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Forest restoration increases global forest area and ecosystem services such as primary productivity and carbon storage. How tree species functional composition impacts the provisioning of these services as forests develop is sparsely studied. We used 10-year data from 478 plots with 191,200 trees in a forest biodiversity experiment in subtropical China to assess the relationship between community productivity and community-weighted mean (CWM) or functional diversity (FD) values of 38 functional traits. We found that effects of FD values on productivity became larger than effects of CWM values after 7 years of forest development and that the FD values also became more reliable predictors of productivity than the CWM values. In contrast to CWM, FD values consistently increased productivity across ten different species-pool subsets. Our results imply that to promote productivity in the long term it is imperative for forest restoration projects to plant multispecies communities with large functional diversity.

International audience; Forests are ecosystems critical to understanding the global carbon budget, due to their carbon sequestration potential in both aboveground and belowground compartments, especially in species-rich forests. Soil carbon sequestration is strongly linked to soil microbial communities, and this link is mediated by the tree community, likely due to modifications of microenvironmental conditions (i.e., biotic conditions, soil properties, and microclimate). We studied soil carbon concentration and the soil microbial biomass of 180 local neighborhoods along a gradient of tree species richness ranging from 1 to 16 tree species per plot in a Chinese subtropical forest experiment (BEF-China). Tree productivity and different tree functional traits were measured at the neighborhood level. We tested the effects of tree productivity, functional trait identity, and dissimilarity on soil carbon concentrations, and their mediation by the soil microbial biomass and microenvironmental conditions. Our analyses showed a strong positive correlation between soil microbial biomass and soil carbon concentrations. In addition, soil carbon concentration increased with tree productivity and tree root diameter, while it decreased with litterfall C:N content. Moreover, tree productivity and tree functional traits (e.g., fungal root association and litterfall C:N ratio) modulated microenvironmental conditions with substantial consequences for soil microbial biomass. We also showed that soil history and topography should be considered in future experiments and tree plantations, as soil carbon concentrations were higher at sites where historical (i.e., at the beginning of the experiment) carbon concentrations were high, themselves being strongly affected by the topography. Altogether, these results implied that the quantification of the different soil carbon pools is critical for understanding microbial community–soil carbon stock relationships and their dependence on tree diversity and microenvironmental conditions.

Previous studies have shown that tree species richness increases forest productivity by allowing for greater spatial complementarity of tree crowns (crown complementarity), which in turn results in more densely packed canopies. However, the mechanisms driving crown complementarity in tree species mixtures remain unclear. Here, we take advantage of a high-resolution, three-dimensional terrestrial laser scanning approach in the context of a large-scale biodiversity-ecosystem functioning experiment in subtropical China (BEF-China) to quantify the extent to which functional dissimilarity and divergences in branch traits between neighbouring trees affect crown complementarity at the scale of tree species pairs (i.e., two adjacent trees). Overall, we found no support that functional dissimilarity (divergence in morphological flexibility, specific leaf area and wood density) promotes crown complementarity. However, we show that the effects of functional dissimilarity (the plasticity of the outer crown structure) on crown complementarity vary in their magnitude and importance depending on branch trait divergences. Firstly, crown complementarity tended to be highest for tree species pairs that strongly differed in their functional traits, but were similar in branch density. In contrast, heterospecific pairs with a low functional trait divergence benefitted the most from a large difference in branch density compared with pairs characterised by a large functional dissimilarity. Secondly, the positive effects of increasing divergence in branching intensity (the plasticity of the inner crown structure) on crown complementarity became most important at low levels of functional dissimilarity, i.e. when species pairs were similar in their branch packing and vice versa. This suggests that species mixing allows trees to occupy canopy space more efficiently mainly due to phenotypic changes associated with crown morphology and branch plasticity. Our findings highlight the importance of considering outer and inner crown structures (e.g. branching architecture) to deepen our understanding of tree-tree interactions in mixed-species communities.

Global change exposes forest ecosystems to many risks including novel climatic conditions, increased frequency of climatic extremes and sudden emergence and spread of pests and pathogens. At the same time, forest landscape restoration has regained global attention as an integral strategy for climate change mitigation. Owing to unpredictable future risks and the need for new forests that provide multiple ecosystem services, mixed-species forests have been advocated for this purpose. However, the successful establishment of mixed forests requires intrinsic knowledge of biodiversity's role for forest ecosystem functioning. In this respect, a better understanding of tree-tree interactions and how they contribute to observed positive tree species richness effects on key ecosystem functions is critical. Here, we review the current knowledge of the underlying mechanisms of tree-tree interactions and argue that positive net biodiversity effects at the community scale may emerge from the dominance of positive over negative interactions at the local neighbourhood scale. In a second step, we demonstrate how tree-tree interactions and the immediate tree neighbourhood's role can be systematically assessed in a tree diversity experiment. The expected results will improve predictions about the effects of tree interactions on ecosystem functioning based on general principles. We argue that this knowledge is urgently required to guide the design of tree species mixtures for the successful establishment of newly planted forests.

Background: Species-specific genotypic features, local neighbourhood interactions and resource supply stronglyinfluence the tree stature and growth rate. In mixed-species forests, diversity-mediated biomass allocation has beensuggested to be a fundamental mechanism underlying the positive biodiversity-productivity relationships. Empiricalevidence, however, is rare about the impact of local neighbourhood diversity on tree characteristics analysed at avery high level of detail. To address this issue we analysed these effects on the individual-tree crown architectureand tree productivity in a mature mixed forest in northern Germany. Methods: Our analysis considers multiple target tree species across a local neighbourhood species richnessgradient ranging from 1 to 4. We applied terrestrial laser scanning to quantify a large number of individual maturetrees (N= 920) at very high accuracy. We evaluated two different neighbour inclusion approaches by analysing botha fixed radius selection procedure and a selection based on overlapping crowns. Results and conclusions: We show that local neighbourhood species diversity significantly increases crowndimension and wood volume of target trees. Moreover, we found a size-dependency of diversity effects on treeproductivity (basal area and wood volume increment) with positive effects for large-sized trees (diameter at breastheight (DBH) > 40 cm) and negative effects for small-sized (DBH < 40 cm) trees. In our analysis, the neighbourinclusion approach has a significant impact on the outcome. For scientific studies and the validation of growthmodels we recommend a neighbour selection by overlapping crowns, because this seems to be the relevant scaleat which local neighbourhood interactions occur. Because local neighbourhood diversity promotes individual-treeproductivity in mature European mixed-species forests, we conclude that a small-scale species mixture should beconsidered in management plans. Background: Species-specific genotypic features, local neighbourhood interactions and resource supply strongly influence the tree stature and growth rate. In mixed-species forests, diversity-mediated biomass allocation has been suggested to be a fundamental mechanism underlying the positive biodiversity-productivity relationships. Empirical evidence, however, is rare about the impact of local neighbourhood diversity on tree characteristics analysed at a very high level of detail. To address this issue we analysed these effects on the individual-tree crown architecture and tree productivity in a mature mixed forest in northern Germany. Methods: Our analysis considers multiple target tree species across a local neighbourhood species richness gradient ranging from 1 to 4. We applied terrestrial laser scanning to quantify a large number of individual mature trees (N = 920) at very high accuracy. We evaluated two different neighbour inclusion approaches by analysing both a fixed radius selection procedure and a selection based on overlapping crowns. Results and conclusions: We show that local neighbourhood species diversity significantly increases crown dimension and wood volume of target trees. Moreover, we found a size-dependency of diversity effects on tree productivity (basal area and wood volume increment) with positive effects for large-sized trees (diameter at breast height (DBH) > 40 cm) and negative effects for small-sized (DBH < 40 cm) trees. In our analysis, the neighbour inclusion approach has a significant impact on the outcome. For scientific studies and the validation of growth models we recommend a neighbour selection by overlapping crowns, because this seems to be the relevant scale at which local neighbourhood interactions occur. Because local neighbourhood diversity promotes individual-tree productivity in mature European mixed-species forests, we conclude that a small-scale species mixture should be considered in management plans.

Previous studies have shown that tree species richness increases forest productivity by allowing for greater spatial complementarity of tree crowns (crown complementarity), which in turn results in more densely packed canopies. However, the mechanisms driving crown complementarity in tree species mixtures remain unclear. Here, we take advantage of a high-resolution, three-dimensional terrestrial laser scanning approach in the context of a large-scale biodiversity-ecosystem functioning experiment in subtropical China (BEF-China) to quantify the extent to which functional dissimilarity and divergences in branch traits between neighbouring trees affect crown complementarity at the scale of tree species pairs (i.e., two adjacent trees). Overall, we found no support that functional dissimilarity (divergence in morphological flexibility, specific leaf area and wood density) promotes crown complementarity. However, we show that the effects of functional dissimilarity (the plasticity of the outer crown structure) on crown complementarity vary in their magnitude and importance depending on branch trait divergences. Firstly, crown complementarity tended to be highest for tree species pairs that strongly differed in their functional traits, but were similar in branch density. In contrast, heterospecific pairs with a low functional trait divergence benefitted the most from a large difference in branch density compared with pairs characterised by a large functional dissimilarity. Secondly, the positive effects of increasing divergence in branching intensity (the plasticity of the inner crown structure) on crown complementarity became most important at low levels of functional dissimilarity, i.e. when species pairs were similar in their branch packing and vice versa. This suggests that species mixing allows trees to occupy canopy space more efficiently mainly due to phenotypic changes associated with crown morphology and branch plasticity. Our findings highlight the importance of considering outer and inner crown structures (e.g. branching architecture) to deepen our understanding of tree-tree interactions in mixed-species communities. Previous studies have shown that tree species richness increases forest productivity by allowing for greater spatial complementarity of tree crowns (crown complementarity), which in turn results in more densely packed canopies. However, the mechanisms driving crown complementarity in tree species mixtures remain unclear. Here, we take advantage of a highresolution,three-dimensional terrestrial laser scanning approach in the context of a large-scale biodiversity-ecosystem functioning experiment in subtropical China (BEF-China) to quantify the extent to which functional dissimilarity and divergences in branch traits between neighbouring trees affect crown complementarity at the scale of tree species pairs (i.e., two adjacent trees). Overall, we found no support that functional dissimilarity (divergence in morphological flexibility, specific leaf area and wood density) promotes crown complementarity. However, we show that the effects of functional dissimilarity (the plasticity of the outer crown structure) on crown complementarity vary in their magnitude and importance depending on branch trait divergences. Firstly, crown complementarity tended tobe highest for tree species pairs that strongly differed in their functional traits, but were similar in branch density. In contrast, heterospecific pairs with a low functional trait divergence benefitted the most from a large difference in branch density compared with pairs characterised by a large functional dissimilarity. Secondly, the positive effects of increasing divergence in branching intensity (the plasticity of the inner crown structure) on crown complementarity became most important at low levels of functional dissimilarity, i.e. when species pairs were similar in their branch packing and vice versa. This suggests that species mixing allows trees to occupy canopy space more efficiently mainly due to phenotypicchanges associated with crown morphology and branch plasticity. Our findings highlight the importance of considering outer and inner crown structures (e.g. branching architecture) to deepen our understanding of tree-tree interactions in mixed-species communities.

Plant diversity loss can alter higher trophic -level communities via non -random species interactions, which in turn may cascade to affect key ecosystem func-tions. These non -random linkages might be best captured by patterns of phyloge-netic diversity, which take into account co -evolutionary dependencies. However,lack of adequate phylogenetic data of higher trophic levels hampers our mecha-nistic understanding of biodiversity relationships in species-rich ecosystems.2. We used DNA barcoding to generate data on the phylogenetic diversity of lepi-dopteran caterpillars in a large- scale forest biodiversity experiment in subtropicalChina. We analysed how different metrics of lepidopteran phylogenetic diversity(Faith's PD, MPD, MNTD) and taxonomic diversity were influenced by multiple components of tree diversity (taxonomic, functional, phylogenetic).3. Our data from six sampling periods represent 7,204 mitochondrial cytochrome c oxidase subunit I (COI) sequences of lepidopteran larvae, clustered into 461 mo-lecular operational taxonomic units. Lepidopteran abundance, the effective num-ber of species (irrespective of the focus on rare or common species) and Faith's PD and MPD (reflecting basal evolutionary splits), but not MNTD (reflecting re-cent evolutionary splits), significantly increased with experimentally manipulated ree species richness. Lepidopteran MNTD decreased with increasing tree MNTD. Path analyses showed that tree phylogenetic and functional diversity explained part, but not all of the effects of tree species richness on lepidopteran diversity. Importantly, tree diversity effects on lepidopteran diversity were to a large extent indirect, operating via changes in lepidopteran abundance. 4. Synthesis. Our study shows that evolutionary dependencies determine the response of herbivore communities to changes in host plant diversity. Incorporating a wider range of diversity metrics both at the level of producers and consumers can thus help to develop a more comprehensive understanding of the functional consequences of biodiversity change across trophic levels. Moreover, the dependence of trophic linkages on herbivore abundances underlines the need to address the consequences of current declines in insect abundances for ecosystem structure and functioning. peerReviewed

Biodiversity is considered to mitigate detrimental impacts of climate change on the functioning of forest ecosystems, such as drought-induced decline in forest productivity. However, previous studies produced controversial results and experimental evidence is rare. Specifically, the biological mechanisms underlying mitigation effects remain unclear, as existing work focuses on biodiversity effects related to the community scale. Using trait-based neighbourhood models, we quantified changes in above-ground wood productivity of 3,397 trees that were planted in a large-scale tree diversity experiment in subtropical China across gradients of neighbourhood diversity and climatic conditions over a 6-year period. This approach allowed us to simultaneously assess to what extent functional traits of a focal tree and biodiversity at the local neighbourhood scale mediate the growth response of individual trees to drought events. We found that neighbourhood tree species richness can mitigate for drought-induced growth decline of young trees. Overall, positive net biodiversity effects were strongest during drought and increased with increasing taxonomic diversity of neighbours. In particular, drought-sensitive species (i.e. those with a low cavitation resistance) benefitted the most from growing in diverse neighbourhoods, suggesting that soil water partitioning among local neighbours during drought particularly facilitated most vulnerable individuals. Thus, diverse neighbourhoods may enhance ecosystem resistance to drought by locally supporting drought-sensitive species in the community. Synthesis. Our findings demonstrate that mechanisms operating at the local neighbourhood scale are a key component for regulating forests responses to drought and improve insights into how local species interactions vary along stress gradients in highly diverse tree communities.

A large proportion of temperate forest plant diversity is found in the herb layer.However, for many of its species, little is known about their autecology, whichmakes it difficult to assess potential threats and efficiently safeguard the diversity of understorey herbaceous communities. This also applies to Gageaspathacea (Liliaceae), a globally rare spring geophyte, which mainly occurs indeciduous forests of northern Central Europe. We investigated the causal relationships between population characteristics of G. spathacea and abiotic siteconditions across different forest communities in the center of its distributionalrange. Leaf length (a surrogate of the species' vegetative propagation) was positively related to soil moisture and soil nitrogen. Consequently, mean leaflength was highest in moist forest communities of the alliance Alno-Ulmion.Moreover, mean variability in leaf length was lowest in those forests, indicatinga higher and more stable vegetative propagation via bulbils. We found no support for a significant relationship between leaf length and leaf density orbetween leaf length and flower formation. Population density varied stronglyamong forest sites, but was not related to soil moisture and hardly influenced by soil nitrogen. Our results suggest that soil water and nutrient supply play avital role in determining the species' vegetative propagation, whereas the duration of habitat continuity is most likely an important determinant of population size and density. Conservation strategies therefore require a betterunderstanding of the complex interrelationships between abiotic site conditionsand the historical context-dependency of habitats. A large proportion of temperate forest plant diversity is found in the herb layer. However, for many of its species, little is known about their autecology, which makes it difficult to assess potential threats and efficiently safeguard the diversity of understorey herbaceous communities. This also applies to Gagea spathacea (Liliaceae), a globally rare spring geophyte, which mainly occurs in deciduous forests of northern Central Europe. We investigated the causal relationships between population characteristics of G. spathacea and abiotic site conditions across different forest communities in the center of its distributional range. Leaf length (a surrogate of the species' vegetative propagation) was positively related to soil moisture and soil nitrogen. Consequently, mean leaf length was highest in moist forest communities of the alliance Alno‐Ulmion. Moreover, mean variability in leaf length was lowest in those forests, indicating a higher and more stable vegetative propagation via bulbils. We found no support for a significant relationship between leaf length and leaf density or between leaf length and flower formation. Population density varied strongly among forest sites, but was not related to soil moisture and hardly influenced by soil nitrogen. Our results suggest that soil water and nutrient supply play a vital role in determining the species' vegetative propagation, whereas the duration of habitat continuity is most likely an important determinant of population size and density. Conservation strategies therefore require a better understanding of the complex interrelationships between abiotic site conditions and the historical context‐dependency of habitats.

1. Diversity of producers (e.g. plants) usually increases the diversity of associated organisms, but the scale (i.e. the spatial area of plant diversity considered) at which plant diversity acts on other taxa has rarely been studied. Most evidence for cross‐taxon diversity relations come from above‐ground consumers that directly interact with plants.2. Experimental tests of plant diversity effects on elusive organisms inhabiting the leaf litter layer, which are important for nutrient cycling and decomposition, are rare.3. Using a large tree diversity experiment, we tested whether tree diversity at the larger plot (i.e. community) or the smaller neighbourhood scale relates to the abundance, species richness, functional and phylogenetic diversity of leaf litter ants, which are dominant organisms in brown food webs.4. Contrary to our expectations of scale‐independent positive tree diversity effects, ant diversity increased only with plot but not neighbourhood tree diversity. While the exact causal mechanisms are unclear, nest relocation or small‐scale competition among ants may explain the stronger tree diversity effects at the plot scale.5. Our results indicate that even for small and less mobile organisms in the leaf litter, effects of tree diversity are stronger at relatively larger scales. The finding emphasizes the importance of diverse forest stands, in which mixing of tree species is not restricted to small patches, for supporting arthropod diversity in the leaf litter. Diversity of producers (e.g. plants) usually increases the diversity of associated organisms, but the scale (i.e. the spatial area of plant diversity considered) at which plant diversity acts on other taxa has rarely been studied. Most evidence for cross-taxon diversity relations come from above-ground consumers that directly interact with plants. Experimental tests of plant diversity effects on elusive organisms inhabiting the leaf litter layer, which are important for nutrient cycling and decomposition, are rare. Using a large tree diversity experiment, we tested whether tree diversity at the larger plot (i.e. community) or the smaller neighbourhood scale relates to the abundance, species richness, functional and phylogenetic diversity of leaf litter ants, which are dominant organisms in brown food webs. Contrary to our expectations of scale-independent positive tree diversity effects, ant diversity increased only with plot but not neighbourhood tree diversity. While the exact causal mechanisms are unclear, nest relocation or small-scale competition among ants may explain the stronger tree diversity effects at the plot scale. Our results indicate that even for small and less mobile organisms in the leaf litter, effects of tree diversity are stronger at relatively larger scales. The finding emphasizes the importance of diverse forest stands, in which mixing of tree species is not restricted to small patches, for supporting arthropod diversity in the leaf litter.

The maintenance or enhancement of structural heterogeneity (SH) of forest stands is considered a key element of sustainable forest management in central Europe, making the monitoring of stand structure an important task. However, existing approaches that enable the quantification of SH usually neglect the three-dimensional (3D) representation of forest structures. To overcome this limitation, terrestrial laser scanning (TLS) can be applied to provide an efficient, non-destructive and high resolution record of 3D structures. In this study, we present a new structural heterogeneity index (SHITLS) at the plot level which is based on 3D point clouds of individual trees that were derived from multiple scans. Because tree crowns are of particular importance for the 3D SH of forests, we focus particularly on outer and inner crown attributes. To test the robustness and the explanatory power of our index we sampled data on 84 plots across 12 different forest stand types in two study areas of Germany. We analyzed whether our index is able to explain differences between the different stand types. In addition, we compared our SHITLS to an existing stand structural complexity index (SSCI) that is based purely on single TLS scans. We found different trends of SHITLS between stand types dominated by Fagus sylvatica compared to those dominated by coniferous tree species (mainly Picea abies and Pinus sylvestris). Correlation analyses showed a significant, but weak, correlation with SSCI. The advantage of the SHITLS is the use of multiple scans from several scan positions. Thus, a better and very high resolution coverage of the forest structures is achieved. Due to its TLS-based nature this method provides many benefits in terms of reproducibility, objectivity and simplicity of data evaluation.

The abandonment of traditional pastoralism as well as the use of heath areas for military purposes has had a major impact on dry heaths in the Continental biogeographical region of Europe, causing severe degradation of its key species Calluna vulgaris (L.) HULL. The reproductive potential of this species in a Continental climate is assumed to be low, although there is yet no observational or experimental evidence for this. More knowledge is also needed about cost-effective and sustainable measures to restore abandoned dry heaths in this biogeographical region, because traditional management options are often too expensive (e.g., sod-cutting) or restricted due to environmental laws and the danger of unexploded ammunition (e.g., burning). Using as an example an 800 ha Continental heathland in Germany that has been abandoned for about two decades, we studied the reproductive potential (seed production, soil seed bank, and germination ability) of degenerate C. vulgaris stands. In addition, we conducted a comprehensive field experiment to test the effects of low-intensity, year-round grazing by Heck cattle and Konik horses as well as one-time mowing and patchy exposure of bare soil on the generative rejuvenation (i.e., recruitment and survival) of degenerate C. vulgaris stands over 3 years. We used generalized linear mixed models for statistical analyses. Seed production of degenerate C. vulgaris stands was high as well as the germination ability of their seeds, being similar to Atlantic heathlands. However, soil seed-bank densities were lower than those found in managed or abandoned Atlantic heaths. Overall seedling recruitment in the field was considerably lower in comparison with Atlantic heaths. Low-intensity grazing or one-time mowing did not induce a substantial increase in C. vulgaris recruitment, whereas an additional one-time creation of bare soil patches or the one-time creation of bare soil without subsequent management significantly facilitated seedling recruitment and survival in the first year. However, from the second year on, the positive effect of the creation of bare soil without subsequent management was no longer present. In the third year, survival of juveniles was significantly supported by low-intensity grazing in combination with shallow soil disturbances as well as in combination with one-time mowing and shallow soil disturbances, whereas mowing alone resulted in marginally significant lower survival. The extremely low seedling recruitment requires a careful choice of suitable management measures to promote the survival of sufficient numbers of Calluna individuals. Therefore, we recommend low-intensity grazing with free-ranging robust breeds and the combination of this with one-time mowing as an effective means of supporting generative rejuvenation of C. vulgaris in degraded heaths. However, at the beginning of the restoration process, the creation of bare soil patches for seedling recruitment is crucial. For implementation into practice, we present different strategies to enhance the proportion of bare soil after long-term abandonment of heaths when traditional management options are no longer feasible.

1. Understanding how species diversity affects plant performance is a central question in biodiversity–ecosystem functioning (BEF) research. At the community level, functional trait means and trait dissimilarities have been used to explain biodiversity effects, but with mixed success. To disentangle how functional traits explain community growth and underpin biodiversity–ecosystem functioning relationships, we should elucidate how plant traits affect individual growth across species richness levels, because the role of functional traits on growth depends on the ecological context of the individual.2. We addressed this topic by using detailed data of twelve functional traits and annual growth for 529 individual trees of 31 species at five species‐richness levels in a large forest biodiversity experiment in south‐east China from 4 to 9 years after planting.3. Our analyses show first that individual trait values can change with species richness, indicating that trait values can change due to the biotic context. Secondly, we show that early tree growth is more strongly affected by traits than by species richness. Finally, our data show that growth–trait relationships can change with species richness and with forest age. Trait effects on growth are more pronounced at higher richness levels, indicating that measuring traits on individual trees across richness levels can improve growth predictions and inference of BEF relationships that are shaped by functional traits.4. Synthesis. This study shows that functional trait values and their effect on individual tree growth depend on species richness. Our data support the notion that to elucidate how functional traits shape biodiversity–ecosystem functioning relationships, an important step is to consider the biotic context of individual trees within a community. We have made an initial step by analysing how functional traits affect individual‐tree growth in a diversity‐dependent manner and future research should continue by elucidating the role of traits on tree–tree interactions across diversity levels. 1. Understanding how species diversity affects plant performance is a central question in biodiversity–ecosystem functioning (BEF) research. At the community level, functional trait means and trait dissimilarities have been used to explain biodiversity effects, but with mixed success. To disentangle how functional traits explain community growth and underpin biodiversity–ecosystem functioning relationships, we should elucidate how plant traits affect individual growth across species richness levels, because the role of functional traits on growth depends on the ecological context of the individual.2. We addressed this topic by using detailed data of twelve functional traits and annual growth for 529 individual trees of 31 species at five species‐richness levels in a large forest biodiversity experiment in south‐east China from 4 to 9 years after planting.3. Our analyses show first that individual trait values can change with species richness, indicating that trait values can change due to the biotic context. Secondly, we show that early tree growth is more strongly affected by traits than by species richness. Finally, our data show that growth–trait relationships can change with species richness and with forest age. Trait effects on growth are more pronounced at higher richness levels, indicating that measuring traits on individual trees across richness levels can improve growth predictions and inference of BEF relationships that are shaped by functional traits.4. Synthesis. This study shows that functional trait values and their effect on individual tree growth depend on species richness. Our data support the notion that to elucidate how functional traits shape biodiversity–ecosystem functioning relationships, an important step is to consider the biotic context of individual trees within a community. We have made an initial step by analysing how functional traits affect individual‐tree growth in a diversity‐dependent manner and future research should continue by elucidating the role of traits on tree–tree interactions across diversity levels.

Local neighbourhood interactions are considered a main driver for biodiversity–productivity relationships in forests. Yet, the structural responses of individual trees in species mixtures and their relation to crown complementarity remain poorly understood. Using a large-scale forest experiment, we studied the impact of local tree species richness and structural variability on above-ground wood volume allocation patterns and crown morphology. We applied terrestrial laser scanning to capture the three-dimensional structure of trees and their temporal dynamics. We found that crown complementarity and crown plasticity increased with species richness. Trees growing in species-rich neighbourhoods showed enhanced aboveground wood volume both in trunks and branches. Over time, neighbourhood diversity induced shifts in wood volume allocation in favour of branches, in particular for morphologically flexible species. Our results demonstrate that diversity-mediated shifts in allocation pattern and crown morphology are a fundamental mechanism for crown complementarity and may be an important driver of overyielding. publishedVersion

Light-related interactions can increase productivity in tree-species mixtures compared with monocultures due to higher stand-level absorption of photosynthetically active radiation (APAR) or light-use efficiency (LUE). However, the effects of different light-related interactions, and their relative importance, have rarely been quantified. Here, measurements of vertical leaf-area distributions, tree sizes, and stand density were combined with a tree-level light model (Maestra) to examine how crown architecture and vertical or horizontal canopy structure influence the APAR of 16 monocultures and eight different two-species mixtures with 16 different species in a Chinese subtropical tree diversity experiment. A higher proportion of crown leaf area occurred in the upper crowns of species with higher specific leaf areas. Tree-level APAR depended largely on tree leaf area and also, but to a lesser extent, on relative height (i.e., tree dominance) and leaf-area index (LAI). Stand-level APAR depended on LAI and canopy volume, but not on the vertical stratification or canopy leaf-area density. The mixing effects, in terms of relative differences between mixtures and monocultures, on stand-level APAR were correlated with the mixing effects on basal area growth, indicating that light-related interactions may have been responsible for part of the mixing effects on basal area growth. While species identity influences the vertical distributions of leaf area within tree crowns, this can have a relatively small effect on tree and stand APAR compared with the size and vertical positioning of the crowns, or the LAI and canopy volume.

AimsGrowth rates of plants are driven by factors that influence the amount of resources captured and the efficiency of resource use. In trees, the amount of light captured and the efficiency of light use strongly depends on crown characteristics and leaf traits. Although theory predicts that both crown and leaf traits affect tree growth, few studies have yet to integrate these two types of traits to explain species-specific growth rates. Using 37 broad-leaved tree species of subtropical forests in SE China, we investigated how interspecific differences in wood volume growth rates were affected by crown and leaf traits. We tested the hypotheses that (i) larger crown dimensions promote growth rates, (ii) species-specific growth rates are positively related to leaf stomatal conductance, leaf water potential and leaf chemical components, and negatively related to leaf C/N and leaf toughness and (iii) the two sets of traits better explain growth rates in combination than either alone.MethodsOur study was conducted in a large-scale forest Biodiversity and Ecosystem Functioning experiment in China (BEF-China), located in a mountainous region in Jiangxi Province. We related 17 functional traits (two crown dimension and three crown structure traits; six physiological and six morphological leaf traits) to the mean annual growth rate of wood volume of young trees of the studied species. Interrelationships between crown and leaf traits were analyzed using principal component analysis. Simple linear regression analysis was used to test the effect of each trait separately. We used multiple regression analysis to establish the relationship of growth rate to each set of traits (crown traits, physiological and morphological leaf traits) and to the combination of all types of traits. The coefficients of determination (R2adj) of the best multiple regression models were compared to determine the relative explanatory power of crown and leaf traits and a combination of both.Important FindingsThe species-specific growth rates were not related to any of the single crown traits, but were related positively to leaf stomatal conductance and leaf water potential individually, and negatively to leaf toughness, with approximately 13% variance explained by each of the traits. Combinations of different crown traits did not significantly explain the species-specific growth rates, whereas combinations of either physiological or morphological leaf traits explained 24% and 31%, respectively. A combination of both crown and leaf traits explained 42% of variance in species-specific growth rates. We concluded that sets of traits related to carbon assimilation at the leaf-level and to overall amount of leaves exposed at the crown-level jointly explained species-specific growth rates better than either set of traits alone.

Aims Although the net biodiversity effect (NE) can be statistically partitioned into complementarity and selection effects (CE and SE), there are different underlying mechanisms that can cause a certain partitioning. Our objective was to assess the role of resource partitioning and species interactions as two important mechanisms that can bring about CEs by interspecific and intraspecific trait variation. Methods We measured tree height of 2493 living individuals in 57 plots and specific root length (SRL) on first-order roots of 368 of these individuals across different species richness levels (1, 2, 4, 8 species) in a large-scale forest biodiversity and ecosystem functioning experiment in subtropical China (BEF-China) established in 2009. We describe the effects of resource partitioning between species by a fixed component of interspecific functional diversity (RaoQ) and further effects of species interactions by variable components of interspecific and intraspecific functional diversity (community weighted trait similarity and trait dissimilarity, CWS and CWD). Finally, we examined the relationships between biodiversity effects on stand-level tree height and functional diversity (RaoQ, CWS and CWD) in SRL using linear regression and assessed the relative importance of these three components of functional diversity in explaining the diversity effects. Important Findings Our results show that species richness significantly affected SRL in five and tree height in ten out of 16 species. A positive NE was generally brought about by a positive CE on stand-level tree height and related to high values of RaoQ and CWS in SRL. A positive CE was related to high values of all three components of root functional diversity (RaoQ, CWS and CWD). Our study suggests that both resource partitioning and species interactions are the underlying mechanisms of biodiversity effects on stand-level tree growth in subtropical forest.

Forest canopies and tree crown structures are of high ecological importance.Measuring canopies and crowns by direct inventory methods is time-consuming and of limited accuracy. High-resolution inventory tools, in particular terrestrial laser scanning (TLS), is able to overcome these limitations and obtain three-dimensional (3D) structural information about the canopy with a very high level of detail. The main objective of this study was to introduce a novel method to analyze spatiotemporal dynamics in canopy occupancy at the individual tree and local neighborhood level using high-resolution 3D TLS data. For the analyses, a voxel grid approach was applied. The tree crowns were modeled through the combination of two approaches: the encasement of all crown points with a 3D α-shape, which was then converted into a voxel grid, and the direct voxelization of the crown points. We show that canopy occupancy at individual tree level can be quantified as the crown volume occupied only by the respective tree or shared with neighboring trees. At the local neighborhood level, our method enables the precise determination of the extent of canopy space filling, the identification of tree–tree interactions, and the analysis of complementary space use. Using multitemporal TLS data recordings, this method allows the precise detection and quantification of changes in canopy occupancy through time. The method is applicable to a wide range of investigations in forest ecology research, including the study of tree diversity effects on forest productivityor growing space analyses for optimal tree growth. Due to the high accuracy of this novel method, it facilitates the precise analyses even of highly plastic individual tree crowns and, thus, the realistic representation of forest canopies. Moreover, our voxel grid framework is flexible enough to allow for the inclusion of further biotic and abiotic variables relevant to complex analyses of forest canopy dynamics. Forest canopies and tree crown structures are of high ecological importance. Measuring canopies and crowns by direct inventory methods is time‐consuming and of limited accuracy. High‐resolution inventory tools, in particular terrestrial laser scanning (TLS), is able to overcome these limitations and obtain three‐dimensional (3D) structural information about the canopy with a very high level of detail. The main objective of this study was to introduce a novel method to analyze spatiotemporal dynamics in canopy occupancy at the individual tree and local neighborhood level using high‐resolution 3D TLS data. For the analyses, a voxel grid approach was applied. The tree crowns were modeled through the combination of two approaches: the encasement of all crown points with a 3D α‐shape, which was then converted into a voxel grid, and the direct voxelization of the crown points. We show that canopy occupancy at individual tree level can be quantified as the crown volume occupied only by the respective tree or shared with neighboring trees. At the local neighborhood level, our method enables the precise determination of the extent of canopy space filling, the identification of tree–tree interactions, and the analysis of complementary space use. Using multitemporal TLS data recordings, this method allows the precise detection and quantification of changes in canopy occupancy through time. The method is applicable to a wide range of investigations in forest ecology research, including the study of tree diversity effects on forest productivity or growing space analyses for optimal tree growth. Due to the high accuracy of this novel method, it facilitates the precise analyses even of highly plastic individual tree crowns and, thus, the realistic representation of forest canopies. Moreover, our voxel grid framework is flexible enough to allow for the inclusion of further biotic and abiotic variables relevant to complex analyses of forest canopy dynamics.

While functional diversity (FD) has been shown to be positively related to a number of ecosystem functions including biomass production, it may have a much less pronounced effect than that of environmental factors or species-specific properties. Leaf and wood traits can be considered particularly relevant to tree growth, as they reflect a trade-off between resources invested into growth and persistence. Our study focussed on the degree to which early forest growth was driven by FD, the environment (11 variables characterizing abiotic habitat conditions), and community-weighted mean (CWM) values of species traits in the context of a large-scale tree diversity experiment (BEF-China). Growth rates of trees with respect to crown diameter were aggregated across 231 plots (hosting between one and 23 tree species) and related to environmental variables, FD, and CWM, the latter two of which were based on 41 plant functional traits. The effects of each of the three predictor groups were analyzed separately by mixed model optimization and jointly by variance partitioning. Numerous single traits predicted plot-level tree growth, both in the models based on CWMs and FD, but none of the environmental variables was able to predict tree growth. In the best models, environment and FD explained only 4 and 31% of variation in crown growth rates, respectively, while CWM trait values explained 42%. In total, the best models accounted for 51% of crown growth. The marginal role of the selected environmental variables was unexpected, given the high topographic heterogeneity and large size of the experiment, as was the significant impact of FD, demonstrating that positive diversity effects already occur during the early stages in tree plantations. While functional diversity (FD) has been shown to be positively related to a number of ecosystem functions including biomass production, it may have a much less pronounced effect than that of environmental factors or species‐specific properties. Leaf and wood traits can be considered particularly relevant to tree growth, as they reflect a trade‐off between resources invested into growth and persistence. Our study focussed on the degree to which early forest growth was driven by FD, the environment (11 variables characterizing abiotic habitat conditions), and community‐weighted mean (CWM) values of species traits in the context of a large‐scale tree diversity experiment (BEF‐China). Growth rates of trees with respect to crown diameter were aggregated across 231 plots (hosting between one and 23 tree species) and related to environmental variables, FD, and CWM, the latter two of which were based on 41 plant functional traits. The effects of each of the three predictor groups were analyzed separately by mixed model optimization and jointly by variance partitioning. Numerous single traits predicted plot‐level tree growth, both in the models based on CWMs and FD, but none of the environmental variables was able to predict tree growth. In the best models, environment and FD explained only 4 and 31% of variation in crown growth rates, respectively, while CWM trait values explained 42%. In total, the best models accounted for 51% of crown growth. The marginal role of the selected environmental variables was unexpected, given the high topographic heterogeneity and large size of the experiment, as was the significant impact of FD, demonstrating that positive diversity effects already occur during the early stages in tree plantations.

Despite the importance of herbivory for the structure and functioning of species-rich forests, little is known about how herbivory is affected by tree species richness, and more specifically by random vs. non-random species loss.We assessed herbivore damage and its effects on tree growth in the early stage of a large-scale forest biodiversity experiment in subtropical China that features random and non-random extinction scenarios of tree mixtures numbering between one and 24 species. In contrast to random species loss, the non-random extinction scenarios were based on the tree species' local rarity and specific leaf area – traits that may strongly influence the way herbivory is affected by plant species richness.Herbivory increased with tree species richness across all scenarios and was unaffected by the different species compositions in the random and non-random extinction scenarios. Whereas tree growth rates were positively related to herbivory on plots with smaller trees, growth rates significantly declined with increasing herbivory on plots with larger trees. Our results suggest that the effects of herbivory on growth rates increase from monocultures to the most species-rich plant communities and that negative effects with increasing tree species richness become more pronounced with time as trees grow larger.Synthesis. Our results indicate that key trophic interactions can be quick to become established in forest plantations (i.e. already 2.5 years after tree planting). Stronger herbivory effects on tree growth with increasing tree species richness suggest a potentially important role of herbivory in regulating ecosystem functions and the structural development of species-rich forests from the very start of secondary forest succession. The lack of significant differences between the extinction scenarios, however, contrasts with findings from natural forests of higher successional age, where rarity had negative effects on herbivory. This indicates that the effects of non-random species loss could change with forest succession.

Biodiversity-ecosystem functioning (BEF) research has extended its scope from communities that are short-lived or reshape their structure annually to structurally complex forest ecosystems. The establishment of tree diversity experiments poses specific methodological challenges for assessing the multiple functions provided by forest ecosystems. In particular, methodological inconsistencies and nonstandardized protocols impede the analysis of multifunctionality within, and comparability across the increasing number of tree diversity experiments. By providing an overview on key methods currently applied in one of the largest forest biodiversity experiments, we show how methods differing in scale and simplicity can be combined to retrieve consistent data allowing novel insights into forest ecosystem functioning. Furthermore, we discuss and develop recommendations for the integration and transferability of diverse methodical approaches to present and future forest biodiversity experiments. We identified four principles that should guide basic decisions concerning method selection for tree diversity experiments and forest BEF research: (1) method selection should be directed toward maximizing data density to increase the number of measured variables in each plot. (2) Methods should cover all relevant scales of the experiment to consider scale dependencies of biodiversity effects. (3) The same variable should be evaluated with the same method across space and time for adequate larger-scale and longer-time data analysis and to reduce errors due to changing measurement protocols. (4) Standardized, practical and rapid methods for assessing biodiversity and ecosystem functions should be promoted to increase comparability among forest BEF experiments. We demonstrate that currently available methods provide us with a sophisticated toolbox to improve a synergistic understanding of forest multifunctionality. However, these methods require further adjustment to the specific requirements of structurally complex and long-lived forest ecosystems. By applying methods connecting relevant scales, trophic levels, and above- and belowground ecosystem compartments, knowledge gain from large tree diversity experiments can be optimized. Biodiversity–ecosystem functioning (BEF) research has extended its scope from communities that are short‐lived or reshape their structure annually to structurally complex forest ecosystems. The establishment of tree diversity experiments poses specific methodological challenges for assessing the multiple functions provided by forest ecosystems. In particular, methodological inconsistencies and nonstandardized protocols impede the analysis of multifunctionality within, and comparability across the increasing number of tree diversity experiments. By providing an overview on key methods currently applied in one of the largest forest biodiversity experiments, we show how methods differing in scale and simplicity can be combined to retrieve consistent data allowing novel insights into forest ecosystem functioning. Furthermore, we discuss and develop recommendations for the integration and transferability of diverse methodical approaches to present and future forest biodiversity experiments. We identified four principles that should guide basic decisions concerning method selection for tree diversity experiments and forest BEF research: (1) method selection should be directed toward maximizing data density to increase the number of measured variables in each plot. (2) Methods should cover all relevant scales of the experiment to consider scale dependencies of biodiversity effects. (3) The same variable should be evaluated with the same method across space and time for adequate larger‐scale and longer‐time data analysis and to reduce errors due to changing measurement protocols. (4) Standardized, practical and rapid methods for assessing biodiversity and ecosystem functions should be promoted to increase comparability among forest BEF experiments. We demonstrate that currently available methods provide us with a sophisticated toolbox to improve a synergistic understanding of forest multifunctionality. However, these methods require further adjustment to the specific requirements of structurally complex and long‐lived forest ecosystems. By applying methods connecting relevant scales, trophic levels, and above‐ and belowground ecosystem compartments, knowledge gain from large tree diversity experiments can be optimized.

Studies on tree communities have demonstrated that species diversity can enhance forest productivity, but the driving mechanisms at the local neighbourhood level remain poorly understood. Here, we use data from a large-scale biodiversity experiment with 24 subtropical tree species to show that neighbourhood tree species richness generally promotes individual tree productivity. We found that the underlying mechanisms depend on a focal tree's functional traits: For species with a conservative resource-use strategy diversity effects were brought about by facilitation, and for species with acquisitive traits by competitive reduction. Moreover, positive diversity effects were strongest under low competition intensity (quantified as the total basal area of neighbours) for acquisitive species, and under high competition intensity for conservative species. Our findings demonstrate that net biodiversity effects in tree communities can vary over small spatial scales, emphasising the need to consider variation in local neighbourhood interactions to better understand effects at the community level.

We investigated long-term responses (since 1850) of Fagus sylvatica (Luxembourg; central Europe) to shifts in temperature, precipitation, and nitrogen deposition by analyzing diameter at breast height (DBH) increment, basal area increment (BAI), and tree-ring stable isotopes (δ13C, δ15N). We compared stands on soils with contrasting water supply (Regosols and Cambisols with an available water capacity of ca. 40 and 170 mm, respectively) and of two different age classes (ca. 60 vs. 200 years). All stands showed a peak in DBH increment in the decade 1978–1987, but a decline in increment growth in subsequent decades. In addition, BAI declined in mature stands in the last two decades. Decreasing increment rates were attributable to an increasing drought limitation of stands, mainly induced by increasing temperatures in the last two decades. Contrary to our expectations, stands on Cambisols showed a similar susceptibility to shifts in temperature and precipitation as stands on Regosols, suggesting a strong adaptation of the respective ecotypes grown at dryer sites. This result was in line with long-term trends for tree-ring δ13C signatures, which did not differ significantly between stands on Cambisols and Regosols. Climate impacts on tree-ring δ15N signatures were low. High spring precipitation and temperatures caused increasing and decreasing δ15N values, respectively, but only in mature stands on Cambisols. Stands on Regosols tended to have lower δ15N values than stands on Cambisols. Decreasing δ15N values in recent decades suggest an increasing impact of allochthonous N loads with isotopically lighter N. We investigated long-term responses (since 1850) of Fagus sylvatica (Luxembourg; central Europe) to shifts in temperature, precipitation, and nitrogen deposition by analyzing diameter at breast height (DBH) increment, basal area increment (BAI), and tree-ring stable isotopes (δ13C, δ15N). We compared stands on soils with contrasting water supply (Regosols and Cambisols with an available water capacity of ca. 40 and 170 mm, respectively) and of two different age classes (ca. 60 vs. 200 years). All stands showed a peak in DBH increment in the decade 1978–1987, but a decline in increment growth in subsequent decades. In addition, BAI declined in mature stands in the last two decades. Decreasing increment rates were attributable to an increasing drought limitation of stands, mainly induced by increasing temperatures in the last two decades. Contrary to our expectations, stands on Cambisols showed a similar susceptibility to shifts in temperature and precipitation as stands on Regosols, suggesting a strong adaptation of the respective ecotypes grown at dryer sites. This result was in line with long-term trends for tree-ring δ13C signatures, which did not differ significantly between stands on Cambisols and Regosols. Climate impacts on tree-ring δ15N signatures were low. High spring precipitation and temperatures caused increasing and decreasing δ15N values, respectively, but only in mature stands on Cambisols. Stands on Regosols tended to have lower δ15N values than stands on Cambisols. Decreasing δ15N values in recent decades suggest an increasing impact of allochthonous N loads with isotopically lighter N.

AimsThe aim of our research was to understand small-scale effects of topography and soil fertility on tree growth in a forest biodiversity and ecosystem functioning (BEF) experiment in subtropical SE China.MethodsGeomorphometric terrain analyses were carried out at a spatial resolution of 5×5 m. Soil samples of different depth increments and data on tree height were collected from a total of 566 plots (667 m2 each). The soils were analyzed for carbon (soil organic carbon [SOC]), nitrogen, acidity, cation exchange capacity (CEC), exchangeable cations and base saturation as soil fertility attributes. All plots were classified into geomorphological units. Analyses of variance and linear regressions were applied to all terrain, soil fertility and tree growth attributes.Important FindingsIn general, young and shallow soils and relatively small differences in stable soil properties suggest that soil erosion has truncated the soils to a large extent over the whole area of the experiment. This explains the concurrently increasing CEC and SOC stocks downslope, in hollows and in valleys. However, colluvial, carbon-rich sediments are missing widely due to the convexity of the footslopes caused by uplift and removal of eroded sediments by adjacent waterways. The results showed that soil fertility is mainly influenced by topography. Monte–Carlo flow accumulation (MCCA), curvature, slope and aspect significantly affected soil fertility. Furthermore, soil fertility was affected by the different geomorphological positions on the experimental sites with ridge and spur positions showing lower exchangeable base cation contents, especially potassium (K), due to leaching. This geomorphological effect of soil fertility is most pronounced in the topsoil and decreases when considering the subsoil down to 50cm depth. Few soil fertility attributes affect tree height after 1–2 years of growth, among which C stocks proved to be most important while pHKCl and CEC only played minor roles. Nevertheless, soil acidity and a high proportion of Al on the exchange complex affected tree height even after only 1–2 years growth. Hence, our study showed that forest nutrition is coupled to a recycling of litter nutrients, and does not only depend on subsequent supply of nutrients from the mineral soil. Besides soil fertility, topography affected tree height. We found that especially MCCA as indicator of water availability affected tree growth at small-scale, as well as aspect. Overall, our synthesis on the interrelation between fertility, topography and tree growth in a subtropical forest ecosystem in SE China showed that topographic heterogeneity lead to ecological gradients across geomorphological positions. In this respect, small-scale soil–plant interactions in a young forest can serve as a driver for the future development of vegetation and biodiversity control on soil fertility. In addition, it shows that terrain attributes should be accounted for in ecological research.

tAnthropogenic deposition of reactive nitrogen (N) has increased during the 20th century, and is con-sidered an important driver of shifts in ecosystem functions and biodiversity loss. The objective of thepresent study was to identify those ecosystem functions that best evidence a target ecosystem’s sensitiv-ity to N deposition, taking coastal heathlands as an example. We conducted a three-year field experimentin heathlands of the island Fehmarn (Baltic Sea, North Germany), which currently are subject to a back-ground deposition of 9 kg N ha−1yr−1. We experimentally applied six levels of N fertilisation (applicationof 0, 2.5, 5, 10, 20, and 50 kg N ha−1yr−1), and quantified the growth responses of different plant speciesof different life forms (dwarf shrubs, graminoids, bryophytes, lichens) as well as shifts in the C:N ratios ofplant tissue and humus horizons. For an applicability of the experimental findings (in terms of heathlandmanagement and critical load assessment) fertilisation effects on response variables were visualised bycalculating the treatment ‘effect sizes’. The current year’s shoot increment of the dominant dwarf shrubCalluna vulgaris proved to be the most sensitive indicator to N fertilisation. Shoot increment significantlyresponded to additions of ≥ 5 kg N ha−1yr−1already in the first year, whereas flower formation of Callunavulgaris increased only in the high-N treatments. Similarly, tissue C:N ratios of vascular plants (Callunavulgaris and the graminoids Carex arenaria and Festuca ovina agg.) only decreased in the highest N treat-ments (50 and 20 kg N ha−1yr−1, respectively). In contrast, tissue C:N ratios of cryptogams respondedmore quickly and sensitively than vascular plants. For example, Cladonia spp. tissue C:N ratios respondedto N additions ≥ 5 kg N ha−1yr−1in the second study year. After three years we observed an increase incover of graminoids and a corresponding decrease of cryptogams at N fertilisation rates of ≥ 10 kg N ha−1yr−1. Soil C:N ratios proved to be an inappropriate indicator for N fertilisation at least within our three-year study period. Although current critical N loads for heathlands (10−20 kg N ha−1yr−1) were confirmedin our experiment, the immediate and highly sensitive response of the current year’s shoots of Callunavulgaris suggests that at least some ecosystem functions (e.g. dwarf shrub growth) also might respond tolow (i.e. < 10 kg N ha−1yr−1) but chronic inputs of N. Anthropogenic deposition of reactive nitrogen (N) has increased during the 20th century, and is considered an important driver of shifts in ecosystem functions and biodiversity loss. The objective of the present study was to identify those ecosystem functions that best evidence a target ecosystem’s sensitivity to N deposition, taking coastal heathlands as an example. We conducted a three-year field experiment in heathlands of the island Fehmarn (Baltic Sea, North Germany), which currently are subject to a background deposition of 9 kg N ha−1 yr−1. We experimentally applied six levels of N fertilisation (application of 0, 2.5, 5, 10, 20, and 50 kg N ha−1 yr−1), and quantified the growth responses of different plant species of different life forms (dwarf shrubs, graminoids, bryophytes, lichens) as well as shifts in the C:N ratios of plant tissue and humus horizons. For an applicability of the experimental findings (in terms of heathland management and critical load assessment) fertilisation effects on response variables were visualised by calculating the treatment ‘effect sizes’. The current year’s shoot increment of the dominant dwarf shrub Calluna vulgaris proved to be the most sensitive indicator to N fertilisation. Shoot increment significantly responded to additions of ≥ 5 kg N ha−1 yr−1 already in the first year, whereas flower formation of Calluna vulgaris increased only in the high-N treatments. Similarly, tissue C:N ratios of vascular plants (Calluna vulgaris and the graminoids Carex arenaria and Festuca ovina agg.) only decreased in the highest N treatments (50 and 20 kg N ha−1 yr−1, respectively). In contrast, tissue C:N ratios of cryptogams responded more quickly and sensitively than vascular plants. For example, Cladonia spp. tissue C:N ratios responded to N additions ≥ 5 kg N ha−1 yr−1 in the second study year. After three years we observed an increase in cover of graminoids and a corresponding decrease of cryptogams at N fertilisation rates of ≥ 10 kg N ha−1 yr−1. Soil C:N ratios proved to be an inappropriate indicator for N fertilisation at least within our three-year study period. Although current critical N loads for heathlands (10−20 kg N ha−1 yr−1) were confirmed in our experiment, the immediate and highly sensitive response of the current year’s shoots of Calluna vulgaris suggests that at least some ecosystem functions (e.g. dwarf shrub growth) also might respond to low (i.e. < 10 kg N ha−1 yr−1) but chronic inputs of N.

The specialized fauna of peat bogs declines strongly both in species numbers and population density and becomes fragmented because of anthropogenic land use changes. We investigated 15 populations of the stenotopic ground beetle species Agonum ericeti in south Sweden to address the question of reduced genetic variability in populations inhabiting smaller habitat patches. Our results reveal a generally low differentiation and a significant positive relationship between habitat size and allelic richness The specialized fauna of peat bogs declines strongly both in species numbers and population density and becomes fragmented because of anthropogenic land use changes. We investigated 15 populations of the stenotopic ground beetle species Agonum ericeti in south Sweden to address the question of reduced genetic variability in populations inhabiting smaller habitat patches. Our results reveal a generally low differentiation and a significant positive relationship between habitat size and allelic richness.

Summary 1. Insect herbivory can strongly affect ecosystem processes, and its relationship with plant diversity is a central topic in biodiversity–functioning research. However, very little is known about this relationship from complex ecosystems dominated by long-lived individuals, such as forests, especially over gradients of high plant diversity. 2. We analysed insect herbivory on saplings of 10 tree and shrub species across 27 forest stands differing in age and tree species richness in an extraordinarily diverse subtropical forest ecosystem in China. We tested whether plant species richness significantly influences folivory in these highly diverse forests or whether other factors play a more important role at such high levels of phytodiversity. 3. Leaf damage was assessed on 58 297 leaves of 1284 saplings at the end of the rainy season in 2008, together with structural and abiotic stand characteristics. 4. Species-specific mean damage of leaf area ranged from 3% to 16%. Herbivory increased with plant species richness even after accounting for potentially confounding effects of stand characteristics, of which stand age-related aspects most clearly covaried with herbivory. Intraspecific density dependence or other abiotic factors did not significantly influence overall herbivory across forest stands. 5. Synthesis. The positive herbivory–plant diversity relationship indicates that effects related to hypotheses of resource concentration, according to which a reduction in damage by specialized herbivores might be expected as host plant concentration decreases with increasing plant diversity, do not seem to be major determinants for overall herbivory levels in our phytodiverse subtropical forest ecosystem. We discuss the potential role of host specificity of dominant herbivores, which are often expected to show a high degree of specialization in many (sub)tropical forests. In the forest system we studied, a much higher impact of polyphagous species than traditionally assumed might explain the observed patterns, as these species can profit from a broad dietary mix provided by high plant diversity. Further testing is needed to experimentally verify this assumption.

In the face of ongoing atmospheric nutrient loads the employment of management measures to remove nutrients from heathland ecosystems has increased in importance. The present study is the first to analyse whether Calluna vulgaris is a suitable bio-monitor of management-mediated nutrient pools in heathland ecosystems. If Calluna vulgaris proves to be an appropriate indicator, its bio-indicative usage may prove to be a helpful tool for an assessment of management success in heathland ecosystems. In the Luneburger Heide nature reserve (NW Germany) we analysed the impacts of grazing, mowing, prescribed burning, choppering and sod-cutting on the nutritional status of Calluna vulgaris by measuring nutrient contents (N, P, Ca, Mg, K) of current year's shoots I and 5 years after application of management measures. Results were related to management-induced nutrient flows and nutrient pools at the focal heath sites. Our results indicate that the less the physical environment of a heath site was affected by management measures the better the nutrient contents of current year's shoots of Calluna vulgaris mirrored changes in nutrient pools. For low-intensity measures (i.e. grazing, mowing, prescribed burning), shoot nutrient contents were a suitable indicator for changes in nutrient pools, particularly for nutrients with conservative cycles such as P. At grazed and mown sites high output rates of P caused by these measures were well reflected by decreased shoot P content. At burned sites, Calluna vulgaris proved to be a good indicator of changes in nutrient pools of the organic layer, mainly attributable to the deposition of nutrients with ash. In contrast, at sites subjected to high-intensity measures, shoot nutrient contents did not reflect management-mediated shifts in nutrient pools, despite the high nutrient losses caused by choppering and sod-cutting. At these sites, shoot nutrient contents mirrored only the effects of altered mineralisation rates attributable to changes in the physical environment following high-intensity measures. As plant growth and competition in heathlands is considered to be controlled by N or P, shoot N:P ratios are recommended as a tool to indicate whether plant growth tends to be limited by N, by P or by N and P. This, in turn, allows for an assessment of long-term effects of both atmospheric nutrient loads and management-mediated shifts in N and P pools at a focal heath site. (C) 2008 Elsevier Ltd. All rights reserved. In the face of ongoing atmospheric nutrient loads the employment of management measures to remove nutrients from heathland ecosystems has increased in importance. The present study is the first to analyse whether Calluna vulgaris is a suitable bio-monitor of management-mediated nutrient pools in heathland ecosystems. If Calluna vulgaris proves to be an appropriate indicator, its bio-indicative usage may prove to be a helpful tool for an assessment of management success in heathland ecosystems. In the Lüneburger Heide nature reserve (NW Germany) we analysed the impacts of grazing, mowing, prescribed burning, choppering and sod-cutting on the nutritional status of Calluna vulgaris by measuring nutrient contents (N, P, Ca, Mg, K) of current year's shoots 1 and 5 years after application of management measures. Results were related to management-induced nutrient flows and nutrient pools at the focal heath sites. Our results indicate that the less the physical environment of a heath site was affected by management measures the better the nutrient contents of current year's shoots of Calluna vulgaris mirrored changes in nutrient pools. For low-intensity measures (i.e. grazing, mowing, prescribed burning), shoot nutrient contents were a suitable indicator for changes in nutrient pools, particularly for nutrients with conservative cycles such as P. At grazed and mown sites high output rates of P caused by these measures were well reflected by decreased shoot P content. At burned sites, Calluna vulgaris proved to be a good indicator of changes in nutrient pools of the organic layer, mainly attributable to the deposition of nutrients with ash. In contrast, at sites subjected to high-intensity measures, shoot nutrient contents did not reflect management-mediated shifts in nutrient pools, despite the high nutrient losses caused by choppering and sod-cutting. At these sites, shoot nutrient contents mirrored only the effects of altered mineralisation rates attributable to changes in the physical environment following high-intensity measures. As plant growth and competition in heathlands is considered to be controlled by N or P, shoot N:P ratios are recommended as a tool to indicate whether plant growth tends to be limited by N, by P or by N and P. This, in turn, allows for an assessment of long-term effects of both atmospheric nutrient loads and management-mediated shifts in N and P pools at a focal heath site.

Sclerophyllous oak forests are an important ecosystem type of the natural vegetation in the Mediterranean region. As a part of the mosaic-like landscape, old-growth oak forests, in particular, provide a wide range of ecosystem functions and services. However, due to the general scarcity of older forests in the Mediterranean region, studies on structure and dynamics of oak forests are mostly restricted to younger stands. The aim of this study was to analyze stand structure, dead wood, and natural regeneration of an old Quercus calliprinos forest in northern Israel (nature reserve Ya'ar Bar'am) and to compare the results with those reported from other evergreen Mediterranean oak forests. The mean density of trees was 735 ha-1, and the mean basal area amounted to 25.0 m2 ha-1. The mean tree height was 6.6 m, and the maximum tree height was 13.2 m. The mean dead wood volume was 11 m3 ha-1. The sapling density ranged from 1,050 to 6,250 ha-1. We conclude that Bar'am Forest is of great value for conservation bi...

In the present study we analyzed the combined effects of management (grazing, mowing, prescribed burning, sod-cutting) and atmospheric deposition on N and P budgets of heathland ecosystems (Lüneburger Heide nature reserve; N Germany). We hypothesize that management measures such as grazing and mowing can accelerate a deposition-induced imbalance of N and P pools as a result of a disproportionally high output of P. We analyzed management and deposition affected input-output flows of N and P and related them to changes in the nutritional status of Calluna vulgaris 5 years after treatment application. We found that grazing and mowing caused the highest net loss of P due to high P concentrations in the aboveground biomass. In contrast, prescribed burning only slightly affected P pools, as P remained in the system due to ash deposition. Management-mediated effects on N and P pools were mirrored in the nutritional status of Calluna vulgaris: at the grazed and mown sites, the P content of current season's shoots significantly decreased within 5 years after treatments, whereas the N content remained unchanged. We conclude that grazing and mowing can accelerate declining availability of P and, thus, accelerate a deposition-induced shift from N- to P-limited plant growth in the medium term. In the face of ongoing atmospheric N loads management schemes need to combine high- and low-intensity measures to maintain both a diverse structure and balanced nutrient budgets in the long term. © 2009 Springer Science+Business Media, LLC. In the present study we analyzed the combined effects of management (grazing, mowing, prescribed burning, sod-cutting) and atmospheric deposition on N and P budgets of heathland ecosystems (Lüneburger Heide nature reserve; N Germany). We hypothesize that management measures such as grazing and mowing can accelerate a deposition-induced imbalance of N and P pools as a result of a disproportionally high output of P. We analyzed management and deposition affected inputoutput flows of N and P and related them to changes in the nutritional status of Calluna vulgaris 5 years after treatment application. We found that grazing and mowing caused the highest net loss of P due to high P concentrations in the aboveground biomass. In contrast, prescribed burning only slightly affected P pools, as P remained in the system due to ash deposition. Management-mediated effects on N and P pools were mirrored in the nutritional status of Calluna vulgaris: at the grazed and mown sites, the P content of current season’s shoots significantly decreased within 5 years after treatments, whereas the N content remained unchanged. We conclude that grazing and mowing can accelerate declining availability of P and, thus, accelerate a deposition-induced shift from N- to P-limited plant growth in the medium term. In the face of ongoing atmospheric N loads management schemes need to combine high- and low-intensity measures to maintain both a diverse structure and balanced nutrient budgets in the long term.

In this study, we report on 67 years of secondary succession in the temperate deciduous forest Dalby Söderskog, southern Sweden. In 2002 vegetation analyses were conducted in 74 systematically distributed plots and the results compared with similar investigations during the years 1935, 1969 and 1976. Additionally, a floristic survey of the forest was made and compared with inventories of the years 1925, 1935 and 1970. From 1970 to 2002, overall species richness of the forest continued to decline, but at a lower rate than before. In 2002, for the first time, shade-tolerant forest species constituted the largest group. In the upper tree layer Fraxinus excelsior became the most important species, at the expense of Quercus robur and Ulmus glabra. In the herb layer, no changes in total or mean species richness were exhibited at plot scale between 1976 and 2002. In contrast to the lack of quantitative changes in species richness, we observed major changes in species composition at the plot level. In particular, the former dominant species Mercurialis perennis had declined markedly. After release from a historical regime of livestock grazing and irregular cuttings, the predicted steady-state community dominated by Ulmus and Mercurialis only prevailed for a few decades before unexpected external events, in this particular case Dutch elm disease and invasion by the slug Arion lusitanicus, initiated a successional change. We conclude that unexpected diseases or pests or rare climatic extremes can play a decisive role in forest dynamics and strongly interact with more gradual autogenic changes of forest structure and composition.

Climate change and atmospheric deposition of nitrogen (N) affect the biodiversity patterns and functions of ecosystems world-wide. While many single-factor studies have quantified ecosystem responses to single global change drivers, less is known about the interaction effects of these drivers on ecosystem functions. Here, we present the results of a three-year field and a two-year glasshouse experiment, in which we assessed responses of Calluna vulgaris heathlands to the single and combined effects of drought events (D) and N fertilization (D: 25% precipitation reduction in the field experiment and 20-50% soil water content reduction in the glasshouse experiment; N fertilization: 35 kg N ha-1 year-1). We examined the effects of D and N treatments on growth responses of the dominant dwarf shrub Calluna vulgaris (in terms of biomass production and allocation, tissue δ13C signatures and C:N ratios) in relation to two plant life-history stages and different 'ecotypes' (sub-Atlantic vs. subcontinental heathlands). Plant responses varied strongly with life-history stage, and the interaction of N and D showed lower effects than would be expected based on additive responses to single factors. While D treatments had no effects on Calluna in the building phase (ca. ten-year-old plants), seedlings (particularly one-year-old plants) were highly susceptible to drought. Differences in response patterns were attributable to the high shoot-root ratios typical of young Calluna plants. These ratios decreased with progressing life history as a result of increasing below-ground biomass investments. Below-ground biomass production and shoot-root ratios differed between plants from the different heathland sites. Tissue δ13C signatures decreased and C:N ratios increased with plant age as a result of decreasing evaporative demands (per unit root biomass). N fertilization increases the shoot-root ratios and thereby the drought susceptibility of Calluna plants. Synthesis. Our findings suggest that plant responses to global change are difficult to anticipate by means of single-factor studies or by focusing on a single life-history stage. This highlights the need for global change research to include multiple factors and life-history stages when assessing an ecosystem's susceptibility to shifts in environmental conditions.

The impact of forest management on the species diversity and species composition of epiphytic bryophytes and lichens in beech forests (Fagus sylvatica ) of NE Germany was analysed. The investigations were carried out in one unmanaged and in one managed forest, with 45 sample plots of 400 m2 each. In the sample plots, the presence of the epiphytes on 2–4 randomly chosen trees (100 trees per forest) and seven environmental variables were recorded. The total number of species and the mean lichen density per mantle area was higher in the unmanaged forest than in the managed forest. Regression analyses and DCA revealed the diameter at breast height (dbh) as the most important factor affecting the species diversity and species composition in both forest types. The dbh was positively correlated with the number of growth anomalies and with the bark roughness of the phorophytes and thus a good indicator for the habitat quality. Moreover, the light conditions had a significant impact on the species composition in both forests types and on the species diversity in the managed forest. In particular, epiphytes demanding shady conditions and a consistent humidity may suffer from the abrupt exposition to radiation and the lower humidity after logging. Species typical for the unmanaged forest and most rare species must be considered as stenoecious as they demand the shady and humid conditions of the forest interior and grow on old, large trees. With the aim of sustaining the diversity of epiphytic bryophytes and lichens, forest management should ensure the continuous occurrence of big trees above the target diameter. In addition, a single tree selection felling method should be applied to minimise strong changes in the microclimate. (© 2006 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim) Artenvielfalt und Artenzusammensetzung epiphytischer Moose und Flechten – ein Vergleich zwischen bewirtschafteten und unbewirtschafteten Buchenwaldern in Nordostdeutschland Der Einfluss der Bewirtschaftung auf die Artenvielfalt und Artenzusammensetzung epiphytischer Moose und Flechten in Buchenwaldern Nordostdeutschlands wurde analysiert. Die Untersuchungen erfolgten in einem unbewirtschafteten und einem benachbarten, bewirtschafteten Waldgebiet in jeweils 45 Untersuchungsflachen a 400 m2. In den Untersuchungsflachen wurden die Stetigkeit der Epiphyten an jeweils zwei bis vier zufallig ausgewahlten Tragerbaumen (100 Baume je Waldtyp) sowie sieben Umweltvariablen aufgenommen. Im Naturwald waren die Gesamtartenzahl sowie die mittlere Artendichte der Flechten bezogen auf die Mantelflache hoher als im Wirtschaftswald. Die Regressionsanalysen und die DCA wiesen den Brusthohendurchmesser (BHD) als den fur die Artenvielfalt und Artenzusammensetzung bedeutsamsten Faktor aus. Der BHD korrelierte positiv mit der Anzahl an Wuchsanomalien und der Rindenrauigkeit der Tragerbaume und ist deshalb ein guter Indikator fur die Habitatqualitat. Weiterhin zeigten die Lichtverhaltnisse einen signifikanten Einfluss auf die Artenzusammensetzung in beiden Waldtypen sowie auf die Artenvielfalt im Wirtschaftswald. Insbesondere schattentolerante und austrocknungsempfindliche Arten konnen durch die plotzliche Lichtstellung und Veranderung des Mikroklimas infolge forstlicher Eingriffe geschadigt werden. Die Arten mit deutlich hoherer Stetigkeit im Naturwald sowie die meisten seltenen Arten gelten als stenok, da sie die schattigen und luftfeuchten Bedingungen eines geschlossenen Waldinnenklimas benotigen und uberwiegend an alten, starken Baumen siedeln. Zur Erhaltung der Artenvielfalt epiphytischer Moose und Flechten sollten daher im Rahmen forstlichen Managements starke Baume uber den Zieldurchmesser hinaus erhalten werden. Weiterhin ist eine einzelstammweise Nutzung anzustreben, um Schwankungen im Mikroklima zu minimieren.

Measurements of the soil pH are often used to characterize the nutritional conditions for the vegetation at a particular site, for example in ecograms describing the ecological behaviour of a (tree) species. In spite of this use and interpretation of pH measurements, certain prerequisites must be fulfilled when pH-values are to be used as an expression of the nutritional conditions for plants. Our study in deciduous forests of northern Germany shows that in strongly acid soils (pH < 4) the correlations be-tween pH-values on the one hand and S-value, base saturation and C/N-ratios on the other hand are generally low, particularly for measurements in the upper mineral horizon. By contrast, in moderately acid to neutral soils (pH 4–7), pH-values could be employed as a useful indicator of the nutritional conditions for the vegetation. The main reason for the observed discrepancy in the indicator quality of pH measurements is the presence of distinct differences in the cation exchange capacity (CEC) of the soils, which are often not taken into account when interpreting pH measurements. CEC-values close to zero may be associated with either low or comparatively high pH-values, depending on whether or not the few available cation exchange sites are occupied by protons or bases, respectively. In general, pH – S-value/base saturation/CN-ratio relationships are stronger in soil samples taken in the main (fine and coarse) root horizon than in those from the upper mineral horizon. Consequently, ecograms give a more precise description of vegetation-site relationships if the considered nutrient gradients refer to appropriate soil parameters (e.g. S-value, C/N-ratio) and to measurements in the main root horizon. Measurements of the soil pH are often used to characterize the nutritional conditions for the vegetation at a particular site, for example in ecograms describing the ecological behaviour of a (tree) species. In spite of this use and interpretation of pH measurements, certain prerequisites must be fulfilled when pH-values are to be used as an expression of the nutritional conditions for plants. Our study in deciduous forests of northern Germany shows that in strongly acid soils (pH < 4) the correlations be-tween pH-values on the one hand and S-value, base saturation and C/N-ratios on the other hand are generally low, particularly for measurements in the upper mineral horizon. By contrast, in moderately acid to neutral soils (pH 4–7), pH-values could be employed as a useful indicator of the nutritional conditions for the vegetation. The main reason for the observed discrepancy in the indicator quality of pH measurements is the presence of distinct differences in the cation exchange capacity (CEC) of the soils, which are often not taken into account when interpreting pH measurements. CEC-values close to zero may be associated with either low or comparatively high pH-values, depending on whether or not the few available cation exchange sites are occupied by protons or bases, respectively. In general, pH – S-value/base saturation/CN-ratio relationships are stronger in soil samples taken in the main (fine and coarse) root horizon than in those from the upper mineral horizon. Consequently, ecograms give a more precise description of vegetation-site relationships if the considered nutrient gradients refer to appropriate soil parameters (e.g. S-value, C/N-ratio) and to measurements in the main root horizon.

In the present study we investigated long-term climate imprints (160 year period) on tree-ring δ15N signatures of sessile oak (Quercus petraea) at sites with contrasting water availability (i.e. Cambisols vs. Regosols in Luxembourg, Central Europe, with 175 mm and 42 mm available water capacity, respectively). We hypothesized that tree-ring δ15N signatures constitute a sensitive indicator to long-term shifts in climatic conditions. Our findings revealed a close positive correlation between winter and spring temperatures and tree-ring δ15N signatures. These relationships were stronger for Cambisol than for Regosol sites. If entire chronologies were considered, peaks in annual mean temperatures closely corresponded with peaks in tree-ring δ15N signatures, with both annual mean temperatures and δ15N signatures reaching their maxima within the last two decades. In addition, we found a weak but significant impact of February precipitation on δ15N signatures, but only for Cambisols. We hypothesize that these findings are attributable to climate- (particularly temperature-) mediated nitrification rates in forest soils. As nitrification is a strongly fractionating process that produces 15N-depleted nitrate and higher isotopic ratios for ammonium in the topsoil, increased nitrification leads to 15N-enriched pools of Ninorg in the upper soil horizons and therefore higher δ15N signatures in plant tissues. Weaker correlations at Regosol sites were likely related to dryer and more acidic site conditions, both of which may reduce nitrification rates. Comparisons of oak and beech trees in the study area point to species-specific trajectories of wood nitrogen isotopes, likely related to the partitioning of ammonium and nitrate among species. In conclusion, tree-ring δ15N signatures may serve as an integrator of terrestrial N cycling and as such constitute a valuable tool in the identification of spatial and temporal patterns of N cycling in relation to environmental changes. Due to the mediating effects of the isotopic composition of the respective N sources, analyses of the isotopic composition of airborne N loads would support the interpretation of wood δ15N patterns, particularly in areas that are subject to high N pollution.

Migration of the herb species Anemone nemorosa from older woodlands into adjacent, recently established deciduous woodlands on former arable land was studied. The wood anemone had colonized part of all studied recent woods, varying in age between 30 and 75 years. Cover of A. nemorosa in the recent woods decreased with increasing distance from the older woodland at all sites but one, indicating dispersal limitation during colonization. The advancing edge of most populations was characterized by negative logarithmic decrease in cover with establishment of isolated pioneer individuals and later gradual infill between pioneers. Migration rates were calculated by three methods. The mean migration rate of A. nemorosa based on observed maximum cover in the recent woods was 0.20 m year1, 0.40 m year-1 based on half maximum cover and 0.85 m year1 based on the individual found farthest from the former woodland border. The calculated migration rates were consistently higher than the rate of possible rhizome growth. Seed dispersal and establishment is thus very important for colonization of new woodlands. Migration rates increased with tree canopy cover–especially cover of broad-leaved species with quickly decomposing litter–and with soil pH. Migration rates decreased with increasing grass cover in the field layer. These differences in migration rates may be due to increased micro-site availability for establishment at high canopy cover, low grass cover and high soil pH. Our results show that the wood anemone generally colonizes recent woods from nearby source populations. However, colonization proceeds relatively slowly and is limited by both seed dispersal and availability of suitable micro-sites. Gradients in abundance of A. nemorosa within secondary woods may be detectable for long periods of time and indicate the recent origin of a woodland.

1 We studied the migration of field layer plants across ecotones between ancient woodlands and recent deciduous woods on former arable land varying in age between 30 and 75 years. 2 Number and percentage cover of woodland species in recent woods decreased with increasing distance to the ancient woods, and increased with the age of the recent woods, indicating dispersal limitation during secondary succession. 3 Colonization by typical woodland plants was observed in 183 of 200 species × site combinations. In 72 combinations, a colonization front was characterized by logarithmic or linear decrease in species cover, indicating establishment of isolated individuals and gradual infill of gaps. This pattern was most common in ant-dispersed species and less frequent in species with adhesive or ingested seeds. 4 Migration rates were calculated for 49 woodland species. Mean migration rates based on maximum cover in recent woods varied from 0.00 to 1.00 m year−1 between species, with a median migration rate of 0.30 m year−1. Migration rates calculated on occurrence of the farthest individual ranged from 0.00 to 1.25 m year−1, with a median rate of 0.53 m year−1. 5 Ant-dispersed species had lower migration rates based on maximum cover compared with species with adhesive or ingested seeds. No differences between dispersal modes were found when comparing migration rates based on the farthest individuals. Most of the calculated migration rates (84%) exceeded the rate of possible vegetative spread of woodland species. 6 Establishment of a field layer vegetation in secondary woods comparable to that of the adjacent ancient stands proceeded at a rate of c. 0.3–0.5 m year−1. We conclude that scale and intensity of temperate forest management should be adjusted to the relatively slow migration of the field layer flora in order to enable complete recovery during a management cycle.

Summary: Biodiversity-ecosystem functioning (BEF) experiments address ecosystem-level consequences of species loss by comparing communities of high species richness with communities from which species have been gradually eliminated. BEF experiments originally started with microcosms in the laboratory and with grassland ecosystems. A new frontier in experimental BEF research is manipulating tree diversity in forest ecosystems, compelling researchers to think big and comprehensively. We present and discuss some of the major issues to be considered in the design of BEF experiments with trees and illustrate these with a new forest biodiversity experiment established in subtropical China (Xingangshan, Jiangxi Province) in 2009/2010. Using a pool of 40 tree species, extinction scenarios were simulated with tree richness levels of 1, 2, 4, 8 and 16 species on a total of 566 plots of 25·8 × 25·8 m each. The goal of this experiment is to estimate effects of tree and shrub species richness on carbon storage and soil erosion; therefore, the experiment was established on sloped terrain. The following important design choices were made: (i) establishing many small rather than fewer larger plots, (ii) using high planting density and random mixing of species rather than lower planting density and patchwise mixing of species, (iii) establishing a map of the initial 'ecoscape' to characterize site heterogeneity before the onset of biodiversity effects and (iv) manipulating tree species richness not only in random but also in trait-oriented extinction scenarios. Data management and analysis are particularly challenging in BEF experiments with their hierarchical designs nesting individuals within-species populations within plots within-species compositions. Statistical analysis best proceeds by partitioning these random terms into fixed-term contrasts, for example, species composition into contrasts for species richness and the presence of particular functional groups, which can then be tested against the remaining random variation among compositions. We conclude that forest BEF experiments provide exciting and timely research options. They especially require careful thinking to allow multiple disciplines to measure and analyse data jointly and effectively. Achieving specific research goals and synergy with previous experiments involves trade-offs between different designs and requires manifold design decisions.

The aim of this study was to analyse the relation of horizontal and vertical canopy structure to tree functional diversity of a highly diverse subtropical broad-leaved slope forest, stratified for different successional stages. This is of particular interest because many key ecosystem processes and functions are related to the arrangement of forest canopies. We assessed the effect of stand-related functional diversity (FDQ, measured as Rao’s quadratic entropy of leaf traits), together with other environmental variables on horizontal [measured as relative crown projection areas (CPAr)] and vertical [relative crown overlap, coefficients of variation (CV) of crown positioning variables] structure of the upper canopy at the local neighbourhood level. The analyses with mixed effects models revealed a negative relation (p = 0.025; estimate −0.07) between FDQ and CPAr. No significant effect of FDQ on vertical canopy structure has been found (p > 0.05). The findings are discussed with regard to resource partitioning and niche differentiation of canopy and sub-canopy species. Successional stage positively impacted the CV of crown length (p = 0.019; estimate 0.03) but did not affect other response variables. The sloping terrain strongly influenced vertical canopy structure as revealed by the significant effect of slope inclination on CV of crown length (p = 0.004; estimate −0.05) and of slope aspect on CV of mean crown height (p = 0.036; estimate −0.03). The high complexity of vertical crown positioning depending on the heterogeneous sloping terrain of the study area may have obscured relations of FDQ to vertical canopy structure.

Subtropical broad-leaved forests in southeastern China support a high diversity of woody plants. Using a comparative study design with 30×30 m plots (n1/427) from five successional stages (1 m in height in each plot and counted all woody recruits (bank of all seedlings ≤1 m in height) in each central 10×10 m quadrant of each plot. In addition, we measured a number of environmental variables (elevation, slope, aspect, soil moisture, pH, C, N, and C/N ratio) and biotic structural variables (height and cover of layers). Adult species richness varied from 25 to 69 species per plot, and in total 148 woody species from 46 families were recorded. There was a clear successional gradient in species composition as revealed by nonmetric multidimensional scaling (NMDS), but only a poor differentiation of different successional stages with respect to particular species. Adult richness per 100 individuals (rarefaction method) increased with successional stage. None of the measured abiotic variables were significantly correlated with adult species richness. We found no evidence that rare species were responsible for the increasing adult species richness, as richness of rare species among both adults and recruits was independent of the successional stage. Furthermore, the similarity between established adults and recruits did not increase with successional stage. There was a constant number of recruit species and also of exclusive recruit species, i.e., those that had not been present as adult individuals, across all successional stages, suggesting a continuous random immigration over time. © 2011 by the Ecological Society of America.

There is growing evidence from different sources that prolonged high N deposition causes a shift from nitrogen (N) limitation to nitrogen and phosphorus (P) co-limitation or even P limitation in many terrestrial ecosystems. However, the number of ecosystems where the type of limitation has been directly tested by longer-term full-factorial field experiments is very limited. We conducted a 5-year fertilization experiment with N and P in the Lüneburger Heide (NW Germany) to test the hypothesis that, following decades of elevated atmospheric N inputs, plant growth in dry lowland heaths may have shifted from N to N-P co-limitation or P limitation. We also tested whether the plant tissue N:P ratio reflects the type of nutrient limitation in a continental lowland heathland. Experimental plots dominated by Calluna vulgaris received regular additions of N (50 kg N ha -1 y -1), P (20 kg P ha -1 y -1), a combination of both, or water only (control) from 2004 to 2008. Over the whole study period, a highly significant positive N effect on shoot length was found, thus indicating N limitation. We conclude that a clear shift from N limitation to N-P co-limitation or P limitation has not yet occurred. Tissue N:P ratios showed a high temporal variability and no relationship between tissue N:P ratio and the shoot length response of Calluna to nutrient addition was found. The N:P tool is thus of limited use at the local scale and within the range of N:P ratio observed in this study, and should only be used as a rough indicator for the prediction of the type of nutrient limitation in lowland heathland on a larger geographical scale with a broader interval of N:P ratio.

Only very few cases have documented that an increase in connectivity after a period of fragmentation in ecological time has had an effect on the distribution, genetic structure and morphology of stenotopic species. In this study we present an example of clinal variability in a woodland ground beetle as a result of changes in the connectivity of a landscape during the last two centuries. The study area hosts both the nominate form C. violaceus s. str. and the subspecies C. v. purpurascens, which is ranked as a distinct species by some authors. We studied 12 Carabus violaceus populations from a 30 km transect of ancient and recent forests in north-western Germany. We analyzed three polymorphic enzyme loci, classified the elytron sculpture and measured the shape of the aedeagus tip of the specimens. C. violaceus showed secondary gradients both in allozyme markers and morphometric characters in our study area. A genetic differentiation of 16% between the populations is high but lies within the range of intraspecific variability in habitat specialists of the genus Carabus. Populations had no significant deficit of heterozygotes. We found many hybrid populations in terms of morphological properties. This study highlights the conservation value of ancient woodland and the consequences of landscape connectivity and defragmentation on the genetic setting of a ground beetle. Moreover, it shows that differences in the external shape of male genitalia do not prevent gene flow within the genus Carabus. Thus, the establishment of species status should not exclusively be based on this property.

Herb layer richness and composition of four forest types and three different management treatments were investigated in 16 deciduous forest stands of northern Germany. Specifically, we compared the species richness and composition occurring in mature forest stands that were single-tree and group selection harvested to those in unmanaged reference stands. Mean species richness of all herb layer species increased significantly with increasing harvest severity. When analyzing plant groups separately, it became obvious that this overall pattern was not consistent. While a negative relationship was detected between vernal herb richness and harvest severity, group selection harvest significantly increased species richness of summer herbaceous forest species and generalists. Woody species richness was not related to harvest severity. Community composition of the spring aspect was not significantly affected by selection harvest, whereas herb layer species composition in the summer aspect differed significantly among the three harvest treatments. A dominance of highly competitive shrub and generalist species was confined to some parts of the most intensively harvested stands. Overall, our results indicate that the herb layer community was not severely adversely affected by selection harvest at the intensities used in the studied stands. It is suggested that selection harvest systems may be feasible tools with which to conserve local forest vascular plant diversity and at the same time to meet the demand for timber products. However, information about forest history and the implementation of the selection harvest system are basic requirements when interpreting the results of studies on understorey response to selection harvest. Herb layer richness and composition of four forest types and three different management treatments were investigated in 16 deciduous forest stands of northern Germany. Specifically, we compared the species richness and composition occurring in mature forest stands that were single-tree and group selection harvested to those in unmanaged reference stands. Mean species richness of all herb layer species increased significantly with increasing harvest severity. When analyzing plant groups separately, it became obvious that this overall pattern was not consistent. While a negative relationship was detected between vernal herb richness and harvest severity, group selection harvest significantly increased species richness of summer herbaceous forest species and generalists. Woody species richness was not related to harvest severity. Community composition of the spring aspect was not significantly affected by selection harvest, whereas herb layer species composition in the summer aspect differed significantly among the three harvest treatments. A dominance of highly competitive shrub and generalist species was confined to some parts of the most intensively harvested stands. Overall, our results indicate that the herb layer community was not severely adversely affected by selection harvest at the intensities used in the studied stands. It is suggested that selection harvest systems may be feasible tools with which to conserve local forest vascular plant diversity and at the same time to meet the demand for timber products. However, information about forest history and the implementation of the selection harvest system are basic requirements when interpreting the results of studies on understorey response to selection harvest.

A large-scale field experiment with different management options (free-range grazing, onetime cutting, additional disturbances and their interactions) was carried out in a continental, dry heathland. Within the framework of the experiment seed production and germination ability of over-aged Calluna individuals as well as specific safe site conditions were investigated to reveal restricting factors for the generative rejuvenation of Calluna. The calculated values of seed production and germination ability of over-aged Calluna individuals can be compared with those from literature or exceed the values reported there. Therefore, seed production and germination ability are not the limiting factors for the generative rejuvenation of Calluna. Free-range grazing with large herbivores combined with onetime cutting of over-aged Calluna stands supports the generative rejuvenation, especially due to the creation of specific safe sites, which are crucial for a successful survival of Calluna seedlings. Particularly a light shadowing seems to be the restricting factor for a successful generative rejuvenation of Calluna in continental, dry heathlands.

Current knowledge about processes that generate long-distance dispersal of plants is still limited despite its importance for persistence of populations and colonization of new potential habitats. Today wild large mammals are presumed to be important vectors for long-distance transport of diaspores within and between European temperate forest patches, and in particular wild boars recently came into focus. Here we use a specific habit of wild boar, i.e. wallowing in mud and subsequent rubbing against trees, to evaluate epizoic dispersal of vascular plant diaspores. We present soil seed bank data from 27 rubbing trees versus 27 control trees from seven forest areas in Germany. The mean number of viable seeds and the plant species number were higher in soil samples near rubbing trees compared with control trees. Ten of the 20 most frequent species were more frequent, and many species exclusively appeared in the soil samples near rubbing trees. The large number of plant species and seeds – approximated > 1000 per tree – in the soils near rubbing trees is difficult to explain unless the majority were dispersed by wild boar. Hooked and bristly diaspores, i.e. those adapted to epizoochory, were more frequent, above that many species with unspecialised diaspores occurred exclusively near rubbing trees. Different to plant species closely tied to forest species which occur both in forest and open vegetation, and non-forest species were more frequent near rubbing trees compared with controls. These findings are consistent with previous studies on diaspore loads in the coats and hooves of shot wild boars. However, our method allows to identify the transport of diaspores from the open landscape into forest stands where they might especially emerge after disturbance, and a clustered distribution of epizoochorically dispersed seeds. Moreover, accumulation of seeds of wetness indicators near rubbing trees demonstrates directed dispersal of plant species inhabiting wet places between remote wallows. Das aktuelle Wissen über Prozesse, die zur Fernausbreitung von Pflanzen führen, ist trotz ihrer Bedeutung für das Überleben von Populationen und die Besiedlung neuer potenzieller Habitate noch immer sehr begrenzt. Wildlebende Großsäuger sind heutzutage vermutlich wichtige Vektoren für den Ferntransport von Diasporen innerhalb und zwischen den einzelnen Waldflächen in Mitteleuropa, und speziell das Wildschwein (Sus scrofa L.) spielt dabei offenbar eine herausragende Rolle. Wir nutzen hier ein spezifisches Verhalten des Wildschweins – Suhlen im Schlamm und nachfolgendes Scheuern an sogenannten Malbäumen – um die epizoochore Ausbreitung von Gefäßpflanzen-Diasporen einzuschätzen. Dargestellt werden die Ergebnisse von Samenbank-Untersuchungen von 27 Malbäumen im Vergleich zu 27 Kontrollbäumen aus sieben Waldgebieten in Deutschland. Sowohl die mittlere Zahl lebensfähiger Samen als auch die Artenzahl waren höher in Bodenproben neben Malbäumen. Zehn der 20 in der Samenbank verbreitetsten Pflanzenarten hatten hier ihren Schwerpunkt, und viele Arten kamen ausschließlich in den neben Malbäumen gewonnenen Proben vor. Die große Zahl von Pflanzenarten und Samen – zumindest > 1000 pro Baum – im Boden an Malbäumen lässt sich nur durch die Aktivität der Wildschweine erklären. Mit Haken oder Borsten ausgestattete, d.h. an Epizoochorie angepasste Diasporen waren häufiger, aber auch viele Arten mit unspezialisierten Diasporen kamen ausschließlich in der Samenbank bei Malbäumen vor. Anders als weitgehend an Wald gebundene Pflanzenarten waren solche, die sowohl im Wald und im Offenland vorkommen, sowie nicht im Wald vorkommende Arten häufiger neben Malbäumen als neben Kontrollbäumen. Diese Befunde stimmen mit denen früherer Untersuchungen von Diasporenladungen im Fell und in den Hufen geschossener Wildschweine überein. Unsere Methode erlaubt darüber hinaus aber die Identifizierung des Diasporentransports aus dem Offenland in die Waldbestände, wo sie insbesondere nach Störungen keimen dürften, sowie einer ungleichmäßigen Verteilung epizoochor ausgebreiteter Diasporen. Außerdem zeigt die Akkumulation von Samen von Nässezeigern neben den Malbäumen eine gezielte Ausbreitung nasse Standorte bewohnender Pflanzenarten zwischen entfernt gelegenen Suhlen.