Luka, Stefanie and Meissner, Justus and M{\"u}ller, Josef and Newton, Adrian and Peppler-Lisbach, Cord and Rosenthal, Gert and van den Berg, Leon J. L. and Vergeer, Philippine and Wesche, Karsten}, title = {Patterns of long-term vegetation change vary between different types of semi-natural grasslands in Western and Central Europe}, series = {Journal of vegetation science}, volume = {30}, journal = {Journal of vegetation science}, number = {2}, publisher = {Wiley}, address = {Hoboken}, issn = {1100-9233}, doi = {10.1111/jvs.12727}, pages = {187 -- 202}, year = {2019}, abstract = {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.}, language = {en} } @article{GraaeDeFrenneKolbetal.2012, author = {Graae, Bente J. and De Frenne, Pieter and Kolb, Annette and Brunet, Jorg and Chabrerie, Olivier and Verheyen, Kris and Pepin, Nick and Heinken, Thilo and Zobel, Martin and Shevtsova, Anna and Nijs, Ivan and Milbau, Ann}, title = {On the use of weather data in ecological studies along altitudinal and latitudinal gradients}, series = {Oikos}, volume = {121}, journal = {Oikos}, number = {1}, publisher = {Wiley-Blackwell}, address = {Hoboken}, issn = {0030-1299}, doi = {10.1111/j.1600-0706.2011.19694.x}, pages = {3 -- 19}, year = {2012}, abstract = {Global warming has created a need for studies along climatic gradients to assess the effects of temperature on ecological processes. Altitudinal and latitudinal gradients are often used as such, usually in combination with air temperature data from the closest weather station recorded at 1.52 m above the ground. However, many ecological processes occur in, at, or right above the soil surface. To evaluate how representative the commonly used weather station data are for the microclimate relevant for soil surface biota, we compared weather station temperatures for an altitudinal (500900 m a.s.l.) and a latitudinal gradient (4968 degrees N) with data obtained by temperature sensors placed right below the soil surface at five sites along these gradients. The mean annual temperatures obtained from weather stations and adjusted using a lapse rate of -5.5 degrees C km-1 were between 3.8 degrees C lower and 1.6 degrees C higher than those recorded by the temperature sensors at the soil surface, depending on the position along the gradients. The monthly mean temperatures were up to 10 degrees C warmer or 5 degrees C colder at the soil surface. The within-site variation in accumulated temperature was as high as would be expected from a 300 m change in altitude or from a 4 degrees change in latitude or a climate change scenario corresponding to warming of 1.63.8 degrees C. Thus, these differences introduced by the decoupling are significant from a climate change perspective, and the results demonstrate the need for incorporating microclimatic variation when conducting studies along altitudinal or latitudinal gradients. We emphasize the need for using relevant temperature data in climate impact studies and further call for more studies describing the soil surface microclimate, which is crucial for much of the biota.}, language = {en} } @article{KurzeHeinkenFartmann2017, author = {Kurze, Susanne and Heinken, Thilo and Fartmann, Thomas}, title = {Nitrogen enrichment of host plants has mostly beneficial effects on the life-history traits of nettle-feeding butterflies}, series = {Acta oecologica : international journal of ecology}, volume = {85}, journal = {Acta oecologica : international journal of ecology}, publisher = {Elsevier}, address = {Amsterdam}, issn = {1146-609X}, doi = {10.1016/j.actao.2017.11.005}, pages = {157 -- 164}, year = {2017}, abstract = {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.}, language = {en} } @article{KurzeHeinkenFartmann2018, author = {Kurze, Susanne and Heinken, Thilo and Fartmann, Thomas}, title = {Nitrogen enrichment in host plants increases the mortality of common Lepidoptera species}, series = {Oecologia}, volume = {188}, journal = {Oecologia}, number = {4}, publisher = {Springer}, address = {New York}, issn = {0029-8549}, doi = {10.1007/s00442-018-4266-4}, pages = {1227 -- 1237}, year = {2018}, abstract = {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.}, language = {en} } @article{HeinkenZippel2004, author = {Heinken, Thilo and Zippel, Elke}, title = {Natural re-colonization of experimental gaps by terricolous bryophytes in Central European pine forests}, issn = {0029-5035}, year = {2004}, abstract = {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}, language = {en} } @article{DeFrenneRodriguezSanchezCoomesetal.2013, author = {De Frenne, Pieter and Rodriguez-Sanchez, Francisco and Coomes, David Anthony and B{\"a}ten, Lander and Verstr{\"a}ten, Gorik and Vellend, Mark and Bernhardt-R{\"o}mermann, Markus and Brown, Carissa D. and Brunet, J{\"o}rg and Cornelis, Johnny and Decocq, Guillaume M. and Dierschke, Hartmut and Eriksson, Ove and Gilliam, Frank S. and Hedl, Radim and Heinken, Thilo and Hermy, Martin and Hommel, Patrick and Jenkins, Michael A. and Kelly, Daniel L. and Kirby, Keith J. and Mitchell, Fraser J. G. and Naaf, Tobias and Newman, Miles and Peterken, George and Petrik, Petr and Schultz, Jan and Sonnier, Gregory and Van Calster, Hans and Waller, Donald M. and Walther, Gian-Reto and White, Peter S. and Woods, Kerry D. and Wulf, Monika and Graae, Bente Jessen and Verheyen, Kris}, title = {Microclimate moderates plant responses to macroclimate warming}, series = {Proceedings of the National Academy of Sciences of the United States of America}, volume = {110}, journal = {Proceedings of the National Academy of Sciences of the United States of America}, number = {46}, publisher = {National Acad. of Sciences}, address = {Washington}, issn = {0027-8424}, doi = {10.1073/pnas.1311190110}, pages = {18561 -- 18565}, year = {2013}, abstract = {Recent global warming is acting across marine, freshwater, and terrestrial ecosystems to favor species adapted to warmer conditions and/or reduce the abundance of cold-adapted organisms (i.e., "thermophilization" of communities). Lack of community responses to increased temperature, however, has also been reported for several taxa and regions, suggesting that "climatic lags" may be frequent. Here we show that microclimatic effects brought about by forest canopy closure can buffer biotic responses to macroclimate warming, thus explaining an apparent climatic lag. Using data from 1,409 vegetation plots in European and North American temperate forests, each surveyed at least twice over an interval of 12-67 y, we document significant thermophilization of ground-layer plant communities. These changes reflect concurrent declines in species adapted to cooler conditions and increases in species adapted to warmer conditions. However, thermophilization, particularly the increase of warm-adapted species, is attenuated in forests whose canopies have become denser, probably reflecting cooler growing-season ground temperatures via increased shading. As standing stocks of trees have increased in many temperate forests in recent decades, local microclimatic effects may commonly be moderating the impacts of macroclimate warming on forest understories. Conversely, increases in harvesting woody biomass-e.g., for bioenergy-may open forest canopies and accelerate thermophilization of temperate forest biodiversity.}, language = {en} } @article{MaesBlondeelPerringetal.2019, author = {Maes, Sybryn L. and Blondeel, Haben and Perring, Michael P. and Depauw, Leen and Brumelis, Guntis and Brunet, J{\"o}rg and Decocq, Guillaume and den Ouden, Jan and Haerdtle, Werner and Hedl, Radim and Heinken, Thilo and Heinrichs, Steffi and Jaroszewicz, Bogdan and Kirby, Keith J. and Kopecky, Martin and Malis, Frantisek and Wulf, Monika and Verheyen, Kris}, title = {Litter quality, land-use history, and nitrogen deposition effects on topsoil conditions across European temperate deciduous forests}, series = {Forest ecology and management}, volume = {433}, journal = {Forest ecology and management}, publisher = {Elsevier}, address = {Amsterdam}, issn = {0378-1127}, doi = {10.1016/j.foreco.2018.10.056}, pages = {405 -- 418}, year = {2019}, abstract = {Topsoil conditions in temperate forests are influenced by several soil-forming factors, such as canopy composition (e.g. through litter quality), land-use history, atmospheric deposition, and the parent material. Many studies have evaluated the effects of single factors on physicochemical topsoil conditions, but few have assessed the simultaneous effects of multiple drivers. Here, we evaluate the combined effects of litter quality, land-use history (past land cover as well as past forest management), and atmospheric deposition on several physicochemical topsoil conditions of European temperate deciduous forest soils: bulk density, proportion of exchangeable base cations, carbon/nitrogen-ratio (C/N), litter mass, bio-available and total phosphorus, pH(KCI)and soil organic matter. We collected mineral soil and litter layer samples, and measured site characteristics for 190 20 x 20 m European mixed forest plots across gradients of litter quality (derived from the canopy species composition) and atmospheric deposition, and for different categories of past land cover and past forest management. We accounted for the effects of parent material on topsoil conditions by clustering our plots into three soil type groups based on texture and carbonate concentration. We found that litter quality was a stronger driver of topsoil conditions compared to land-use history or atmospheric deposition, while the soil type also affected several topsoil conditions here. Plots with higher litter quality had soils with a higher proportion of exchangeable base cations, and total phosphorus, and lower C/N-ratios and litter mass. Furthermore, the observed litter quality effects on the topsoil were independent from the regional nitrogen deposition or the soil type, although the soil type likely (co)-determined canopy composition and thus litter quality to some extent in the investigated plots. Litter quality effects on topsoil phosphorus concentrations did interact with past land cover, highlighting the need to consider land-use history when evaluating canopy effects on soil conditions. We conclude that forest managers can use the canopy composition as an important tool for influencing topsoil conditions, although soil type remains an important factor to consider.}, language = {en} }