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Via sheltering, decoupling and buffering mechanisms, tree canopies have the capacity to mitigate impacts of multiple global‐change drivers on below‐canopy processes and organisms in forests. As a result, canopies have an important potential as nature‐based solution. The optimal combinations of forest canopy structural attributes to jointly mitigate the impacts of multiple global‐change drivers on below‐canopy organisms and processes have received little attention to date. To help solving this research gap, here we review how forest canopies modulate the effects of four important global‐change drivers—climate warming, drought, air pollution and biological invasions—on below‐canopy conditions. Particular attention is paid to mitigating canopy attributes that can be influenced by forest management, including canopy cover, tree species composition and vertical and horizontal structure. Synthesis. We show that the potential of forest canopies to mitigate global‐change effects is highly context‐dependent and that optimal canopy‐based solutions strongly depend on the environmental context and the targeted subcanopy organisms. Hence, holistic approaches, which maximize synergies and minimize trade‐offs, are needed to optimize the solution potential of forest canopies. [ABSTRACT FROM AUTHOR]

Summary: Species are altering their phenology to track warming temperatures. In forests, understorey plants experience tree canopy shading resulting in light and temperature conditions, which strongly deviate from open habitats. Yet, little is known about understorey phenology responses to forest microclimates.We recorded flowering onset, peak, end and duration of 10 temperate forest understorey plant species in two mesocosm experiments to understand how phenology is affected by sub‐canopy warming and how this response is modulated by illumination, which is related to canopy change. Furthermore, we investigated whether phenological sensitivities can be explained by species' characteristics, such as thermal niche.We found a mean advance of flowering onset of 7.1 d per 1°C warming, more than previously reported in studies not accounting for microclimatic buffering. Warm‐adapted species exhibited greater advances. Temperature sensitivity did not differ between early‐ and later‐flowering species. Experimental illumination did not significantly affect species' phenological temperature sensitivities, but slightly delayed flowering phenology independent from warming.Our study suggests that integrating sub‐canopy temperature and light availability will help us better understand future understorey phenology responses. Climate warming together with intensifying canopy disturbances will continue to drive phenological shifts and potentially disrupt understorey communities, thereby affecting forest biodiversity and functioning. [ABSTRACT FROM AUTHOR]

Coffee is an important crop in the global south. However, ongoing changes in the climate system reinforce the need to quantify coffee plants' ecological and eco‐physiological traits to assure coffee production in the future. One way to assess how environmental changes affect coffee performance is via leaf traits, most notably leaf carbon and nitrogen concentrations (to reflect the nutrient status), leaf stable carbon isotope composition (δ13C) to determine intrinsic water use efficiency (WUEi), and specific leaf area (SLA) to describe carbon gain relative to water loss within a plant canopy as these traits are related to yields. Therefore, we sampled coffee plants growing at contrasting elevations using a space‐for‐time substitution approach for warming and superimposed a canopy cover gradient to assess whether increasing canopy cover could modulate responses to temperature. Three coffee shrubs were sampled in each of 59 coffee farms in southwest Ethiopia across elevations of 1500–2160 m a.s.l. and along canopy cover gradients from open to deep shade. Soil nutrient concentrations, light availability, soil temperature and moisture were quantified for each coffee shrub. Elevation and shade tree canopy cover significantly and interactively affected WUEi. Elevation was found to be the driving factor for microclimate and soil factors which indirectly influenced both SLA and WUEi. Both of these coffee leaf traits are moderately governed by soil temperature whereas leaf N and C:N are mainly controlled by soil temperature and soil chemical variables. As elevation increased, WUEi kept increasing at light (< 35%) to intermediate shade levels (35–65%), and the values decreased at dense shade levels (65–100%) at high elevations, suggesting that coffee plants growing at high elevations with light shade can assimilate more CO2 with minimum evaporative water loss. SLA declined with elevation. Leaf N and leaf C:N responded negatively and positively to shade canopy cover, respectively. In sum, elevation and canopy cover interactively determined microclimate and coffee leaf traits. Our findings are useful to adjust the intensity of shade, along with other tree‐level management tools, to modulate climate‐change effects on coffee at the farm level. [ABSTRACT FROM AUTHOR]

Trees have a strong influence on the chemical properties of the soil in which they grow. Establishing plantations with different tree species combinations thus potentially leads to divergence in soil chemistry. To study the degree to which differentiation already occurs during the first years after establishment, we made use of a biodiversity–ecosystem functioning experiment in Belgium, FORBIO. The multi-site experiment replicates tree species richness (1–4 species) and composition in three sites. The sites represent contrasting site contexts, mostly land-use history, soil, and climate. Soil samples (0–10 cm) were taken at the time of planting and approximately 8 years later. We measured the total C and N concentration, the Olsen P, the pH-H2O, and the concentration of base and Al cations. The change in chemical composition was strongly dependent on site conditions including former land use. Afforestation on former cropland had a positive impact on total C and Olsen P and a negative effect on base cations and pH. On sites reforested after clear-cut, soil texture and particular site preparation played an important role. On top of that, we found several significant effects of species composition. Strongest composition effects were detected in the reforested site on loamy soil with little soil disturbance. This study highlights that species choice can already affect soil chemistry in early stages of forest development, but that the nature of the effects may strongly depend on the context in which the plantations are established. Further research is needed to identify the most important contextual factors. [ABSTRACT FROM AUTHOR]

Questions: Human disturbance is increasingly affecting forest dynamics across the tropics. Forests can recover via natural secondary succession to pre‐disturbance levels of biodiversity, species composition, and ecosystem carbon stocks. Central Africa will be subject to increasingly high shifting‐cultivation pressure in the next decades, but succession trajectories of these ecosystem properties are still poorly known for the Congo basin. We addressed two questions: (1) how does taxonomic and functional composition and diversity shift during secondary succession; and (2) how fast do above‐ground carbon stocks recover during secondary succession in tropical forests? Location: Central Congo basin. Methods: We conducted an inventory of trees (diameter at breast height [DBH] ≥ 10 cm), measured species traits and soil texture and carbon content in 18 plots, located along six secondary succession stages (i.e., from agricultural to old growth forest sites). We measured tree diameter, height for 20% of trees distributed across diameter classes, wood traits from all species, and leaf traits from species that contributed to 85% of the plot basal area. Results: We showed that secondary forests recover relatively fast in terms of tree species diversity, alpha functional diversity, and fine‐root carbon, with near‐old‐growth forest values after six decades past disturbance, while floristic composition exhibited slower recovery. Secondary forests only partially shifted from acquisitive to a conservative life history, with shifts in leaf traits being largely decoupled from wood traits. Only 43% of above‐ground carbon recovered after 60 years of forest regrowth, potentially through a slow recovery of the large‐sized tree stems that dominate carbon stocks of old‐growth forests. Conclusions: Our findings underline the capacity of Afrotropical forests to recover species and alpha functional diversity after clear‐cutting through shifting cultivation. Simultaneously, old‐growth forests harbor a particular floristic community and store a large quantity of carbon with much longer recovery trajectories, stressing the need for conservation of these forests in the Congo basin. [ABSTRACT FROM AUTHOR]

Climatic drivers alone do not adequately explain the regional variation in budburst timing in deciduous forests across Europe. Stand‐level factors, such as tree species richness, might affect budburst timing by creating different microclimates under the same site macroclimate. We assessed different phases of the spring phenology (start, midpoint, end, and overall duration of the budburst period) of four important European tree species (Betula pendula, Fagus sylvatica, Quercus robur and Tilia cordata) in monocultures and four‐species mixture stands of a common garden tree biodiversity experiment in Belgium (FORBIO) in 2021 and 2022. Microclimatic differences between the stands in terms of bud chilling, temperature forcing, and soil temperature were considerable, with four‐species mixtures being generally colder than monocultures in spring, but not in winter. In the colder spring of 2021, at the stand level, the end of the budburst period was advanced, and its overall duration shortened, in the four‐species mixtures. At species level, this response was significant for F. sylvatica. In the warmer spring of 2022, advances in spring phenology in four‐species stands were observed again in F. sylvatica and, less markedly, in B. pendula but without a general response at the stand level. Q. robur showed specific patterns with delayed budburst start in 2021 in the four‐species mixtures and very short budburst duration for all stands in 2022. Phenological differences between monocultures and four‐species mixtures were linked to microclimatic differences in light availability rather than in temperature as even comparatively colder microclimates showed an advanced phenology. Compared to weather conditions, tree species richness had a lower impact on budburst timing, but this impact can be of importance for key species like F. sylvatica and colder springs. These results indicate that forest biodiversity can affect budburst phenology, with wider implications, especially for forest‐ and land surface models. [ABSTRACT FROM AUTHOR]

Aim: Revisits of non‐permanent, relocatable plots first surveyed several decades ago offer a direct way to observe vegetation change and form a unique and increasingly used source of information for global change research. Despite the important insights that can be obtained from resurveying these quasi‐permanent vegetation plots, their use is prone to both observer and relocation errors. Studying the combined effects of both error types is important since they will play out together in practice and it is yet unknown to what extent observed vegetation changes are influenced by these errors. Methods: We designed a study that mimicked all steps in a resurvey study and that allowed determination of the magnitude of observer errors only vs the joint observer and relocation errors. Communities of vascular plants growing in the understorey of temperate forests were selected as study system. Ten regions in Europe were covered to explore generality across contexts and 50 observers were involved, which deliberately differed in their experience in making vegetation records. Results: The mean geographic distance between plots in the observer+relocation error data set was 24 m. The mean relative difference in species richness in the observer error and the observer+relocation data set was 15% and 21%, respectively. The mean "pseudo‐turnover" between the five records at a quasi‐permanent plot location was on average 0.21 and 0.35 for the observer error and observer+relocation error data sets, respectively. More detailed analyses of the compositional variation showed that the nestedness and turnover components were of equal importance in the observer data set, whereas turnover was much more important than nestedness in the observer+relocation data set. Interestingly, the differences between the observer and the observer+relocation data sets largely disappeared when looking at temporal change: both the changes in species richness and species composition over time were very similar in these data sets. Conclusions: Our results demonstrate that observer and relocation errors are non‐negligible when resurveying quasi‐permanent plots. A careful interpretation of the results of resurvey studies is warranted, especially when changes are assessed based on a low number of plots. We conclude by listing measures that should be taken to maximally increase the precision and the strength of the inferences drawn from vegetation resurveys. Revisits to quasi‐permanent plots offer a direct way to observe vegetation change. Since their use is prone to both observer and relocation errors, we designed a study that allowed determination of the magnitude of these errors. Our results demonstrate that errors are non‐negligible and that a careful interpretation of the results of resurvey studies is warranted. [ABSTRACT FROM AUTHOR]

Abstract: Understanding the processes that underlie drought‐related tree vitality loss is essential for anticipating future forest dynamics, and for developing management plans aiming at increasing the resilience of forests to climate change. Forest vitality has been continuously monitored in Europe since the acid rain alert in the 1980s, and the intensive monitoring plots of ICP Forests offer the opportunity to investigate the effects of air pollution and climate change on forest condition. By making use of over 100 long‐term monitoring plots, where crown defoliation has been assessed extensively since 1990, we discovered a progressive shift from a negative to a positive effect of species richness on forest health. The observed tipping point in the balance of net interactions, from competition to facilitation, has never been reported from real ecosystems outside experimental conditions; and the strong temporal consistency of our observations with increasing drought stress emphasizes its climate change relevance. Furthermore, we show that higher species diversity has reduced the severity of defoliation in the long term. Our results confirm the greater resilience of diverse forests to future climate change‐induced stress. More generally, they add to an accumulating body of evidence on the large potential of tree species mixtures to face manifold disturbances in a changing world. [ABSTRACT FROM AUTHOR]

Shade trees are used in many coffee production systems across the globe. Beyond the benefits on biodiversity conservation, climate buffering, carbon sequestration and pathogen regulation, shade trees can impact the soil nutrient status via, for instance, litter inputs and nitrogen fixation. Since soil nutrients affect coffee quality and taste, there is also a potential indirect effect of shade tree species on coffee quality. Yet, in spite of the potentially large impact of shade tree species, quantitative data on the effects of shade trees on (i) soil biogeochemistry and (ii) the associated coffee bean quality remain scarce. To what extent four widely used shade trees species (Acacia abyssinica L. , Albizia gummifera L. , Cordia africana L. and Croton macrostachyus L.) in a plantation coffee agroforestry system impact soil biogeochemistry, and how this in turn affects coffee quality, measured as cupping scores and bean size. A significant negative impact of N-fixing shade tree species on soil pH and base cation concentrations was found. Plant-available and total phosphorus was enhanced by the presence of Albizia gummifera L. Thus, the present findings demonstrate that careful selection and integration of shade tree species such as Acacia abyssinica L. and Albizia gummifera L. into coffee production systems is a good practice for sustaining soil chemical properties in coffee agroecosystem. Despite the impacts on soil chemical characteristics, the shade tree species had no effect on coffee cup quality but did affect the bean mass. In this particular study, an attempt was made to quantify the impacts of widely used shade tree species on soil biogeochemistry and the subsequent effect on coffee bean quality in a plantation agroforestry system over the course of one season in southwest Ethiopia. However, it might be feasible to accommodate both relatively sparse time-series experimental data consisting of coffee farms from plantations and smallholders, which needs to be the goal of future research to accurately examine the impacts on the outcome variables. • N-fixing shade tree species impacted soil pH negatively (increase in soil acidity and decrease in base cation concentrations). • Plant-available and total phosphorus contents are enhanced by the presence of N-fixing shade tree species. • Careful selection and integration of shade tree species such as Acacia abyssinica and Albizia gummifera into coffee agroecosystem could be a good practice for sustaining soil biogeochemical parameters. • Shade tree species identity did not impact coffee beverages. [ABSTRACT FROM AUTHOR]

Abstract: Temperate forests across Europe and eastern North America have become denser since the 1950s due to less intensive forest management and global environmental changes such as nitrogen deposition and climate warming. Denser tree canopies result in lower light availability at the forest floor. This shade may buffer the effects of nitrogen deposition and climate warming on understorey plant communities. We conducted an innovative in situ field experiment to study the responses of co‐occurring soil microbial and understorey plant communities to nitrogen addition, enhanced light availability and experimental warming in a full‐factorial design. We determined the effects of multiple environmental drivers and their interactions on the soil microbial and understorey plant communities, and assessed to what extent the soil microbial and understorey plant communities covary. High light led to lower biomass of the soil microbes (analysed by phospholipid fatty acids), but the soil microbial structure, i.e. the ratio of fungal biomass to bacterial biomass, was not affected by light availability. The composition of the soil bacterial community (analysed by high‐throughput sequencing) was affected by both light availability and warming (and their interaction), but not by nitrogen addition. Yet, the number of unique operational taxonomic units was higher in plots with nitrogen addition, and there were significant interactive effects of light and nitrogen addition. Light availability also determined the composition of the plant community; no effects of nitrogen addition and warming were observed. The soil bacterial and plant communities were co‐structured, and light availability explained a large part of the variance of this co‐structure. We provide robust evidence for the key role of light in affecting both the soil microbial and plant communities in forest understoreys. Our results advocate for more multifactor global change experiments that investigate the mechanism underlying the (in) direct effects of light on the plant–soil continuum in forests. A plain language summary is available for this article. [ABSTRACT FROM AUTHOR]

Abstract: Overyielding in mixed‐species forests has been demonstrated in a vast body of literature, and the focus of functional biodiversity research is now shifting towards a mechanistic understanding of these observations. We explored diversity–productivity relationships at two sites of a large‐scale tree diversity experiment, with harsh (Ged) and benign (Zed) environmental conditions for plantation establishment. Additive partitioning methodologies were adopted to detect phenomenological patterns in the productivity data, and the trait structure of mixed communities was used to advance insights into compositional effects. After 6 years of plantation development, biomass productivity was significantly higher in mixtures compared to the monocultures of component species. We observed that processes operated through direct tree–tree interactions, as the diversity signal disappeared where trees in mixed stands were surrounded by conspecific neighbours only. This result is particularly relevant for mixed‐species plantation systems, as trees are commonly planted in monospecific patches to simplify management. Partitioning unveiled strong selection effects at both plantation sites. However, at the harsh Ged‐site this was caused by competitive dominance of species with fast young growth, whereas at the benign Zed‐site, species with slow young growth improved their performances but not at the expense of others (i.e. trait‐dependent complementarity). Species tolerance to shading is an influential trait for predicting biodiversity effects, with community‐weighted means in shade tolerance mediating dominance effects (Ged) and functional diversity in shade tolerance mediating (trait‐dependent) complementarity effects (Zed). Synthesis. This study highlights that biodiversity effects in young tree plantations could be explained by the functional composition of mixed communities, with a key role for species levels of shade tolerance. As contrasting results between plantation sites were observed, future research should target the context‐dependency of diversity–productivity relationships. [ABSTRACT FROM AUTHOR]

Small forest patches embedded in agricultural (and peri-urban) landscapes in Western Europe play a key role for biodiversity conservation with a recognized capacity of delivering a wide suite of ecosystem services. Measures aimed to preserve these patches should be both socially desirable and ecologically effective. This study presents a joint ecologic and economic assessment conducted on small forest patches in Flanders (Belgium) and Picardie (N France). In each study region, two contrasted types of agricultural landscapes were selected. Open field (OF) and Bocage (B) landscapes are distinguished by the intensity of their usage and higher connectivity in the B landscapes. The social demand for enhancing biodiversity and forest structure diversity as well as for increasing the forest area at the expenses of agricultural land is estimated through an economic valuation survey. These results are compared with the outcomes of an ecological survey where the influence of structural features of the forest patches on the associated herbaceous diversity is assessed. The ecological and economic surveys show contrasting results; increasing tree species richness is ecologically more important for herbaceous diversity in the patch, but both tree species richness and herbaceous diversity obtain insignificant willingness to pay estimates. Furthermore, although respondents prefer the proposed changes to take place in the region where they live, we find out that social preferences and ecological effectiveness do differ between landscapes that represent different intensities of land use. Dwellers where the landscape is perceived as more “degraded” attach more value to diversity enhancement, suggesting a prioritization of initiatives in these area. In contrast, the ecological analyses show that prioritizing the protection and enhancement of the relatively better-off areas is more ecologically effective. Our study calls for a balance between ecological effectiveness and welfare benefits, suggesting that cost effectiveness studies should consider these approaches jointly. [ABSTRACT FROM AUTHOR]

Due to forest fragmentation, forest edges have become dominant features in landscapes around the world. Forest edges are exposed to a different microclimate and to higher atmospheric nitrogen (N) deposition compared to the forest interior. It is still unclear how both factors affect N cycling at temperate forest edges. In this study, the microbial community structure was mapped using phospholipid fatty acids (PLFA) in forest edge (0–5 m) and interior (64 m) in two oak ( Quercus robur ) stands, two pine ( Pinus nigra ) stands and one spruce ( Picea sitchensis ) stand in northern Belgium and Denmark. Nitrogen mineralization, nitrification and immobilization rates were obtained via the in situ 15 N pool dilution technique in the forest edge and interior, and linked to the microbial community structure. Furthermore, we assessed 15 N recovery in simulated throughfall via the 15 N tracing method in edge and interior as a proxy for the long-term fate of mineral N. Microbial biomass was higher at the forest edges compared to the forest interiors and was associated to the higher gross mineralization rates. Gross nitrification rates were not increased at the edge, hereby preventing NO 3 − leaching losses. Gross and net nitrification rates differed between the forest types, where the oak stands were characterized by higher nitrification rates than the pine and spruce stands. The oak stand retained 15 NO 3 − in the mineral soil at the edge, while in the pine stand the polyphenol-rich litter layer retained more 15 NO 3 − in the forest interior. Overall, our results indicated that the specific characteristics of the forest edge (atmospheric deposition, microclimate, pH of mineral soil and C/N ratio of the forest floor) increased microbial biomass and gross mineralization rates. Given the omnipresence of forest edges, more research should be conducted to validate our observations for other forest and soil types. [ABSTRACT FROM AUTHOR]

Aims The environmental legacies of past land use generally have an important impact on the establishment and persistence of forest understorey plant species in post-agricultural forests. Here we studied how the nature of such environmental limitation changes during the first decades of forest development. Location Deciduous forest 'Muizenbos', northern Belgium. Methods Two forest herbs with comparable life-history traits, but contrasting rates of post-agricultural forest colonization ( Primula elatior, Geum urbanum) were introduced 18 yr ago. Introduction sites were ancient or post-agricultural forest; seeds or adults were added to permanent plots where the understorey vegetation was initially cleared or left undisturbed. We measured chemical soil conditions and recorded the understorey plant community to characterize the sites. The effects of the treatments (land use, vegetation clearing) were tested on the present-day occurrence and abundance of the introduced species across the plots. We also quantified whether the temporal changes in the total number of individuals per site was different between the two land-use types and consistent within land-use types. Results Land use had an important effect on soil and vegetation. In post-agricultural forest, the P availabilities were many times higher, the plant community diversity was lower and the competitive species were significantly more abundant compared with ancient forest. The present-day abundance of P. elatior was significantly higher in the ancient forest sites, both for planted and seeded plots, while for G. urbanum land use had no effect. The initial vegetation clearing had a persistent effect on the establishment success of both species. The 18-yr-long time series showed consistent changes in the total population sizes in ancient forest: P. elatior has been expanding and G. urbanum initially increased and then declined over the past decade. In post-agricultural forest, temporal patterns were more divergent and partly related to differences in dominant tree species: differences in acidification rates and changes in insolation caused by canopy dieback have put the populations on new and unforeseen trajectories. Conclusions The longer the experiment is running, the more the individual sites show separate dynamics. In general, we showed that environmental limitation affects plant colonization at least during the first two decades of secondary forest succession, but also that as the post-agricultural forests develop, the nature of environmental limitation changes. [ABSTRACT FROM AUTHOR]

More and more ecologists have started to resurvey communities sampled in earlier decades to determine long-term shifts in community composition and infer the likely drivers of the ecological changes observed. However, to assess the relative importance of and interactions among multiple drivers, joint analyses of resurvey data from many regions spanning large environmental gradients are needed. In this article, we illustrate how combining resurvey data from multiple regions can increase the likelihood of driver orthogonality within the design and show that repeatedly surveying across multiple regions provides higher representativeness and comprehensiveness, allowing us to answer more completely a broader range of questions. We provide general guidelines to aid the implementation of multiregion resurvey databases. In so doing, we aim to encourage resurvey database development across other community types and biomes to advance global environmental change research. [ABSTRACT FROM AUTHOR]

Biodiversity can insure ecosystems against declines in their functioning by increasing the mean level of ecosystem processes and decreasing the spatial or temporal variance of these processes. On this basis, mixing tree species is expected to be an effective management strategy to reduce the risk of planting failure in young plantations., We examined the effects of biodiversity insurance on sapling survival in three tree diversity experiments across Belgium. Based on the survival scoring of 89 254 saplings, planted in 126 plots with up to four-species mixtures, we tested two hypotheses: (i) variability in plot-level survival is lower for mixtures compared to monocultures due to compensation among the species (i.e. buffering effect) and (ii) mean survival is higher due to facilitation (i.e. performance-enhancing effect)., Variation in plot-level survival decreased strongly with diversity, indicating a buffering effect. The risk of severe planting failure was reduced in mixtures because species exhibit different survival rates; therefore, mixing ensures that not all trees in the plantation are equally susceptible to environmental stressors. In contrast, the mean plot-level survival did not increase with diversity, and thus, an overall performance-enhancing effect was lacking. However, species-level analyses did show performance-enhancing effects, where some species profited from mixing while others did not., Synthesis and applications. We conclude that biodiversity through mixing tree species insures young experimental plantations against planting failure and is therefore highly recommended as a planting management strategy. The risk of large mortality gaps is reduced if tree plantation saplings are mixed at the scale of individual trees or small cells of trees. [ABSTRACT FROM AUTHOR]

• We investigated effects of tree species diversity on rainfall partitioning. • Rainfall partitioning was highly dependant on tree species identity. • Non-additive diversity effects were detected in mixed plots. • Tree species diversity did not affect rainfall partitioning directly. • Species richness affected rainfall partitioning indirectly by increased basal area. Rainfall partitioning is a key process in forest ecosystems, affecting evaporation, transpiration and water storage in the canopy, litter and soil. Although rainfall partitioning has been well studied in forest ecosystems, less is known about how rainfall partitioning is modified by tree species diversity. To disentangle the effects of tree species identity and diversity on rainfall partitioning, we installed 128 rain gauges in an experimental plantation, located in Zedelgem (Belgium), part of a large, multi-site forest diversity experiment in Belgium (FORBIO). With these rain gauges, we monitored throughfall for a full year, with a temporal resolution of two weeks. Based on these data, we analysed identity (i.e. whether throughfall is species-specific), non-additive (i.e. whether throughfall in mixtures is different from the expected amounts based on monocultures) and richness effects (i.e. how species richness affects throughfall). First of all, we found that rainfall partitioning highly differed between tree species, due to pronounced species identity effects. The presence of Pinus sylvestris [pine] was always associated with lower throughfall. Year-round throughfall was, for example, the lowest in mixtures of pine and Tilia cordata [lime] (43.8 ± 14.5% of the open-field rainfall) and monocultures of pine (48.6 ± 11.8%). Additionally, non-additive effects were significant, leading to a 10% decrease of rainfall reaching the forest floor in mixtures compared to the expected throughfall based on measurements in monocultures. Tree species richness effects on rainfall partitioning acted indirectly, mostly via an increased basal area in mixed stands. Our findings suggest that mixing tree species leads to a higher than expected reduction in throughfall, which lowered soil moisture in mixed plots. Whether this reduced throughfall and soil moisture affects ecosystem functioning remains to be quantified. [ABSTRACT FROM AUTHOR]

The area of forest plantations is increasing worldwide helping to meet timber demand and protect natural forests. However, with global change, monospecific plantations are increasingly vulnerable to abiotic and biotic disturbances. As an adaption measure we need to move to plantations that are more diverse in genotypes, species, and structure, with a design underpinned by science. TreeDivNet, a global network of tree diversity experiments, responds to this need by assessing the advantages and disadvantages of mixed species plantations. The network currently consists of 18 experiments, distributed over 36 sites and five ecoregions. With plantations 1-15 years old, TreeDivNet can already provide relevant data for forest policy and management. In this paper, we highlight some early results on the carbon sequestration and pest resistance potential of more diverse plantations. Finally, suggestions are made for new, innovative experiments in understudied regions to complement the existing network. [ABSTRACT FROM AUTHOR]

Questions How do land-use legacies and distance to forest patches influence tree colonization at a post-agricultural heathland? Are colonizing species with different life-history traits affected differently by these factors? Is the effect of increased nutrient availability from land-use legacies mediated by the understorey vegetation? Location Nørholm hede, a 350-ha heathland in southwest Denmark developing naturally after abandonment of traditional agricultural practices in 1895. Methods Using 140 plots (0.03 ha), we quantified the vegetation structure, soil properties and natural recruitment of tree/shrubs in the heathland. Further, we used high-resolution Li DAR data to classify the vegetation and identify forest patches. In the analysis, we first used a logistic mixed model to test whether colonization of tree and shrub species differed between areas with different land-use history and whether it was influenced by the distance to forest patches and life-history traits (seed mass) of colonizing species. Then, to determine how different factors influence colonization, we explored the direct and indirect relationships among nutrient availability, density of adult trees, canopy cover, cover of Deschampsia flexuosa and the probability of seedling colonization in each plot using a confirmatory multilevel path analysis. Results The probability of seedling colonization for small- and large-seeded species decreased with distance to forest patches. This response was more pronounced at the previously cultivated area. Multilevel path analysis showed that colonizing species with different life-history traits were affected by different factors. Small-seeded species were negatively affected by increased nutrient availability mediated by an increase in the cover of D. flexuosa. This effect was not found for large-seeded species. Moreover, the density of trees/shrubs (>2 m) increased the probability of colonization of small-seeded species, whereas higher canopy cover had a positive effect on large-seeded species. Conclusions Our analyses demonstrate that the interactions between abiotic factors, biotic interactions and life-history traits of colonizing species can lead to distinct patterns of tree colonization. Land-use legacies can have long-lasting, indirect effects on tree colonization by altering the composition of the understorey vegetation, which in turn can delay tree encroachment and slow down the succession from heathland to forest. [ABSTRACT FROM AUTHOR]

We compared communities of vascular plants and arthropods in ten Short Rotation Coppice (SRC)–maize pairs, to (1) quantify the difference in diversity and composition between these two alternative land-use types and (2) to assess the potential of SRC plantations to increase functional biodiversity values in agricultural landscapes. In each SRC plantation and maize field, the vegetation was surveyed and arthropods were sampled by applying pitfall and pan trapping. The composition of the vegetation and of the epigeic and vegetation inhabiting arthropod communities strongly differed between the crop types. This differentiation was mainly due to true species turnover and only to a lesser extent to the occurrence of nested subsets. On average, the total cover of the vegetation was 10 times higher in the SRC plantations and taxonomic and trait diversity were also consistently higher in SRC. Arthropod activity densities were significantly higher, sometimes almost double, in SRC plantations. Significantly higher effective species numbers in SRC were only retrieved for Hymenoptera and Coleoptera. Regarding functional groups, the activity densities of omnivores, detritivores, mycophages, phytophages and parasitoids were significantly higher in SRC. While activity densities of predators were not different among the crop types, their effective species number was higher in SRC, indicating a more evenly distributed and diverse predator community. To conclude, we have shown that SRC can significantly increase vegetation and arthropod abundance and/or diversity in agricultural landscapes when replacing annual biomass crops, such as maize. [ABSTRACT FROM AUTHOR]

Abstract: One of the current advances in functional biodiversity research is the move away from short-lived test systems towards the exploration of diversity-ecosystem functioning relationships in structurally more complex ecosystems. In forests, assumptions about the functional significance of tree species diversity have only recently produced a new generation of research on ecosystem processes and services. Novel experimental designs have now replaced traditional forestry trials, but these comparatively young experimental plots suffer from specific difficulties that are mainly related to the tree size and longevity. Tree species diversity experiments therefore need to be complemented with comparative observational studies in existing forests. Here we present the design and implementation of a new network of forest plots along tree species diversity gradients in six major European forest types: the FunDivEUROPE Exploratory Platform. Based on a review of the deficiencies of existing observational approaches and of unresolved research questions and hypotheses, we discuss the fundamental criteria that shaped the design of our platform. Key features include the extent of the species diversity gradient with mixtures up to five species, strict avoidance of a dilution gradient, special attention to community evenness and minimal covariation with other environmental factors. The new European research platform permits the most comprehensive assessment of tree species diversity effects on forest ecosystem functioning to date since it offers a common set of research plots to groups of researchers from very different disciplines and uses the same methodological approach in contrasting forest types along an extensive environmental gradient. [Copyright &y& Elsevier]

Questions How do different substrate types and soil nutrient levels influence species composition and successional sequences, as represented by different life-history traits? Is the rate of succession increasing or decreasing along a gradient from low to high fertility? Location Ecological field station Dahlem, Berlin, Germany. Methods An 18-yr permanent plot study of succession on soils of differing fertility. The original substrates were a nutrient-poor sandy soil from a sandpit, ruderal subsoil of moderate fertility and nutrient-rich topsoil. Results In plots with sand, monocarpic perennials were dominant at the beginning, which were then replaced by perennial grasses, those being then replaced by woody perennials. On ruderal soil, monocarpic perennials were co-dominant with perennial herbs at first, then were replaced by perennial herbs, which were then also replaced by woody plants. Dominance of woody species was attained after about 10 yr, both on sand and on ruderal soil. In plots with topsoil, there was a short phase at the start where monocarpic perennials were co-dominant with perennial herbs, followed by a relatively long period of perennial herb dominance. There were about twice as many woody species growing on the substrates of low and intermediate fertility compared to the nutrient-rich soil. If we compare the time necessary for woody species to reach 10% and 50% cover, woody colonization was much slower on the resource-rich site. DCA ordination revealed a clear separation of samples along the nutrient gradient. It also showed slower community change with increasing soil fertility. Conclusions This study demonstrates that woodlands can be created on bare ground in a temperate climate through natural colonization in a relatively short time, if substrates are not too nutrient-rich. The assumption that the rate of succession increases with increasing productivity could not be confirmed. Succession towards woodland can be rapid on soils of low and intermediate fertility. In contrast, a nutrient-rich substrate favours perennial herbaceous vegetation, which inhibits woody colonization and arrests succession. [ABSTRACT FROM AUTHOR]

Summary 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. [ABSTRACT FROM AUTHOR]

Question: (i) How does former land use and land use intensity affect seed bank development during post-agricultural succession? (ii) How does time since the last clear-cut change seed bank composition during post-clear-cut succession? Methods: One data set was compiled per succession type using the following selection criteria: (i) the data set included a successional series, (ii) plots were located in mesotrophic forest plant communities and (iii) vegetation data were available. The post-agricultural succession data set comprised 76 recent forest plots (eight studies); the post-clear-cut succession data set comprised 218 ancient forest plots (three studies). Each data set was analysed separately using either linear mixed models or generalized linear models, controlling for both environmental heterogeneity and variation between study locations. Results: In the post-agricultural succession data set, land use and time significantly affected nearly all the studied seed bank characteristics. Seed banks on former arable land recovered poorly even after 150 year of restored forest cover, whereas moderate land use intensities (grasslands, heathlands) yielded more rapid seed bank recovery. Time was a significant determinant of all but two soil seed bank characteristics during post-clear-cut succession. Seed banks in managed ancient forest differed strongly in their characteristics compared to primary forest seed banks. Conclusions: Forest seed banks bear the marks of former land use and/or forest management and continue to do so for at least 150 years. Nevertheless, time since the last major disturbance, being either former land use or clear-cutting, remains a significant determinant of the seed bank. [ABSTRACT FROM AUTHOR]

Seed banks are often crucial for the survival of plant species, especially short-lived ones. Nevertheless, empirical studies about the fate of seeds under natural conditions are scarce, particularly for parasitic plants. Therefore, fresh seeds of the holoparasitic Cuscuta epithymum were buried in the field under natural conditions in Belgium or kept at room temperature for up to 31 months, and germination was tested periodically by incubating seeds in light at 238C. Buried C. epithymum seeds had cyclic changes in their germinability, while those stored dry at room temperature never germinated without scarification. Buried seeds exhibited a seasonal cycle of physiological dormancy, the first to be reported for a species with combinational (i.e. physical × physiological) dormancy. The physiological dormancy of the embryo was broken during winter (≤5°C) but induced during exposure to late spring temperatures (>10°C). Therefore, germination of C. epithymum seeds is fine tuned to seasonal temperature changes. Each year only a portion of the viable seeds could germinate. The portion of seeds buried in the soil that remained available for delayed germination, i.e. in the seed bank, decreased rather quickly, with only 8.5% of them being viable after 31 months of burial; all seeds stored in the laboratory remained viable. Nonetheless, we concluded that some C. epithymum seeds are capable of surviving for several years in the soil. These data indicate that the parasite is well adapted to survive in a dynamic heathland landscape where conditions for survival change constantly, but it is still sensitive to local extinction. [ABSTRACT FROM AUTHOR]

By creating transient patch mosaics, disturbance can influence the dynamics of interacting populations in many ecosystems. In European heathland, traditional land use created such dynamic systems favourable for both early and later successional species. Little empirical evidence is, however, available on the impact of current management on metapopulations occurring in such landscapes. This paper looks at the metapopulation viability of the endangered holoparasite Cuscuta epithymum, a species that typically occurs in early successional stages of recently managed heathlands. We used both observational and experimental data from a 4-yr study to parameterise a spatially explicit metapopulation model. This model explores the impact of demographic characteristics and spatiotemporal landscape patterns created by management events on metapopulation viability. Both occasional long-distance dispersal and dormant seeds are shown to be critical for the long-term survival of C. epithymum in a dynamic heathland landscape subjected to a fixed rotational mowing of 15 yr. A relatively high management frequency (<15 yr between two consecutive mowing events) appeared to be necessary to sustain a viable C. epithymum metapopulation. When there is a longer interval between management events, grazing can counterbalance the negative effects of vegetation succession. Our results indicate that small-scale cyclical management events combined with extensive grazing are the most appropriate management strategy to maintain viable populations of C. epithymum instead of the current large-scale management events. Our results further emphasise the importance of incorporating both spatiotemporal patch availability and key demographic characteristics, especially seed banks, for a realistic view of metapopulation dynamics in disturbed landscapes. This study clearly demonstrates the usefulness of metapopulation models to understand the impact of management events and to provide new ecological insights into processes acting at a landscape scale. [ABSTRACT FROM AUTHOR]

Abstract: A good understanding of the relationship between plant establishment and the ecosystem of which they are part of is needed to conserve rare plant species. Introduction experiments offer a direct test of recruitment limitation, but generally only the seed germination and seedling phases are monitored. Thus the relative importance of different establishment stages in the process of recruitment is not considered. This is particularly true for parasitic plants where empirical data are generally missing. During two consecutive growing seasons we examined the effect of heathland management applications, degree of heathland succession (pioneer, building and mature phase) and seed-density on the recruitment and establishment of the endangered holoparasite Cuscuta epithymum. In general, recruitment after two growing seasons was low with 4.79% of the sown seeds that successfully emerged to the seedling stage and a final establishment of 89 flowering adults (i.e. <1.5% of the sown seeds). Although a higher seed-density resulted in a higher number of seedlings, seed-density did not significantly affected relative germination percentages. The management type and subsequent heath succession had no significant effect on seedling emergence; whereas, seedling attachment to the host, establishment and growth to full-grown size were hampered in older heath vegetation (i.e. high, dense, and mature canopy). Establishment was most successful in turf-cut pioneer heathland, characterised by a relatively open and low vegetation of young Calluna vulgaris. The age of C. vulgaris, C. epithymum''s main host, proved to be the most limiting factor. These results emphasise the importance of site quality (i.e. successional phase of its host) on recruitment success of C. epithymum, which is directly affected by the management applied to the vegetation. Lack of any heathland management will thus seriously restrict establishment of the endangered parasite. [Copyright &y& Elsevier]

The impact of forest edge structure on edge patterns of wind speed, turbulence, and atmospheric deposition was studied by means of a model forest in a wind tunnel. Tests were conducted with eight structure configurations, encompassing combinations of stem densities, crown depths, and edge transitions (steep or gradual edge). Mean wind speed and its standard deviation (as a measure for turbulence) were determined within and at the top of the canopy; deposition was simulated using Cl− aerosols. Edge patterns of wind speed, turbulence, and deposition were closely related and were significantly affected by stem density and, particularly, by edge transition. In the dense forests, the edge effect on deposition extended less deeply into the forest than in the sparse forests, so the deposition in the forest edge zone was lowered with 40%. Gradual edges were able to limit the level by which deposition is enhanced at the edge in comparison with the forest interior deposition, and consequently, they reduced the deposition in the forest edge zone with 66%. Even when the deposition on the trees of the gradually ascending vegetation in front of the forest edge was taken into account, gradual edges were still advantageous in comparison with steep edges. A lower crown depth decreased the enhancement of deposition at the edge relative to the interior, but only at steep edges. We conclude that an adjusted layout of forest edges should be able to mitigate the edge effects on atmospheric deposition, through reducing the deposition enhancement at the edge or the penetration depth of the edge effect. [Copyright &y& Elsevier]

In recent decades, European protected forest areas primarily dedicated to biodiversity conservation have been increasingly selected using standard criteria that refer to three guiding principles in conservation planning: sufficient representativeness, adequate spatial design and adequate site suitability. The way these criteria are currently used by land planners was assessed through a standardised questionnaire sent to representatives from 21 European countries involved in the COST E27 action, and compared to prescriptions from the scientific literature. Results show that only 26% of the 101 types of protected forest areas, distributed in nine different countries, have been selected on the basis of all three guiding principles. For many protected area networks, insufficient importance is given to spatial design and site conditions, with the result that long-term persistence of biodiversity is far from guaranteed. The use of quantitative objectives, operational targets and optimisation approaches is also infrequent, in spite of repeated commitments of European forest ministers (MCPFE) to establish coherent, comprehensive and representative forest conservation networks. Overall, the adoption of a more systematic approach in forest conservation planning seems to be hindered by a dearth of operational guidelines to assess spatial design and site suitability, by insufficient collaboration between biologists and land planners, and by a shortage of funding to establish conservation networks. [Copyright &y& Elsevier]

Positive diversity–productivity relationships have repeatedly been found in experimental grassland plots, but mechanistic explanations are still under debate. We tested whether complementarity for the exploitation of the soil water resource helps to explain these relationships. In the dry summer of 2003, evapotranspiration (ET) was assessed at the Swedish BIODEPTH site using two different approaches: snapshot measurements of canopy surface temperature and simulation of time-accumulated ET by means of a soil water balance model. More diverse plots were characterized by lower surface temperatures and higher accumulated ET. Transgressive overyielding tests revealed that ET in polycultures was higher than in the best-performing monocultures, but this pattern was reversed at high degrees of water stress. Our results indicate that a more complete exploitation of soil water by more diverse grassland systems is on the one hand likely to be a driver for their increased biomass production, but on the other hand causes the more diverse communities to be affected earlier by drought. Nevertheless, the results also suggest that productivity may (at least partially) be maintained under dry conditions due to the higher likelihood of including drought-tolerant species in the more diverse communities. [ABSTRACT FROM AUTHOR]

Whether non-native plant invasions are causes, consequences, or independent of the low species diversity in recipient ecosystems remains a debated question. We tried to test these three hypotheses in the special case of the American black cherry ( Prunus serotina Ehrh.), a gap-dependent tree species, which is invading European temperate forests. We compared plant communities, soil properties, and disturbance history between P. serotina-invaded and uninvaded paired-stands in a managed mixed forest. Relationships between invasion, disturbances, plant communities, and environmental conditions were investigated using redundancy analyses with variation partitioning. Several soil characteristics differed between paired stands, but were rather components of stand invasibility than invasion effects, except for topsoil available phosphorus. The disturbance history was similar among paired stands except for the amount of storm-induced tree falls, which correlated with the invader's density. Wild boar-disturbed soil areas were more important beneath P. serotina canopies, suggesting a positive feedback on its own establishment. Overall, species assemblages in invaded and uninvaded stands were similar; their ecological inconsistency suggested a management-sustained non-equilibrium. Habitat conditions and disturbances explained most of the variation in both plant diversity and P. serotina density, the last two factors exhibiting a weak direct association. We conclude that in managed forest ecosystems where plant communities are mainly driven by non-interactive factors and immigration processes, non-native plant species can naturalize without being directly influenced by measured features of the plant community in the receiving environment on the short term. [ABSTRACT FROM AUTHOR]

The design of cost-efficient control strategies for invasive species that are too widespread and abundant for complete eradication, at least in the short term, will benefit from a rigorous analysis of invasion patterns and associated effects on native biodiversity. In this paper, the case of the invasive North American tree Prunus serotina in Flanders (Belgium) is presented. Our main objectives were to determine the susceptibility of forest stands to invasion by P. serotina and the subsequent effects of invasion on the understorey community. We used the large database of the first Flemish Forest Inventory. Multiple logistic regressions indicated that P. serotina occurred more frequently in privately owned, younger forest on coarse-textured, dry soils (podzols), and the combination of these factors allowed us to correctly predict presence/absence of P. serotina in 70% of the validation plots. However, locational variables proved to be important as well, indicating that the invasion process is still ongoing. Prediction of P. serotina densities by means of multiple linear regressions was less successful. Effects on the understorey richness were analysed by comparing the number of species and the mean Ellenberg values between pairs of plots, only differing by the presence of P. serotina in the shrub layer. A reduction of the understorey richness following invasion was only pronounced on the more moist soils, while compositional changes mainly occurred on drier soils. It is concluded that priority for control should be given to landscapes with a low fraction of invaded stands and to forest stands located on more moist soils. However, using its potential to threaten native biodiversity as an argument for control should be done with care as further research is needed whether the observed negative effects are due to a species (i.e. native vs. non-native) or a density effect (high vs. low). [ABSTRACT FROM AUTHOR]

Particularly in the temperate climate zone many forests have, at some moment in their history, been used as agriculture land. Forest cover is therefore often not as stable as it might look. How forest plant communities recovered after agriculture was abandoned allows us to explore some universal questions on how dispersal and environment limit plant species abundance and distribution. All studies looking at the effects of historical land use rely on adequate land use reconstruction. A variety of tools from maps, archival studies, and interviews to field evidence and soil analyses contribute to that. They allow us to distinguish ancient from recent forests and many studies found pronounced differences in forest plant species composition between them. A considerable percentage of our forest flora is associated with ancient forests. These ancient forest plant species (AFS) all have a low colonization capacity, suggesting that dispersal in space (distance related) and time (seed bank related) limit their distribution and abundance. However recent forests generally are suitable for the recruitment of AFS. There is clear evidence that dispersal limitation is more important than recruitment limitation in the distribution of AFS. Dispersal in time, through persistent seed banks, does not play a significant role. Ancient forests are not necessary more species-rich than recent forest, but if diversity is limited to typical forest plant species then ancient forests do have the highest number of plant species, making them highly important for nature conservation. The use of molecular markers, integrated approaches and modelling are all part of the way forward in this field of historical ecology. [ABSTRACT FROM AUTHOR]

1 Disturbance may cause community composition across sites to become more or less homogenous, depending on the importance of different processes involved in community assembly. In north-eastern North America and Europe local (alpha) diversity of forest plants is lower in forests growing on former agricultural fields (recent forests) than in older (ancient) forests, but little is known about the influence of land-use history on the degree of compositional differentiation among sites (beta diversity). 2 Here we analyse data from 1446 sites in ancient and recent forests across 11 different landscapes in north-eastern North America and Europe to demonstrate decreases in beta diversity and in the strength of species–environment relationships in recent vs. ancient forests. 3 The magnitude of environmental variability among sites did not differ between the two forest types. This suggests the difference in beta diversity between ancient and recent forests was not due to different degrees of environmental heterogeneity, but rather to dispersal filters that constrain the pool of species initially colonizing recent forests. 4 The observed effects of community homogenization and weakened relationships between species distributions and environmental gradients appear to persist for decades or longer. The legacy of human land-use history in spatial patterns of biodiversity may endure, both within individual sites and across sites, for decades if not centuries. [ABSTRACT FROM AUTHOR]

Questions: 1. How are the long-term dynamics of the root hemiparasite Rhinanthus angustifolius related to vegetation structure, grassland management and climate? 2. Does R. angustifolius have a long-term impact on standing crop and community composition? Location: A formerly fertilized grassland, part of a larger brook valley system in the nature reserve ‘Drentsche Aa’, near Groningen, The Netherlands. Methods: Vegetation development has been monitored since 1973 in 54 permanent plots in nine management regimes without fertilizer application. Results: 1. The hemiparasite established when standing crop was less than 600 g.m-2 and performed best under annual haymaking using machinery. Since its appearance, the population fluctuated stochastically, with two peaks. Coinciding collapses in six adjacent grasslands and comparison with an integrated climatic index suggest that the population collapses are induced by spring drought. 2. We did not find a relationship between total standing crop and R. angustifolius cover. Cover of grasses was negatively related to the abundance of the hemiparasite in the same and the previous year. Forb cover tended to increase with the parasite. Conclusions: R. angustifolius shows stochastic population fluctuations, mainly determined by spring drought, to which this species is probably highly vulnerable because of its parasitic and annual life style without a persistent seed bank. The hemiparasite also shows long-term relationships with grass cover (negative) and forb cover (positive), but it seems to have no lasting impact on standing crop. [ABSTRACT FROM AUTHOR]

Following habitat fragmentation individual habitat patches may lose species over time as they pay off their "extinction debt." Species with relatively low rates of population extinction and colonization ("slow" species) may maintain extinction debts for particularly prolonged periods, but few data are available to test this prediction. We analyzed two unusually detailed data sets on forest plant distributions and land-use history from Lincolnshire, United Kingdom, and Vlaams-Brabant, Belgium, to test for an extinction debt in relation to species-specific extinction and colonization rates. Logistic regression models predicting the presence-absence of 36 plant species were first parameterized using data from Lincolnshire, where forest cover has been relatively low (∼5-8%) for the past 1000 years. Consistent with extinction debt theory, for relatively slow species (but not fast species) these models systematically underpredicted levels of patch occupancy in Vlaams- Brabant, where forest cover was reduced from ∼25% to <10% between 1775 and 1900 (it is presently 6.5%). As a consequence, the ability of the Lincolnshire models to predict patch occupancy in Vlaams-Brabant was worse for slow than for fast species. Thus, more than a century after forest fragmentation reached its current level an extinction debt persists for species with low rates of population turnover. [ABSTRACT FROM AUTHOR]

Abstract: In this study, the effectiveness of management measures aimed at promoting recruitment in the last large Flemish juniper population at Heiderbos is evaluated. We resurveyed demographic plots 23 years after their establishment in 1980 and linked the population changes with detailed records of the intensive management during the same period. Between 1980 and 2003 the population size has decreased by 36% and has changed from a relatively immature to a mature population with very few young individuals. Based on a simple model that simulated height growth between 1980 and 2003, the recruitment and mortality rates were estimated to be 5 and 24ha−1 yr−1, respectively. Intensive management has thus not been able to promote recruitment and the population might go extinct within 40 years if these rates remain unchanged. Furthermore, some measures, notably the working of the soil, have increased mortality of established junipers. The reasons for the limited recruitment are not entirely clear yet, but it may be due to a combination of the limited availability of bare ground for germination and the extremely low viability of juniper seeds. The latter fact may be a common characteristic of many threatened juniper populations in northwestern Europe. [Copyright &y& Elsevier]

Abstract: Epizoochorous dispersal of plant seeds is an important long-distance dispersal mechanism. Yet little is known about retention times of seeds in animal furs and hence about potential dispersal distances of the seeds. Here, we used marked seeds of 12 plant species to determine seed depletion curves on Galloway cattle and Haflinger horse in three vegetation types (forest, tall herbage vegetation and meadow), in both dry and rainy weather conditions. In the long fur of Galloway cattle, seeds were retained significantly longer than in the short fur of Haflinger horse. In general, seed retention times were not considerably affected by the structure of the surrounding vegetation. The impact of the weather was negligible, only affecting the retention of some plant species. Negative exponential functions were fitted to the seed depletion curves. Using the parameters of curve estimations in the different conditions of animal species and vegetation structure, half-life seed retention times of up to 13h for Galloway cattle and up to more than 4h for Haflinger horse could be calculated, with corresponding potential half-life dispersal distances in the order of magnitude of tens of metres to a few kilometres. Different seed traits correlated with seed retention times in the long cattle fur and in the short horse fur, respectively. [Copyright &y& Elsevier]

Analysing invasive spread from a landscape ecological perspective forms an important challenge in plant invasion ecology. The present study examines the effects of landscape structure on the spatial and temporal dynamics of an expanding black cherryPrunus serotinapopulation within a rural landscape in Flanders, Belgium, carrying a dense network of interconnected hedgerows. The study area, 251 ha in size, harboured a total of 2962P. serotinaindividuals. The population was characterised by a negative exponential age distribution, a high growth rate and an early and continuous reproduction throughout the species’ life cycle. The historical rate of spread of the species through the hedgerow network progressively increased with time, especially during the last decade. Spatial point pattern analysis revealed that the individuals had a significantly clustered distribution pattern and were spatially aggregated around seed sources, hedgerow intersections and roosting trees. Logistic regression analysis confirmed the effect of landscape structure onP. serotinaoccurrence, suggesting directional long distance dispersal by avian dispersal vectors, resulting in a differential seed pressure throughout the hedgerow network due to the preference of dispersing birds for roosting in structurally rich hedgerow with large trees near hedgerow intersections. Hence, the distribution ofP. serotinain agricultural landscapes was strongly mediated by dispersal processes. Furthermore, decreasing spatial aggregation along the species life cycle, with especially seedlings and saplings being significantly aggregated while adult individuals were mostly distributed at random, and a relative outward shift of seedling recruitment curves with time indicate density dependent mortality, probably caused by intraspecific competition. [ABSTRACT FROM AUTHOR]

In fragmented landscapes, habitat patches are often destroyed and created through time, though most metapopulation models treat patch networks as static. Here we present a generally applicable, modified version of Flanski's Incidence Function Model (IFM) that incorporates landscape dynamics (i.e., habitat patch turnover), and we parameterize the model with data on patch occupancy patterns for forest plants in central Lincolnshire, UK. The modified IFM provided a better, or equally good, fit to species' patch occupancy patterns than logistic regression. Estimated colonization and extinction rates, and the results of logistic regression analyses, varied significantly among species with different life history traits. For example, species with low seed production and predominantly short-distance seed dispersal showed lower rates of colonization and extinction and were more likely to show effects of patch age and connectivity on patch-level presence than species with the opposite set of traits. Model simulations demonstrate a profound negative influence of habitat turnover rate on metapopulation dynamics and persistence, particularly for slow-colonizing species. The incorporation of temporal habitat dynamics into the metapopulation paradigm will permit its application to organisms in a much wider range of real landscapes. [ABSTRACT FROM AUTHOR]

Forest herbs differ greatly in their capacity to recolonize secondary forests established on former agricultural land. We investigated whether interspecific differences in recruitment or growth can account for differences in colonization success. Seeds and adults of two species with limited colonization capacities (Anemone nemorosa and Primula elatior) and two species with high colonization capacities (Ranunculus ficaria and Geum urbanum ) were introduced in ancient and recent forest sites within the Muizen forest (Belgium). At all sites, half of the plots were also cleared of above-ground vegetation. Seedling establishment was similar in recent and ancient forest sites. However, both the number and subsequent survival of seedlings of the two hemicryptophytes (G. urbanum and P. elatior ) were significantly lower when vegetation cover was present, while seedling number and survival of the two geophytes (A. nemorosa and R. ficaria) were less influenced by vegetation cover. Adult performance of P. elatior and G. urbanum was significantly better in the nutrient enriched recent forest stands and in the absence of vegetation cover. Performance of A. nemorosa and R. ficaria was not affected by any of these factors. We conclude that the extent of seed limitation is probably the main cause for the different colonization capacities of the species investigated. [ABSTRACT FROM AUTHOR]

Summary 1 A framework that summarizes the direct and indirect effects of past land use on forest herb recolonization is proposed, and used to analyse the colonization patterns of forest understorey herbaceous species in a 360-ha mixed forest, grassland and arable landscape in the Dijle river valley (central Belgium). 2 Fine-scale distribution maps were constructed for 14 species. The species were mapped in 15 946 forest plots and outside forests (along parcel margins) in 5188 plots. Forest stands varied in age between 1 and more than 224 years. Detailed land-use history data were combined with the species distribution maps to identify species-specific colonization sources and to calculate colonization distances. 3 The six most frequent species were selected for more detailed statistical analysis. 4 Logistic regression models indicated that species frequency in forest parcels was a function of secondary forest age, distance from the nearest colonization source and their interaction. Similar age and distance effects were found within hedgerows. 5 In 199 forest stands, data about soils, canopy structure and the cover of competitive species were collected. The relative importance of habitat quality and spatio-temporal isolation for the colonization of the forest herb species was quantified using structural equation modelling (SEM), within the framework proposed for the effects of past land use. 6 The results of the SEM indicate that, except for the better colonizing species, the measured habitat quality variables are of minor importance in explaining colonization patterns, compared with the combination of secondary forest age and distance from colonization sources. 7 Our results suggest the existence of a two-stage colonization process in which diaspore availability determines the initial pattern, which is affected by environmental sorting at later stages. [ABSTRACT FROM AUTHOR]

Summary 1 Classifying species by shared functional characteristics is important if common functional response groups are to be identified among different taxa. 2 We investigated plant traits that determine the response of forest plant species to land use changes using literature data. Sources from eight European countries and four North-eastern American states, comprising 20 field studies yielded information on 216 forest plant species. For these species, data on 13 life history traits were collected. 3 Trait correlation structure was similar in the European and American data-sets and corresponded well to those described in the literature. The European and American herbs and the European graminoids were clustered into distinct emergent groups based on their plant traits. The profiles of the European and American emergent groups were similar. 4 Herb species belonging to emergent groups characterized by low dispersability (i.e. large seeds, low fecundity, unassisted dispersal) were relatively slow colonizers. Dispersability (and not recruitment) seems to be a key factor limiting the colonization of some forest plant species. The relationship between dispersability and colonizing capacity was less clear for graminoids. 5 A life history trait-based approach offers good opportunities to gain insight into the mechanisms behind species response to land-use change. [ABSTRACT FROM AUTHOR]

Global circulation models predict an increase in mean annual temperature between 2.1 and 4.6 °C by 2080 in the northern temperate zone. The associated changes in the ratio of extinctions and colonizations at the boundaries of species ranges are expected to result in northward range shifts for a lot of species. However, net species colonization at northern boundary ranges, necessary for a northward shift and for range conservation, may be hampered because of habitat fragmentation. We report the results of two forest plant colonization studies in two fragmented landscapes in central Belgium. Almost all forest plant species (85%) had an extremely low success of colonizing spatially segregated new suitable forest habitats after c . 40 years. In a landscape with higher forest connectivity, colonization success was higher but still insufficient to ensure large-scale colonization. Under the hypothesis of net extinction at southern range boundaries, forest plant species dispersal limitation will prevent net colonization at northern range boundaries required for range conservation. [ABSTRACT FROM AUTHOR]

Abstract 1 Distribution patterns (frequency and percentage cover) of 18 forest plant species were studied in 34 ha of mixed deciduous forest (Muizen Forest, north Belgium). Stands varied in age between 6 and more than 223 years and both slow and fast colonizing species were studied. 2 Detailed land use history data were combined with the species distribution maps to identify species-specific colonization sources and calculate colonization distances. 3 A multiple logistic regression model was constructed with four covariables: pH (which can impose limits on the potential species-distribution), secondary forest age, distance from nearest colonization source and age–distance interaction, to allow us to account for the gradual completion of colonization over time. 4 We could distinguish species which are limited by both dispersal and recruitment (Primula elatior , Arum maculatum and Lamium galeobdolon ), mainly by dispersal (Anemone nemorosa, Deschampsia cespitosa ), mainly by recruitment (Paris quadrifolia and Polygonatum multiflorum ) and by neither (Geum urbanum , Ranunculus ficaria , Glechoma hederacea , Aegopodium podagraria , Ajuga reptans , Adoxa moschatellina and Oxalis acetosella ). 5 The low colonizing capacity of ancient forest plants cannot be attributed to a single cause; rather both dispersal and recruitment are limiting but the relative importance varies. [ABSTRACT FROM AUTHOR]

An integrated analysis of the colonization patterns of forest plant species was carried out in a 34-ha, mixed deciduous forest in northern Belgium. First, we sought to describe the relationships between land use history and environmental conditions. Land use history and soil type were related and negative correlations between pH and secondary forest age. Litter quantity and cover of Urtica dioica were mainly indirectly influenced by land use history. Litter starts accumulating at low pH values and high shrub density and Urtica dioica grows vigorously on nutrient enriched soils where much light can reach the ground. Next, the importance of these human-altered environmental conditions for the colonization of forest plant species was assessed relative to the importance of dispersal limitation. Therefore, the distribution of 16 forest species was mapped and species-specific spatio-temporal isolation measures were calculated. The analysis revealed that the colonization patterns of the slowly colonizing species (i.e. 'ancient forest plant species') are best explained by a combination of spatio-temporal isolation, soil type, pH and the (non-)cover of Urtica dioica. By contrast, spatio-temporal isolation was never a limiting factor for good colonizing forest species. Our results suggest that a combination of dispersal and recruitment limitation and that the relative importance of both factors is species specific. [ABSTRACT FROM AUTHOR]

Questions: Effective and successful restoration of species-rich semi-natural grasslands requires knowledge of the soil nutrient status, including soil phosphorus availability. Plants are solid indicators of soil nutrient status, because their growth reflects nutrient availability integrated over a certain period. The use of reflectance spectroscopy to estimate vegetation nutrient content has the potential to offer a fast and efficient approach to provide information about soil nutrient availability for plants. Here, we investigated the effect and relative importance of vegetation phosphorus content compared with vegetation nitrogen content and species identity in explaining variation in vegetation reflectance. Location: Pot experiment mimicking Western European grassland communities on a restoration trajectory. Methods: We combined a pot experiment with a broad range of mesotrophic grassland species growing along a soil phosphorus gradient with a multivariate modelling approach. We measured vegetation spectra and vegetation phosphorus and nitrogen content in monocultures and mixtures. Results: Although vegetation biochemistry explained a considerable part of the variation in vegetation reflectance, we found no pronounced absorption features for vegetation nitrogen and phosphorus content across the reflectance spectrum. The relatively large effect of species and community identity suggest that other drivers, for example vegetation architecture, overruled the effect of vegetation biochemistry on the reflectance spectrum. Our findings indicate that species detection and indirect case-specific estimation of nutrients is possible, especially in structurally less-complex canopies such as monospecific grass swards. Conclusions: Disentangling the specific drivers of variation in spectral reflectance is challenging. Many studies confound the effect of species identity on the vegetation reflectance spectrum with the effect of vegetation biochemistry. Here, we showed the importance of explicitly taking species identity into account. Gaining insight into light-vegetation interactions and the in-depth integration of ecological theory in remote sensing are the way forward. [ABSTRACT FROM AUTHOR]

Background: Across the tropics, the share of secondary versus primary forests is strongly increasing. The high rate of biomass accumulation during this secondary succession relies on the availability of essential nutrients, such as nitrogen (N). Nitrogen primarily limits many young secondary forests in the tropics. However, recent studies have shown that forests of the Congo basin are subject to high inputs of atmospheric N deposition, potentially alleviating this N limitation in early succession. Methods: To address this hypothesis, we assessed the N status along a successional gradient of secondary forests in the Congo basin. In a set-up of 18 plots implemented along six successional stages, we quantified year-round N deposition, N leaching, N2O emission and the N flux of litterfall and fine root assimilation. Additionally, we determined the N content and C:N stoichiometry for canopy leaves, fine roots, and litter, as well as δ15N of canopy leaves. Results: We confirmed that these forests receive high amounts of atmospheric N deposition, with an increasing deposition as forest succession proceeds. Additionally, we noted lower C:N ratios, and higher N leaching losses, N2O emission, and foliar δ15N in older secondary forest (60 years). In contrast, higher foliar, litter and root C:N ratios, and lower foliar δ15N, N leaching, and N2O emission in young (< 20 years) secondary forest were observed. Conclusions: Altogether, we show that despite high N deposition, this early forest succession still shows conservative N cycling characteristics, which are likely indicating N limitation early on in secondary forest succession. As secondary succession advances, the N cycle gradually becomes more open. [ABSTRACT FROM AUTHOR]

Summary Recent droughts have strongly impacted forest ecosystems and are projected to increase in frequency, intensity, and duration in the future together with continued warming. While evidence suggests that tree diversity can regulate drought impacts in natural forests, few studies examine whether mixed tree plantations are more resistant to the impacts of severe droughts. Using natural variations in leaf carbon (C) and nitrogen (N) isotopic ratios, that is δ13C and δ15N, as proxies for drought response, we analyzed the effects of tree species richness on the functional responses of tree plantations to the pan‐European 2018 summer drought in seven European tree diversity experiments. We found that leaf δ13C decreased with increasing tree species richness, indicating less drought stress. This effect was not related to drought intensity, nor desiccation tolerance of the tree species. Leaf δ15N increased with drought intensity, indicating a shift toward more open N cycling as water availability diminishes. Additionally, drought intensity was observed to alter the influence of tree species richness on leaf δ15N from weakly negative under low drought intensity to weakly positive under high drought intensity. Overall, our findings suggest that dual leaf isotope analysis helps understand the interaction between drought, nutrients, and species richness. [ABSTRACT FROM AUTHOR]

Decomposition is a key determinant of forest functioning, controlling nutrient and carbon cycling. Although litter-mixing effects on decomposition (that is, using mixtures of litter of different species) have been studied extensively, less is known about the indirect effects of modified microenvironments via overstory tree species mixing. To investigate the effects of tree species diversity on decomposition, we installed 384 standardized litterbags, filled with leaf litter of four broadleaved tree species with contrasting litter quality, in a large, 10-year-old tree diversity experiment. To quantify microenvironments, we used microclimate sensors, below-canopy rain gauges and measured soil characteristics. We then analysed indirect tree species diversity effects, that is, tree species richness effects on mass loss rates via tree species-induced alterations in the microclimate, throughfall and soil characteristics. We found that understory microenvironmental conditions indeed affect mass loss rates, with the main drivers differing among incubation stages. Predominantly soil phosphorus, but also vapour pressure deficit and throughfall amounts, was negatively associated with mass loss rates across litter types during the first 2 months of the decomposition process. After 6 months of the decomposition, soil moisture was found to be the key determinant positively affecting mass loss rates. In sum, our research contributes to a better understanding of the determinants of decomposition and shows an important pathway in which tree species diversity affects decomposition, via modified microenvironmental conditions acting via the soil, microclimate and throughfall. [ABSTRACT FROM AUTHOR]