@article{VerheyenBaetenDeFrenneetal.2012,
  author    = {Verheyen, Kris and Baeten, Lander and De Frenne, Pieter and Bernhardt-R{\"o}mermann, Markus and Brunet, Jorg and Cornelis, Johnny and Decocq, Guillaume and Dierschke, Hartmut and Eriksson, Ove and Hedl, Radim and Heinken, Thilo and Hermy, Martin and Hommel, Patrick and Kirby, Keith J. and Naaf, Tobias and Peterken, George and Petrik, Petr and Pfadenhauer, Joerg and Van Calster, Hans and Walther, Gian-Reto and Wulf, Monika and Verstraeten, Gorik},
  title     = {Driving factors behind the eutrophication signal in understorey plant communities of deciduous temperate forests},
  series = {The journal of ecology},
  volume    = {100},
  journal   = {The journal of ecology},
  number    = {2},
  publisher = {Wiley-Blackwell},
  address   = {Hoboken},
  issn      = {0022-0477},
  doi       = {10.1111/j.1365-2745.2011.01928.x},
  pages     = {352 -- 365},
  year      = {2012},
  abstract  = {1. Atmospheric nitrogen (N) deposition is expected to change forest understorey plant community composition and diversity, but results of experimental addition studies and observational studies are not yet conclusive. A shortcoming of observational studies, which are generally based on resurveys or sampling along large deposition gradients, is the occurrence of temporal or spatial confounding factors. 2. We were able to assess the contribution of N deposition versus other ecological drivers on forest understorey plant communities by combining a temporal and spatial approach. Data from 1205 (semi-)permanent vegetation plots taken from 23 rigorously selected understorey resurvey studies along a large deposition gradient across deciduous temperate forest in Europe were compiled and related to various local and regional driving factors, including the rate of atmospheric N deposition, the change in large herbivore densities and the change in canopy cover and composition. 3. Although no directional change in species richness occurred, there was considerable floristic turnover in the understorey plant community and a shift in species composition towards more shade-tolerant and nutrient-demanding species. However, atmospheric N deposition was not important in explaining the observed eutrophication signal. This signal seemed mainly related to a shift towards a denser canopy cover and a changed canopy species composition with a higher share of species with more easily decomposed litter. 4. Synthesis. Our multi-site approach clearly demonstrates that one should be cautious when drawing conclusions about the impact of atmospheric N deposition based on the interpretation of plant community shifts in single sites or regions due to other, concurrent, ecological changes. Even though the effects of chronically increased N deposition on the forest plant communities are apparently obscured by the effects of canopy changes, the accumulated N might still have a significant impact. However, more research is needed to assess whether this N time bomb will indeed explode when canopies will open up again.},
  language  = {en}
}
@article{VannesteValdesVerheyenetal.2018,
  author    = {Vanneste, Thomas and Valdes, Alicia and Verheyen, Kris and Perring, Michael P. and Bernhardt-Roemermann, Markus and Andrieu, Emilie and Brunet, Jorg and Cousins, Sara A. O. and Deconchat, Marc and De Smedt, Pallieter and Diekmann, Martin and Ehrmann, Steffen and Heinken, Thilo and Hermy, Martin and Kolb, Annette and Lenoir, Jonathan and Liira, Jaan and Naaf, Tobias and Paal, Taavi and Wulf, Monika and Decocq, Guillaume and De Frenne, Pieter},
  title     = {Functional trait variation of forest understorey plant communities across Europe},
  series = {Basic and applied ecology : Journal of the Gesellschaft f{\"u}r {\"O}kologie},
  volume    = {34},
  journal   = {Basic and applied ecology : Journal of the Gesellschaft f{\"u}r {\"O}kologie},
  publisher = {Elsevier GmbH},
  address   = {M{\"u}nchen},
  issn      = {1439-1791},
  doi       = {10.1016/j.baae.2018.09.004},
  pages     = {1 -- 14},
  year      = {2018},
  abstract  = {Global environmental changes are expected to alter the functional characteristics of understorey herb-layer communities, potentially affecting forest ecosystem functioning. However, little is known about what drives the variability of functional traits in forest understories. Here, we assessed the role of different environmental drivers in shaping the functional trait distribution of understorey herbs in fragmented forests across three spatial scales. We focused on 708 small, deciduous forest patches located in 16 agricultural landscape windows, spanning a 2500-km macroclimatic gradient across the temperate forest biome in Europe. We estimated the relative effect of patch-scale, landscape-scale and macroclimatic variables on the community mean and variation of plant height, specific leaf area and seed mass. Macroclimatic variables (monthly temperature and precipitation extremes) explained the largest proportion of variation in community trait means (on average 77\% of the explained variation). In contrast, patch-scale factors dominated in explaining community trait variation (on average 68\% of the explained variation). Notably, patch age, size and internal heterogeneity had a positive effect on the community-level variability. Landscape-scale variables explained only a minor part of the variation in both trait distribution properties. The variation explained by shared combinations of the variable groups was generally negligible. These findings highlight the importance of considering multiple spatial scales in predictions of environmental-change effects on the functionality of forest understories. We propose that forest management sustainability could benefit from conserving larger, historically continuous and internally heterogeneous forest patches to maximise ecosystem service diversity in rural landscapes. (C) 2018 Gesellschaft fur Okologie. Published by Elsevier GmbH. All rights reserved.},
  language  = {en}
}
@article{PlueDeFrenneAcharyaetal.2013,
  author    = {Plue, Jan and De Frenne, Pieter and Acharya, Kamal P. and Brunet, Jorg and Chabrerie, Olivier and Decocq, Guillaume and Diekmann, Martin and Graae, Bente J. and Heinken, Thilo and Hermy, Martin and Kolb, Annette and Lemke, Isgard and Liira, Jaan and Naaf, Tobias and Shevtsova, Anna and Verheyen, Kris and Wulf, Monika and Cousins, Sara A. O.},
  title     = {Climatic control of forest herb seed banks along a latitudinal gradient},
  series = {Global ecology and biogeography : a journal of macroecology},
  volume    = {22},
  journal   = {Global ecology and biogeography : a journal of macroecology},
  number    = {10},
  publisher = {Wiley-Blackwell},
  address   = {Hoboken},
  issn      = {1466-822X},
  doi       = {10.1111/geb.12068},
  pages     = {1106 -- 1117},
  year      = {2013},
  abstract  = {Aim Seed banks are central to the regeneration strategy of many plant species. Any factor altering seed bank density thus affects plant regeneration and population dynamics. Although seed banks are dynamic entities controlled by multiple environmental drivers, climatic factors are the most comprehensive, but still poorly understood. This study investigates how climatic variation structures seed production and resulting seed bank patterns. Location Temperate forests along a 1900km latitudinal gradient in north-western (NW) Europe. Methods Seed production and seed bank density were quantified in 153 plots along the gradient for four forest herbs with different seed longevity: Geum urbanum, Milium effusum, Poa nemoralis and Stachys sylvatica. We tested the importance of climatic and local environmental factors in shaping seed production and seed bank density. Results Seed production was determined by population size, and not by climatic factors. G.urbanum and M.effusum seed bank density declined with decreasing temperature (growing degree days) and/or increasing temperature range (maximum-minimum temperature). P.nemoralis and S.sylvatica seed bank density were limited by population size and not by climatic variables. Seed bank density was also influenced by other, local environmental factors such as soil pH or light availability. Different seed bank patterns emerged due to differential seed longevities. Species with long-lived seeds maintained constant seed bank densities by counteracting the reduced chance of regular years with high seed production at colder northern latitudes. Main conclusions Seed bank patterns show clear interspecific variation in response to climate across the distribution range. Not all seed banking species may be as well equipped to buffer climate change via their seed bank, notably in short-term persistent species. Since the buffering capacity of seed banks is key to species persistence, these results provide crucial information to advance climatic change predictions on range shifts, community and biodiversity responses.},
  language  = {en}
}
@article{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{PerringBernhardtRoemermannBaetenetal.2018,
  author    = {Perring, Michael P. and Bernhardt-Roemermann, Markus and Baeten, Lander and Midolo, Gabriele and Blondeel, Haben and Depauw, Leen and Landuyt, Dries and Maes, Sybryn L. and De Lombaerde, Emiel and Caron, Maria Mercedes and Vellend, Mark and Brunet, Joerg and Chudomelova, Marketa and Decocq, Guillaume and Diekmann, Martin and Dirnboeck, Thomas and Doerfler, Inken and Durak, Tomasz and De Frenne, Pieter and Gilliam, Frank S. and Hedl, Radim and Heinken, Thilo and Hommel, Patrick and Jaroszewicz, Bogdan and Kirby, Keith J. and Kopecky, Martin and Lenoir, Jonathan and Li, Daijiang and Malis, Frantisek and Mitchell, Fraser J. G. and Naaf, Tobias and Newman, Miles and Petrik, Petr and Reczynska, Kamila and Schmidt, Wolfgang and Standovar, Tibor and Swierkosz, Krzysztof and Van Calster, Hans and Vild, Ondrej and Wagner, Eva Rosa and Wulf, Monika and Verheyen, Kris},
  title     = {Global environmental change effects on plant community composition trajectories depend upon management legacies},
  series = {Global change biology},
  volume    = {24},
  journal   = {Global change biology},
  number    = {4},
  publisher = {Wiley},
  address   = {Hoboken},
  issn      = {1354-1013},
  doi       = {10.1111/gcb.14030},
  pages     = {1722 -- 1740},
  year      = {2018},
  abstract  = {The contemporary state of functional traits and species richness in plant communities depends on legacy effects of past disturbances. Whether temporal responses of community properties to current environmental changes are altered by such legacies is, however, unknown. We expect global environmental changes to interact with land-use legacies given different community trajectories initiated by prior management, and subsequent responses to altered resources and conditions. We tested this expectation for species richness and functional traits using 1814 survey-resurvey plot pairs of understorey communities from 40 European temperate forest datasets, syntheses of management transitions since the year 1800, and a trait database. We also examined how plant community indicators of resources and conditions changed in response to management legacies and environmental change. Community trajectories were clearly influenced by interactions between management legacies from over 200 years ago and environmental change. Importantly, higher rates of nitrogen deposition led to increased species richness and plant height in forests managed less intensively in 1800 (i.e., high forests), and to decreases in forests with a more intensive historical management in 1800 (i.e., coppiced forests). There was evidence that these declines in community variables in formerly coppiced forests were ameliorated by increased rates of temperature change between surveys. Responses were generally apparent regardless of sites' contemporary management classifications, although sometimes the management transition itself, rather than historic or contemporary management types, better explained understorey responses. Main effects of environmental change were rare, although higher rates of precipitation change increased plant height, accompanied by increases in fertility indicator values. Analysis of indicator values suggested the importance of directly characterising resources and conditions to better understand legacy and environmental change effects. Accounting for legacies of past disturbance can reconcile contradictory literature results and appears crucial to anticipating future responses to global environmental change.},
  language  = {en}
}
@article{NaafWulf2012,
  author    = {Naaf, Tobias and Wulf, Monika},
  title     = {Plant community assembly in temperate forests along gradients of soil fertility and disturbance},
  series = {Acta oecologica : international journal of ecology},
  volume    = {39},
  journal   = {Acta oecologica : international journal of ecology},
  number    = {2},
  publisher = {Elsevier},
  address   = {Paris},
  issn      = {1146-609X},
  doi       = {10.1016/j.actao.2012.01.009},
  pages     = {101 -- 108},
  year      = {2012},
  abstract  = {Plant community assembly from a regional pool is largely driven by two mechanisms: environmental filtering and niche partitioning, which result in trait convergence or divergence, respectively. Although empirical evidence for both assembly mechanisms exists, the environmental conditions and traits where each of the two assembly patterns is prevalent remain unclear. We studied community assembly mechanisms in herb layer communities of temperate forest patches in NW Germany, looking at distributions of competitive and reproductive traits along gradients of soil fertility and disturbance. We also examined how community assembly patterns changed over a time span of two decades. Canopy height converged toward taller species with increasing soil fertility and increasing light availability. Most reproductive traits diverged with an increasing degree of disturbance and with increasing fertility. Comparisons over time indicated that disturbance events induced the coexistence of species with different reproductive strategies and also selected for tall species as a result of enhanced competitive pressure. Our study demonstrates that in accordance with existing hypotheses, competitive traits (e.g., canopy height) can be convergent in favorable environments. However, this convergence is associated with a divergence of traits related to other challenges (e.g., reproduction), indicating that true functional redundancy within communities does not exist. Moreover, our study shows that the expected divergence of reproductive traits at disturbed sites can be accompanied by a convergence of other traits (e.g., canopy height), indicating that several assembly mechanisms can operate simultaneously.},
  language  = {en}
}
@article{NaafWulf2011,
  author    = {Naaf, Tobias and Wulf, Monika},
  title     = {Traits of winner and loser species indicate drivers of herb layer changes over two decades in forests of NW Germany},
  series = {Journal of vegetation science},
  volume    = {22},
  journal   = {Journal of vegetation science},
  number    = {3},
  publisher = {Wiley-Blackwell},
  address   = {Malden},
  issn      = {1100-9233},
  doi       = {10.1111/j.1654-1103.2011.01267.x},
  pages     = {516 -- 527},
  year      = {2011},
  abstract  = {Questions What are the most likely environmental drivers for compositional herb layer changes as indicated by trait differences between winner and loser species? Location Weser-Elbe region (NW Germany). Methods We resurveyed the herb layer communities of ancient forest patches on base-rich sites of 175 semi-permanent plots. Species traits were tested for their ability to discriminate between winner and loser species using logistic regression analyses and deviance partitioning. Results Of 115 species tested, 31 were identified as winner species and 30 as loser species. Winner species had higher seed longevity, flowered later in the season and more often had an oceanic distribution compared to loser species. Loser species tended to have a higher specific leaf area, were more susceptible to deer browsing and had a performance optimum at higher soil pH compared to winner species. The loser species also represented several ancient forest and threatened species. Deviance partitioning indicated that local drivers (i.e. disturbance due to forest management) were primarily responsible for the species shifts, while regional drivers (i.e. browsing pressure and acidification from atmospheric deposition) and global drivers (i.e. climate warming) had moderate effects. There was no evidence that canopy closure, drainage or eutrophication contributed to herb layer changes. Conclusions The relative importance of the different drivers as indicated by the winner and loser species differs from that found in previous long-term studies. Relating species traits to species performance is a valuable tool that provides insight into the environmental drivers that are most likely responsible for herb layer changes.},
  language  = {en}
}
@article{NaafWulf2012,
  author    = {Naaf, Tobias and Wulf, Monika},
  title     = {Does taxonomic homogenization imply functional homogenization in temperate forest herb layer communities?},
  series = {Plant ecology : an international journal},
  volume    = {213},
  journal   = {Plant ecology : an international journal},
  number    = {3},
  publisher = {Springer},
  address   = {Dordrecht},
  issn      = {1385-0237},
  doi       = {10.1007/s11258-011-9990-3},
  pages     = {431 -- 443},
  year      = {2012},
  abstract  = {Biotic homogenization, the decrease in beta diversity among formerly distinct species assemblages, has been recognized as an important form of biotic impoverishment for more than a decade. Although researchers have stressed the importance of the functional dimension to understand its potential ecological consequences, biotic homogenization has mostly been studied at a taxonomic level. Here, we explore the relationship between taxonomic and functional homogenization using data on temperate forest herb layer communities in NW Germany, for which taxonomic homogenization has recently been demonstrated. We quantified beta diversity by partitioning Rao's quadratic entropy. We found a general positive relationship between changes in taxonomic and functional beta diversity. This relationship was stronger if multiple functional traits were taken into account. Averaged across sites, however, taxonomic homogenization was not consistently accompanied by functional homogenization. Depending on the traits considered, taxonomic homogenization occurred also together with functional differentiation or no change in functional beta diversity. The species shifts responsible for changes in beta diversity differed substantially between taxonomic and functional beta diversity measures and also among functional beta diversity measures based on different traits. We discuss likely environmental drivers for species shifts. Our study demonstrates that functional homogenization must be explicitly studied as an independent phenomenon that cannot be inferred from taxonomic homogenization.},
  language  = {en}
}
@phdthesis{Naaf2011,
  author    = {Naaf, Tobias},
  title     = {Floristic homogenization and impoverishment : herb layer changes over two decades in deciduous forest patches of the Weser-Elbe region (NW Germany)},
  url       = {http://nbn-resolving.de/urn:nbn:de:kobv:517-opus-52446},
  school      = {Universit{\"a}t Potsdam},
  year      = {2011},
  abstract  = {Human-induced alterations of the environment are causing biotic changes worldwide, including the extinction of species and a mixing of once disparate floras and faunas. One type of biological communities that is expected to be particularly affected by environmental alterations are herb layer plant communities of fragmented forests such as those in the west European lowlands. However, our knowledge about current changes in species diversity and composition in these communities is limited due to a lack of adequate long-term studies. In this thesis, I resurveyed the herb layer communities of ancient forest patches in the Weser-Elbe region (NW Germany) after two decades using 175 semi-permanent plots. The general objectives were (i) to quantify changes in plant species diversity considering also between-community (β) and functional diversity, (ii) to determine shifts in species composition in terms of species' niche breadth and functional traits and (iii) to find indications on the most likely environmental drivers for the observed changes. These objectives were pursued with four independent research papers (Chapters 1-4) whose results were brought together in a General Discussion. Alpha diversity (species richness) increased by almost four species on average, whereas β diversity tended to decrease (Chapter 1). The latter is interpreted as a beginning floristic homogenization. The observed changes were primarily the result of a spread of native habitat generalists that are able to tolerate broad pH and moisture ranges. The changes in α and β diversity were only significant when species abundances were neglected (Chapters 1 and 2), demonstrating that the diversity changes resulted mainly from gains and losses of low-abundance species. This study is one of the first studies in temperate Europe that demonstrates floristic homogenization of forest plant communities at a larger than local scale. The diversity changes found at the taxonomic level did not result in similar changes at the functional level (Chapter 2). The likely reason is that these communities are functionally "buffered". Single communities involve most of the functional diversity of the regional pool, i.e., they are already functionally rich, while they are functionally redundant among each other, i.e., they are already homogeneous. Independent of taxonomic homogenization, the abundance of 30 species decreased significantly (Chapter 4). These species included 12 ancient forest species (i.e., species closely tied to forest patches with a habitat continuity > 200 years) and seven species listed on the Red List of endangered plant species in NW Germany. If these decreases continue over the next decades, local extinctions may result. This biotic impoverishment would seriously conflict with regional conservation goals. Community assembly mechanisms changed at the local level particularly at sites that experienced disturbance by forest management activities between the sampling periods (Chapter 3). Disturbance altered community assembly mechanisms in two ways: (i) it relaxed environmental filters and allowed the coexistence of different reproduction strategies, as reflected by a higher diversity of reproductive traits at the time of the resurvey, and (ii) it enhanced light availability and tightened competitive filters. These limited the functional diversity with respect to canopy height and selected for taller species. Thirty-one winner and 30 loser species, which had significantly increased or decreased in abundance, respectively, were characterized by various functional traits and ecological performances to find indications on the most likely environmental drivers for the observed floristic changes (Chapter 4). Winner species had higher seed longevity, flowered later in the season and had more often an oceanic distribution compared to loser species. Loser species tended to have a higher specific leaf area, to be more susceptible to deer browsing and to have a performance optimum at higher soil pH values compared to winner species. Multiple logistic regression analyses indicated that disturbances due to forest management interventions were the primary cause of the species shifts. As one of the first European resurvey studies, this study provides indications that an enhanced browsing pressure due to increased deer densities and increasingly warmer winters are important drivers. The study failed to demonstrate that eutrophication and acidification due to atmospheric deposition substantially drive herb layer changes. The restriction of the sample to the most base-rich sites in the region is discussed as a likely reason. Furthermore, the decline of several ancient forest species is discussed as an indication that the forest patches are still paying off their "extinction debt", i.e., exhibit a delayed response to forest fragmentation.},
  language  = {en}
}
@article{KolkNaafWulf2017,
  author    = {Kolk, Jens and Naaf, Tobias and Wulf, Monika},
  title     = {Paying the colonization credit},
  series = {Biodiversity and conservation},
  volume    = {26},
  journal   = {Biodiversity and conservation},
  publisher = {Springer},
  address   = {Dordrecht},
  issn      = {0960-3115},
  doi       = {10.1007/s10531-016-1271-y},
  pages     = {735 -- 755},
  year      = {2017},
  abstract  = {Massive historical land cover changes in the Central European lowlands have resulted in a forest distribution that now comprises small remnants of ancient forests and more recently established post-agricultural forests. Here, land-use history is considered a key driver of recent herb-layer community changes, where an extinction debt in ancient forest remnants and/or a colonization credit in post-agricultural forests are being paid over time. On a regional scale, these payments should in theory lead toward a convergence in species richness between ancient and post-agricultural forests over time. In this study, we tested this assumption with a resurvey of 117 semi-permanent plots in the well-studied deciduous forests of the Prignitz region (Brandenburg, NE Germany), where we knew that the plant communities of post-agricultural stands exhibit a colonization credit while the extinction debt in ancient stands has largely been paid. We compared changes in the species richness of all herb layer species, forest specialists and ancient forest indicator species between ancient and post-agricultural stands with linear mixed effect models and determined the influence of patch connectivity on the magnitude of species richness changes. Species richness increased overall, but the richness of forest specialists increased significantly more in post-agricultural stands and was positively influenced by higher patch connectivity, indicating a convergence in species richness between the ancient and postagricultural stands. Furthermore, the richness of ancient forest indicator species only increased significantly in post-agricultural stands. For the first time, we were able to verify a gradual payment of the colonization credit in post-agricultural forest stands using a comparison of actual changes in temporal species richness.},
  language  = {en}
}