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1.	Acharya, Kamal Prasad, et al. (author) Latitudinal variation of life-history traits of an exotic and a native impatiens species in Europe 2017 In: Acta Oecologica. - : Elsevier BV. - 1146-609X .- 1873-6238. ; 81, s. 40-47 Journal article (peer-reviewed)abstract Understanding the responses of invasive and native populations to environmental change is crucial for reliable predictions of invasions in the face of global change. While comparisons of responses across invasive species with different life histories have been performed before, comparing functional traits of congeneric native and invasive species may help to reveal driving factors associated with invasion. Here we compared morphological functional trait patterns of an invasive species (Impatiens parviflora) with its congeneric native species (I. noli-tangere) along an approximately 1600 km European latitudinal gradient from France (49 degrees 34'N) to Norway (63 degrees 40'N). Soil nitrogen was recorded during six weeks of the growing season, and light, soil moisture, and nutrient availability were estimated for each sampled population using community weighted means of indicator values for co-occurring species. Temperature data were gathered from nearby weather stations. Both the native and invasive species are taller at higher latitudes and this response is strongest in the invasive species. Seed mass and number of seeds per capsule increase in I. noli-tangere but decrease in I. parviflora towards higher latitudes. Surprisingly, plant height in the invasive I. parviflora decreases with increasing soil nitrogen availability. The latitudinal pattern in seed mass is positively related to temperature in I. noli-tangere and negatively in I. parviflora. Leaf area of both species decreases with increasing Ellenberg indicator values for nitrogen and light but increases with increasing soil moisture. Soil nitrogen concentrations and Ellenberg indicator values for nitrogen have significant positive (I. nolitangere) and negative (I. parviflora) effects on the number of seeds per capsule. Our results show that the native I. noli-tangere has efficient reproduction at its range edge while the invasive I. parviflora shows a marked decrease in seed size and seed number per capsule. These patterns are unrelated to the growth and obtained size of the plants: even low soil nitrogen availability in the north seemed not to limit plant growth and size. Our results suggest that the invasive I. parviflora tends to become more invasive at lower latitudes by producing heavier seeds and more seeds per capsule.
2.	Ampoorter, Evy, et al. (author) Tree diversity is key for promoting the diversity and abundance of forest-associated taxa in Europe 2020 In: Oikos. - : Wiley. - 0030-1299 .- 1600-0706. ; 129:2, s. 133-146 Journal article (peer-reviewed)abstract Plant diversity is an important driver of diversity at other trophic levels, suggesting that cascading extinctions could reduce overall biodiversity. Most evidence for positive effects of plant diversity comes from grasslands. Despite the fact that forests are hotspots of biodiversity, the importance of tree diversity, in particular its relative importance compared to other management related factors, in affecting forest-associated taxa is not well known. To address this, we used data from 183 plots, located in different forest types, from Mediterranean to Boreal, and established along a climatic gradient across six European countries (FunDivEUROPE project). We tested the influence of tree diversity, tree functional composition (i.e. functional trait values), forest structure, climate and soil on the diversity and abundance/activity of nine taxa (bats, birds, spiders, microorganisms, earthworms, ungulates, foliar fungal pathogens, defoliating insects and understorey plants) and on their overall diversity and abundance/activity (multidiversity, multiabundance/activity). Tree diversity was a key driver of taxon-level and overall forest-associated biodiversity, along with tree functional composition, forest structure, climate and soil. Both tree species richness and functional diversity (variation in functional trait values) were important. The effects of tree diversity on the abundance/activity of forest-associated taxa were less consistent. Nonetheless, spiders, ungulates and foliar fungal pathogens were all more abundant/active in diverse forests. Tree functional composition and structure were also important drivers of abundance/activity: conifer stands had lower overall multidiversity (although the effect was driven by defoliating insects), while stands with potentially tall trees had lower overall multiabundance/activity. We found more synergies than tradeoffs between diversity and abundance/activity of different taxa, suggesting that forest management can promote high diversity across taxa. Our results clearly show the high value of mixed forest stands for multiple forest-associated taxa and indicate that multiple dimensions of tree diversity (taxonomic and functional) are important.
3.	Baeten, Lander, et al. (author) A model-based approach to studying changes in compositional heterogeneity 2014 In: Methods in Ecology and Evolution. - 2041-210X. ; 5:2, s. 156-164 Journal article (peer-reviewed)
4.	Baeten, Lander, et al. (author) Identifying the tree species compositions that maximize ecosystem functioning in European forests 2019 In: Journal of Applied Ecology. - : Wiley. - 0021-8901 .- 1365-2664. ; 56:3, s. 733-744 Journal article (peer-reviewed)abstract 1. Forest ecosystem functioning generally benefits from higher tree species richness, but variation within richness levels is typically large. This is mostly due to the contrasting performances of communities with different compositions. Evidence-based understanding of composition effects on forest productivity, as well as on multiple other functions will enable forest managers to focus on the selection of species that maximize functioning, rather than on diversity per se.2. We used a dataset of 30 ecosystem functions measured in stands with different species richness and composition in six European forest types. First, we quantified whether the compositions that maximize annual above-ground wood production (productivity) generally also fulfil the multiple other ecosystem functions (multifunctionality). Then, we quantified the species identity effects and strength of interspecific interactions to identify the "best" and "worst" species composition for multifunctionality. Finally, we evaluated the real-world frequency of occurrence of best and worst mixtures, using harmonized data from multiple national forest inventories.3. The most productive tree species combinations also tended to express relatively high multifunctionality, although we found a relatively wide range of compositions with high- or low-average multifunctionality for the same level of productivity. Monocultures were distributed among the highest as well as the lowest performing compositions. The variation in functioning between compositions was generally driven by differences in the performance of the component species and, to a lesser extent, by particular interspecific interactions. Finally, we found that the most frequent species compositions in inventory data were monospecific stands and that the most common compositions showed below-average multifunctionality and productivity.4. Synthesis and applications. Species identity and composition effects are essential to the development of high-performing production systems, for instance in forestry and agriculture. They therefore deserve great attention in the analysis and design of functional biodiversity studies if the aim is to inform ecosystem management. A management focus on tree productivity does not necessarily trade-off against other ecosystem functions; high productivity and multifunctionality can be combined with an informed selection of tree species and species combinations.
5.	Bernes, Claes, et al. (author) How are biodiversity and dispersal of species affected by the management of roadsides? A systematic map 2017 In: Environmental Evidence. - : Springer Science and Business Media LLC. - 2047-2382. ; 6:1 Journal article (peer-reviewed)abstract Background: In many parts of the world, roadsides are regularly managed for traffic-safety reasons. Hence, there are similarities between roadsides and certain other managed habitats, such as wooded pastures and mown or grazed grasslands. These habitats have declined rapidly in Europe during the last century. For many species historically associated with them, roadsides may function as new primary habitats or as dispersal corridors in fragmented landscapes. Current recommendations for roadside management to promote conservation values are largely based on studies of plants in semi-natural grasslands, although such areas often differ from roadsides in terms of environmental conditions and disturbance regimes. Moreover, roadsides provide habitat not only for plants but also for many insects. For these reasons, stakeholders in Sweden have emphasised the need for more targeted management recommendations, based on actual studies of roadside biodiversity. Methods: This systematic map provides an overview of the available evidence on how biodiversity is affected by various forms of roadside management, and how such management influences the dispersal of species along roads or roadsides. We searched for literature using 13 online publication databases, 4 search engines, 36 specialist websites and 5 literature reviews. Search terms were developed in English, Danish, Dutch, French, German, Norwegian, Spanish and Swedish. Identified articles were screened for relevance using criteria set out in a protocol. No geographical restrictions were applied, and all species and groups of organisms were considered. Descriptions of included studies are available in an Excel file, and also in an interactive GIS application that can be accessed at an external website. Results: Our searches identified more than 7000 articles. The 207 articles included after screening described 301 individual studies considered to be relevant. More than two-thirds of these studies were conducted in North America, with most of the rest performed in Europe. More than half of the studies were published in grey literature such as reports from agencies or consultants. The interventions most commonly studied were herbicide use, sowing and mowing, followed by soil amendments such as mulching and fertiliser additions. The outcomes most frequently reported were effects of interventions on the abundance or species richness of herbs/forbs, graminoids and woody plants. Effects on insects and birds were reported in 6 and 3% of the studies, respectively. Conclusions: This systematic map is based on a comprehensive and systematic screening of all available literature on the effects of roadside management on biodiversity and dispersal of species. As such it should be of value to a range of actors, including managers and policymakers. The map provides a key to finding concrete guidance for conservation- and restoration-oriented roadside management from published research. However, the map also highlights important knowledge gaps: little data was found for some geographical regions, research is heavily biased taxonomically towards plants, and no study was found on how species dispersal was affected by roadside management. The map could therefore be a source of inspiration for new research.
6.	Bernes, Claes, et al. (author) How are biodiversity and dispersal of species affected by the management of roadsides? A systematic map protocol 2016 In: Environmental Evidence. - : Springer Science and Business Media LLC. - 2047-2382. ; 5:4 Journal article (peer-reviewed)abstract Background: In many parts of the world, roadsides are regularly managed for traffic-safety reasons. Hence, there are similarities between roadsides and certain other managed habitats, such as wooded pastures and mown or grazed grasslands. In contrast to roadsides, the latter habitats have declined rapidly in Europe during the last century, and today only a fraction of their former extent remains. For many species historically associated with these habitats, roadsides may therefore function as new primary habitats or as dispersal corridors in fragmented landscapes. Current recommendations for roadside management to promote conservation values are largely based on studies of plants in semi-natural grasslands, although such areas often differ from roadsides in terms of environmental factors and impacts. Moreover, roadsides provide habitats not only for plants but also for many insects, especially if they are sandy and exposed to the sun. For these reasons, stakeholders in Sweden have emphasised the need for more targeted management recommendations, based on actual studies of roadside biodiversity. Methods: The proposed systematic map is intended to provide an overview of the available evidence on how biodiversity is affected by various forms of roadside management, and how such management influences the dispersal of species along roads or roadsides. Relevant interventions include e.g. mowing, shrub removal, control of invasive/nuisance species, sowing or planting, burning, grazing by livestock, scraping and ditching. Non-intervention or alternative forms of roadside management will be used as comparators. Relevant outcomes include measures of species or genetic diversity, the abundance of individual species or groups of organisms, species distribution patterns, and movement rates of individuals or propagules. Searches will be made for peer-reviewed and grey literature in English and several other languages. No geographical restrictions will be applied, and all species and species groups will be considered.
7.	Bernes, Claes, et al. (author) How does roadside vegetation management affect the diversity of vascular plants and invertebrates? A systematic review protocol 2017 In: Environmental Evidence. - : Springer Science and Business Media LLC. - 2047-2382. ; 6:1 Research review (peer-reviewed)abstract Background: Roadsides have been acknowledged as potential substitutes for semi-natural grasslands and other open habitats with high biodiversity, many of which are now declining. Current recommendations for roadside management to promote conservation of biodiversity are largely based on studies of plants in meadows or pastures, although such areas often differ from roadsides in terms of environmental conditions and disturbance regimes. Stakeholders in Sweden have emphasised the need for more targeted guidelines for roadside management, based on actual roadside studies. We recently performed a systematic mapping of the evidence on how roadside management affects biodiversity and the dispersal of species. Through this process, we found 98 studies on how the richness or abundance of species on roadsides is affected by management such as regular mowing, burning, grazing or selective mechanical removal of plants. Since all of these interventions entail removal of plant biomass, they are comparable. Most of the studies recorded management effects on vascular plants, but there were 14 investigations of insects and other invertebrates. We now intend to proceed with a full systematic review of how maintenance or restoration of roadsides based on non-chemical vegetation removal affects the diversity of vascular plants and invertebrates. Methods: Most of the evidence on which the proposed systematic review is to be based will be selected from the systematic map. To identify more recently published literature on the topic of the review, we will perform a search update using a subset of the search terms applied for the systematic map. The criteria for inclusion of studies will be the same as for the map, except that interventions and outcomes will be restricted to those relevant to the review. Relevant studies will be subject to critical appraisal and categorised as having high or low validity for the review. Studies with low validity will be excluded from the review. Utilisable data on outcomes, interventions and other potential effect modifiers will be extracted from included articles. A narrative synthesis will describe the validity and findings of all studies in the review. Where a sufficient number of studies report similar outcome types, meta-analysis will be conducted.
8.	De Frenne, Pieter, et al. (author) An intraspecific application of the leaf-height-seed ecology strategy scheme to forest herbs along a latitudinal gradient 2011 In: Ecography. - : Wiley. - 0906-7590 .- 1600-0587. ; 34:1, s. 132-140 Journal article (peer-reviewed)abstract We measured LHS traits in 41 Anemone nemorosa and 44 Milium effusum populations along a 1900-2300 km latitudinal gradient from N France to N Sweden. We then applied multilevel models to identify the effects of regional (temperature, latitude) and local (soil fertility and acidity, overstorey canopy cover) environmental factors on LHS traits. Both species displayed a significant 4% increase in plant height with every degree northward shift (almost a two-fold plant height difference between the southernmost and northernmost populations). Neither seed mass nor SLA showed a significant latitudinal cline. Temperature had a large effect on the three LHS traits of Anemone. Latitude, canopy cover and soil nutrients were related to the SLA and plant height of Milium. None of the investigated variables appeared to be related to the seed mass of Milium. The variation in LHS traits indicates that the ecological strategy determined by the position of each population in this three-factor triangle is not constant along the latitudinal gradient. The significant increase in plant height suggests greater competitive abilities for both species in the northernmost populations. We also found that the studied environmental factors affected the LHS traits of the two species on various scales: spring-flowering Anemone was affected more by temperature, whereas early-summer flowering Milium was affected more by local and other latitude-related factors. Finally, previously reported cross-species correlations between LHS traits and latitude were generally unsupported by our within-species approach.
9.	De Frenne, Pieter, et al. (author) Biological Flora of the British Isles : Milium effusum 2017 In: Journal of Ecology. - : Wiley. - 0022-0477 .- 1365-2745. ; 105:3, s. 839-858 Journal article (peer-reviewed)abstract 1. This account presents information on all aspects of the biology of Milium effusum L. (Wood Millet) that are relevant to understanding its ecological characteristics and behaviour. The main topics are presented within the standard framework of the Biological Flora of the British Isles: distribution, habitat, communities, responses to biotic factors, responses to environment, structure and physiology, phenology, floral and seed characters, herbivores and disease, history, and conservation.2. The grass Milium effusum is a common species of mature woodland in central and southern England, but is less common in the wetter parts of northern England, Wales, Scotland and Ireland. Worldwide, the species is native to many temperate, boreal, subarctic and subalpine parts of the northern hemisphere: from eastern North America across most of Europe (excluding Mediterranean climates) to the Ural Mountains and Black Sea, extending eastwards to the Himalaya, Korea and Japan.3. Wood Millet is a shade-tolerant, relatively tall grass (up to 1.8 m) producing up to 700 caryopses per individual. It is characteristic of temperate deciduous woodland, but can also occur in other woodland and forest types and even in scrub, alpine meadows, along railways and roads, and on rocks. In woods, it is one of the most conspicuous plants of the herb layer in the early summer after the disappearance of spring flowering species. While the species is generally considered an ancient woodland indicator in England and western Europe, it is also known to colonize secondary, post-agricultural forests relatively rapidly in other areas such as Denmark, southern Sweden and Poland.4. The species has a wide amplitude in terms of soil acidity and nutrient availability, but predominantly grows on soils of intermediate soil fertility and soil pH and with high organic matter concentration. However, M. effusum can tolerate large quantities of tree-leaf litter on the forest floor and is able to grow on very acidic soils.5. Changes in land use, climate, densities of large herbivores and atmospheric deposition of nitrogen are having effects on populations of Wood Millet. Significant responses of the life-history traits and population characteristics have been detected in response to environmental variation and to experimental treatments of temperature, nutrients, light and acidity. In many of its habitats across its range, M. effusum is currently becoming more frequent. During the last century, its mean elevation of occurrence in upland areas of Europe has also increased by several hundreds of metres. Typically, management actions are directed towards the conservation of its main habitat type (e.g. ancient woodlands of the Milio-Fagetum association) rather than to the species specifically.
10.	De Frenne, Pieter, et al. (author) Global buffering of temperatures under forest canopies 2019 In: Nature Ecology & Evolution. - : Springer Science and Business Media LLC. - 2397-334X. ; 3:5, s. 744-749 Journal article (peer-reviewed)abstract Macroclimate warming is often assumed to occur within forests despite the potential for tree cover to modify microclimates. Here, using paired measurements, we compared the temperatures under the canopy versus in the open at 98 sites across 5 continents. We show that forests function as a thermal insulator, cooling the understory when ambient temperatures are hot and warming the understory when ambient temperatures are cold. The understory versus open temperature offset is magnified as temperatures become more extreme and is of greater magnitude than the warming of land temperatures over the past century. Tree canopies may thus reduce the severity of warming impacts on forest biodiversity and functioning.
11.	De Frenne, Pieter, et al. (author) Interregional variation in the floristic recovery of post-agricultural forests 2011 In: Journal of Ecology. - : Wiley. - 0022-0477 .- 1365-2745. ; 99:2, s. 600-609 Journal article (peer-reviewed)abstract 1. Worldwide, the floristic composition of temperate forests bears the imprint of past land use for decades to centuries as forests regrow on agricultural land. Many species, however, display significant interregional variation in their ability to (re)colonize post-agricultural forests. This variation in colonization across regions and the underlying factors remain largely unexplored. 2. We compiled data on 90 species and 812 species x study combinations from 18 studies across Europe that determined species' distribution patterns in ancient (i.e. continuously forested since the first available land use maps) and post-agricultural forests. The recovery rate (RR) of species in each landscape was quantified as the log-response ratio of the percentage occurrence in post-agricultural over ancient forest and related to the species-specific life-history traits and local (soil characteristics and light availability) and regional factors (landscape properties as habitat availability, time available for colonization, and climate). 3. For the herb species, we demonstrate a strong (interactive) effect of species' life-history traits and forest habitat availability on the RR of post-agricultural forest. In graminoids, however, none of the investigated variables were significantly related to the RR. 4. The better colonizing species that mainly belonged to the short-lived herbs group showed the largest interregional variability. Their recovery significantly increased with the amount of forest habitat within the landscape, whereas, surprisingly, the time available for colonization, climate, soil characteristics and light availability had no effect. 5. Synthesis. By analysing 18 independent studies across Europe, we clearly showed for the first time on a continental scale that the recovery of short-lived forest herbs increased with the forest habitat availability in the landscape. Small perennial forest herbs, however, were generally unsuccessful in colonizing post-agricultural forest even in relatively densely forested landscapes. Hence, our results stress the need to avoid ancient forest clearance to preserve the typical woodland flora.
12.	De Frenne, Pieter, et al. (author) Latitudinal gradients as natural laboratories to infer species' responses to temperature 2013 In: Journal of Ecology. - : Wiley. - 0022-0477 .- 1365-2745. ; 101:3, s. 784-795 Research review (peer-reviewed)abstract Macroclimatic variation along latitudinal gradients provides an excellent natural laboratory to investigate the role of temperature and the potential impacts of climate warming on terrestrial organisms. Here, we review the use of latitudinal gradients for ecological climate change research, in comparison with altitudinal gradients and experimental warming, and illustrate their use and caveats with a meta-analysis of latitudinal intraspecific variation in important life-history traits of vascular plants. We first provide an overview of latitudinal patterns in temperature and other abiotic and biotic environmental variables in terrestrial ecosystems. We then assess the latitudinal intraspecific variation present in five key life-history traits [plant height, specific leaf area (SLA), foliar nitrogen:phosphorus (N:P) stoichiometry, seed mass and root:shoot (R:S) ratio] in natural populations or common garden experiments across a total of 98 plant species. Intraspecific leaf N:P ratio and seed mass significantly decreased with latitude in natural populations. Conversely, the plant height decreased and SLA increased significantly with latitude of population origin in common garden experiments. However, less than a third of the investigated latitudinal transect studies also formally disentangled the effects of temperature from other environmental drivers which potentially hampers the translation from latitudinal effects into a temperature signal. Synthesis. Latitudinal gradients provide a methodological set-up to overcome the drawbacks of other observational and experimental warming methods. Our synthesis indicates that many life-history traits of plants vary with latitude but the translation of latitudinal clines into responses to temperature is a crucial step. Therefore, especially adaptive differentiation of populations and confounding environmental factors other than temperature need to be considered. More generally, integrated approaches of observational studies along temperature gradients, experimental methods and common garden experiments increasingly emerge as the way forward to further our understanding of species and community responses to climate warming.
13.	De Frenne, Pieter, et al. (author) Microclimate moderates plant responses to macroclimate warming 2013 In: Proceedings of the National Academy of Sciences of the United States of America. - : Proceedings of the National Academy of Sciences. - 0027-8424 .- 1091-6490. ; 110:46, s. 18561-18565 Journal article (peer-reviewed)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.
14.	De Frenne, Pieter, et al. (author) Plant movements and climate warming : intraspecific variation in growth responses to nonlocal soils 2014 In: New Phytologist. - : Wiley. - 0028-646X .- 1469-8137. ; 202:2, s. 431-441 Journal article (peer-reviewed)abstract Most range shift predictions focus on the dispersal phase of the colonization process. Because moving populations experience increasingly dissimilar nonclimatic environmental conditions as they track climate warming, it is also critical to test how individuals originating from contrasting thermal environments can establish in nonlocal sites. We assess the intraspecific variation in growth responses to nonlocal soils by planting a widespread grass of deciduous forests (Milium effusum) into an experimental common garden using combinations of seeds and soil sampled in 22 sites across its distributional range, and reflecting movement scenarios of up to 1600km. Furthermore, to determine temperature and forest-structural effects, the plants and soils were experimentally warmed and shaded. We found significantly positive effects of the difference between the temperature of the sites of seed and soil collection on growth and seedling emergence rates. Migrant plants might thus encounter increasingly favourable soil conditions while tracking the isotherms towards currently colder' soils. These effects persisted under experimental warming. Rising temperatures and light availability generally enhanced plant performance. Our results suggest that abiotic and biotic soil characteristics can shape climate change-driven plant movements by affecting growth of nonlocal migrants, a mechanism which should be integrated into predictions of future range shifts.
15.	De Frenne, Pieter, et al. (author) Temperature effects on forest herbs assessed by warming and transplant experiments along a latitudinal gradient 2011 In: Global Change Biology. - : Wiley. - 1354-1013 .- 1365-2486. ; 17:10, s. 3240-3253 Journal article (peer-reviewed)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.
16.	De Frenne, Pieter, et al. (author) The response of forest plant regeneration to temperature variation along a latitudinal gradient 2012 In: Annals of Botany. - : Oxford University Press (OUP). - 0305-7364 .- 1095-8290. ; 109:5, s. 1037-1046 Journal article (peer-reviewed)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.
17.	De Frenne, Pieter, et al. (author) The use of open-top chambers in forests for evaluating warming effects on herbaceous understorey plants 2010 In: Ecological research. - : Wiley. - 0912-3814 .- 1440-1703. ; 25:1, s. 163-171 Journal article (peer-reviewed)abstract Open-top chambers (OTCs) are widely used experimental warming devices in open-field ecosystems such as tundra and alpine heath. However, knowledge of their performance in temperate deciduous forest ecosystems is largely lacking. The application of OTCs in forests might become important in the future since the effects of climate warming on growth, reproduction, and future distribution of understorey forest herbs have rarely been investigated. Therefore, polycarbonate OTCs covered with (OTCs+GF) and without permeable polypropylene GardenFleece (OTCs−GF) were installed in a temperate deciduous forest to create an experimental warming gradient. Short-term responses in phenology, growth, and reproduction of a model understorey forest herb (Anemone nemorosa L.) to OTC installation were determined. In a second growing season, an in-depth study of multiple abiotic conditions inside OTCs−GF was performed. Both OTCs+GF and OTCs−GF raised air and soil temperature in a realistic manner (ca. +0.4°C to +1.15°C), but OTCs−GF only in the leafless period (up to +1.5°C monthly average soil temperature). The early flowering forest herb A. nemorosa also showed a clear phenotypic response to OTC installation. Based on these facts and the large ecological drawbacks associated with OTCs+GF (mostly in connection with a higher relative air humidity and a lower light quantity) and very modest abiotic changes in OTCs−GF, we encourage the use of OTCs−GF in deciduous forest ecosystems for evaluating climate-warming effects on early flowering understorey forest herbs. There is also a potential to use this warming method on later flowering species, but this needs further research.
18.	De Pauw, Karen, et al. (author) Forest understorey communities respond strongly to light in interaction with forest structure, but not to microclimate warming 2022 In: New Phytologist. - : Wiley. - 0028-646X .- 1469-8137. ; 233:1, s. 219-235 Journal article (peer-reviewed)abstract Forests harbour large spatiotemporal heterogeneity in canopy structure. This variation drives the microclimate and light availability at the forest floor. So far, we do not know how light availability and sub-canopy temperature interactively mediate the impact of macroclimate warming on understorey communities.We therefore assessed the functional response of understorey plant communities to warming and light addition in a full factorial experiment installed in temperate deciduous forests across Europe along natural microclimate, light and macroclimate gradients. Furthermore, we related these functional responses to the species’ life-history syndromes and thermal niches.We found no significant community responses to the warming treatment. The light treatment, however, had a stronger impact on communities, mainly due to responses by fast-colonizing generalists and not by slow-colonizing forest specialists. The forest structure strongly mediated the response to light addition and also had a clear impact on functional traits and total plant cover.The effects of short-term experimental warming were small and suggest a time-lag in the response of understorey species to climate change. Canopy disturbance, for instance due to drought, pests or logging, has a strong and immediate impact and particularly favours generalists in the understorey in structurally complex forests.
19.	De Pauw, Karen, et al. (author) Nutrient-demanding and thermophilous plants dominate urban forest-edge vegetation across temperate Europe 2024 In: Journal of Vegetation Science. - 1100-9233 .- 1654-1103. ; 35:1 Journal article (peer-reviewed)abstract QuestionsForests are highly fragmented across the globe. For urban forests in particular, fragmentation increases the exposure to local warming caused by the urban heat island (UHI) effect. We here aim to quantify edge effects on herbaceous understorey vegetation in urban forests, and test whether these effects interact with forest structural complexity.LocationWe set up a pan-European study at the continental scale including six urban forests in Zurich, Paris, Katowice, Brussels, Bremen, and Stockholm.MethodsWe recorded understorey plant communities from the edge towards the interior of urban forests. Within each urban forest, we studied edge-to-interior gradients in paired stands with differing forest structural complexity. Community composition was analysed based on species specialism, life form, light, nutrient, acidity and disturbance indicator values and species' thermal niches.ResultsWe found that herbaceous communities at urban forest edges supported more generalists and forbs but fewer ferns than in forests' interiors. A buffered summer microclimate proved crucial for the presence of fern species. The edge communities contained more thermophilous, disturbance-tolerant, nutrient-demanding and basiphilous plant species, a pattern strongly confirmed by corresponding edge-to-interior gradients in microclimate, soil and light conditions in the understorey. Additionally, plots with a lower canopy cover and higher light availability supported higher numbers of both generalists and forest specialists. Even though no significant interactions were found between the edge distance and forest structural complexity, opposing additive effects indicated that a dense canopy can be used to buffer negative edge effects.ConclusionThe urban environment poses a multifaceted filter on understorey plant communities which contributes to significant differences in community composition between urban forest edges and interiors. For urban biodiversity conservation and the buffering of edge effects, it will be key to maintain dense canopies near urban forest edges. The urban environment poses a multifaceted filter on understorey plant communities which contributes to significant differences in community composition between urban forest edges and interiors. For urban biodiversity conservation and the buffering of edge effects, it will be key to maintain dense canopies near urban forest edges.image
20.	De Pauw, Karen, et al. (author) Taxonomic, phylogenetic and functional diversity of understorey plants respond differently to environmental conditions in European forest edges 2021 In: Journal of Ecology. - : Wiley. - 0022-0477 .- 1365-2745. ; 109:7, s. 2629-2648 Journal article (peer-reviewed)abstract Forest biodiversity world-wide is affected by climate change, habitat loss and fragmentation, and today 20% of the forest area is located within 100 m of a forest edge. Still, forest edges harbour a substantial amount of terrestrial biodiversity, especially in the understorey. The functional and phylogenetic diversity of forest edges have never been studied simultaneously at a continental scale, in spite of their importance for the forests' functioning and for communities' resilience to future change.We assessed nine metrics of taxonomic, phylogenetic and functional diversity of understorey plant communities in 225 plots spread along edge-to-interior gradients in deciduous forests across Europe. We then derived the relative effects and importance of edaphic, stand and landscape conditions on the diversity metrics.Here, we show that taxonomic, phylogenetic and functional diversity metrics respond differently to environmental conditions. We report an increase in functional diversity in plots with stronger microclimatic buffering, in spite of their lower taxonomic species richness. Additionally, we found increased taxonomic species richness at the forest edge, but in forests with intermediate and high openness, these communities had decreased phylogenetic diversity.Functional and phylogenetic diversity revealed complementary and important insights in community assembly mechanisms. Several environmental filters were identified as potential drivers of the patterns, such as a colder macroclimate and less buffered microclimate for functional diversity. For phylogenetic diversity, edaphic conditions were more important. Interestingly, plots with lower soil pH had decreased taxonomic species richness, but led to increased phylogenetic diversity, challenging the phylogenetic niche conservatism concept.Synthesis. Taxonomic, phylogenetic and functional diversity of understorey communities in forest edges respond differently to environmental conditions, providing insight into different community assembly mechanisms and their interactions. Therefore, it is important to look beyond species richness with phylogenetic and functional diversity approaches when focusing on forest understorey biodiversity.
21.	De Pauw, Karen, et al. (author) The urban heat island accelerates litter decomposition through microclimatic warming in temperate urban forests 2024 In: Urban Ecosystems. - 1083-8155 .- 1573-1642. Journal article (peer-reviewed)abstract Forests worldwide are experiencing fragmentation, with especially important consequences for ecosystems bordering urbanized areas. Urban forests are exposed to local warming due to the urban heat island which affects their biodiversity and ecosystem functioning. A key ecosystem function affecting carbon and nutrient cycling in forests is litter decomposition, a process driven by the local microclimate. Thus, our aim was to clarify the impact of the urban heat island on litter decomposition in urban forests. We studied soil microclimate and litter decomposition in six urban forests across Europe and along local gradients from the urban forest edge to the interior. To quantify decomposition independent from local forest composition and litter quality, we used standardized green tea and rooibos tea litterbags. We determined the role of the soil microclimate and other environmental drivers for litter decomposition. Secondly, we assessed effects of edge proximity and landscape context on the soil microclimate. Soil characteristics were only driving green tea and not rooibos tea decomposition. On the contrary, higher soil temperatures resulted in faster rates of litter decomposition for both green and rooibos tea and were related to the proximity to the forest edge and the proportion of built-up area in the landscape. Via structural equation modelling we detected cascading effects of the urban heat island on litter decomposition. Such changes in litter decomposition have the potential to alter the soil food web, nutrient cycling and carbon drawdown in urban forests, and could result in significant interactions between urbanisation and ongoing climate change.
22.	De Pauw, Karen, et al. (author) Urban forest microclimates across temperate Europe are shaped by deep edge effects and forest structure 2023 In: Agricultural and Forest Meteorology. - 0168-1923 .- 1873-2240. ; 341 Journal article (peer-reviewed)abstract The urban heat island (UHI) causes strong warming of cities and their urban forests worldwide. Especially urban forest edges are strongly exposed to the UHI effect, which could impact urban forest biodiversity and functioning. However, it is not known to what extent the UHI effect alters edge-to-interior microclimatic gradients within urban forests and whether this depends on the forests' structure.Here we quantified gradients of air temperature, relative air humidity and vapour pressure deficits (VPD) along urban forest edge-to-interior transects with contrasting stand structures in six major cities across Europe. We performed continuous hourly microclimate measurements for two consecutive years and analysed the magnitude and depth of edge effects, as well as forest structural drivers of microclimatic variation.Compared to edge studies in rural temperate forests, we found that edge effects reached deeper into urban forests, at least up to 50 m. Throughout the year, urban forest edges were warmer and drier compared to forest interiors, with the largest differences occurring during summer and daytime. Not only maximum, but also mean and minimum temperatures were higher at the urban forest edge up to large edge distances (at least 85 m). Denser forests with a higher plant area index buffered high air temperatures and VPDs from spring to autumn.We conclude that urban forest edges are unique ecotones with specific microclimates shaped by the UHI effect. Both forest edges and interiors showed increased buffering capacities with higher forest canopy density. We advocate for the conservation and expansion of urban forests which can buffer increasingly frequent and intense climate extremes. To this end, urban forest managers are encouraged to aim for multi-layered dense forest canopies and consider edge buffer zones of at least 50 m wide.
23.	De Schuyter, Wim, et al. (author) Declining potential nectar production of the herb layer in temperate forests under global change 2024 In: Journal of Ecology. - 0022-0477 .- 1365-2745. ; 112:4, s. 832-847 Journal article (peer-reviewed)abstract Wild pollinators are crucial for ecosystem functioning and human food production and often rely on floral resources provided by different (semi-) natural ecosystems for survival. Yet, the role of European forests, and especially the European forest herb layer, as a potential provider of floral resources for pollinators has scarcely been quantified.In this study, we measured the potential nectar production (PNP) of the forest herb layer using resurvey data across 3326 plots in temperate forests in Europe, with an average time interval of 41 years between both surveys in order to assess (i) the importance of the forest herb layer in providing nectar for wild pollinators, (ii) the intra-annual variation of PNP, (iii) the overall change in PNP between survey periods and (iv) the change in intra-annual variation of PNP between survey periods. The PNP estimates nectar availability based on the relative cover of different plant species in the forest herb layer. Although PNP overestimates actual nectar production, relative differences amongst plots provide a valid and informative way to analyse differences across time and space.Our results show that the forest herb layer has a large potential for providing nectar for wild pollinator communities, which is greatest in spring, with an average PNP of almost 16 g sugar/m2/year. However, this potential has drastically declined (mean plot-level decline >24%).Change in light availability, associated with shifts in canopy structure and canopy composition, is the key driver of temporal PNP changes.Synthesis. Our study shows that if management activities are carefully planned to sustain nectar-producing plant species for wild pollinators, European forest herb layers and European forests as a whole can play key roles in sustaining wild pollinator populations.
24.	De Smedt, Pallieter, et al. (author) Desiccation resistance determines distribution of woodlice along forest edge-to-interior gradients 2018 In: European journal of soil biology. - : Elsevier BV. - 1164-5563 .- 1778-3615. ; 85, s. 1-3 Journal article (peer-reviewed)abstract Forest edges show strong abiotic and biotic gradients potentially altering community composition and ecosystem processes such as nutrient cycling. While abiotic gradients are well studied, short-scale biotic gradients, like detritivore species composition and their associated trait distribution remains a poorly explored research-field. We sampled woodlice in 160 forest patches across Europe at varying distances from the forest edge and discovered that species desiccation resistance determines distribution along forest edge-to-interior gradients. Forest edges are warmer and dryer compared to interiors and favour drought-tolerant species, while abundance and activity of drought-sensitive species is reduced at the edge. Key ecological factors for litter-dwelling detritivores (i.e. humidity) act as environmental filter, because of species-specific differences in desiccation resistance. Future research should focus on quantifying the consequences of a changing detritivore community and their associated functional traits for nutrient cycling.
25.	De Smedt, Pallieter, et al. (author) Forest edges reduce slug (but not snail) activity-density across Western Europe 2019 In: Pedobiologia. - : Elsevier BV. - 0031-4056 .- 1873-1511. ; 75, s. 34-37 Journal article (peer-reviewed)abstract Fragmentation strongly shapes the distribution of organisms within forest patches through contrasting environmental conditions between the edge and interior habitat. Edge-to-interior distribution patterns are, however, poorly studied for litter- and soil-dwelling fauna, such as terrestrial gastropods, despite their high densities and significant impact on ecosystem processes, as both herbivores and detritivores. Therefore, we investigated edge-to-interior abundance patterns of terrestrial gastropods in 224 fragmented forest patches across Western Europe. Catching over 15,000 gastropods, we found that slug abundance is reduced in forest edges, while snail abundance shows no response on the edge effect. We hypothesize that these patterns could be explained by higher drought tolerance of snails, since forest edges have reduced air and soil humidity and elevated temperatures compared to forest interiors. Reduced slug abundance in forest edges potentially has ecological consequences for herbivory in and outside forest patches and nutrient cycling.

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