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Tree species richness, forest structure, and seasonal fluctuations between rainy and dry seasons can strongly affect trophic interactions in forest ecosystems, but the inter- and scale dependence of these variables remains unclear. Using artificial caterpillars (~18,000 replicates), we analyzed predation pressure by arthropods, birds, and rodents along a tree species richness gradient across seasons in a subtropical tree diversity experiment (BEF-China). The aim of the study was to test if forest structure, in addition to tree species richness, has an effect on predation pressure and to further specify which structural variables are important in driving predation. We assessed the effects of tree species richness and forest structure at the plot and local neighborhood levels. We also included fine-scale placement covariates, plot size, and topographical covariates of the study site. Forest structure and tree species richness independently and interactively affected predation pressure. The spatial scale was an important determinant for tree species richness and structural effects, extending from within plot scales to the overall heterogeneity of the plots’ surrounding environment. For example, the effect of branch density in the local neighborhood depended on both surrounding tree species richness and plot-level vegetation density. Similarly, visibility-enhancing factors increased attacks by arthropods (lack of branches in close surroundings) and by birds (open area), depending on the surrounding vegetation. A comparison of structural measures showed that predation pressure can be addressed in much greater detail with multiple specific structural features than with overall forest complexity. Seasonal change also affected predation pressure, with foliage being a stronger attractant in spring, but also by presumable topography-driven study plot differences in sun exposure and humidity between rainy and dry seasons. Our study demonstrates that predation pressure is not simply a function of tree species richness or structure but is shaped by the interplay of structural elements, spatial scale, and seasonal dynamics along gradients of tree species richness and forest structure. The structural and seasonal effects are important to take into account when addressing how current and future biodiversity loss may change top‐down control of herbivory and overall ecosystem functioning.

Measurement(s) vegetation-plot resurvey data of vascular plant species Technology Type(s) vegetation-plot records Factor Type(s) Cover of species in plots Sample Characteristic - Organism Vascular plant species Sample Characteristic - Environment Terrestrial habitats Sample Characteristic - Location Germany

Efforts to naturally remove atmospheric CO2 demand that largely intact forests be maintained. Our inter-cultural research initiative tested the hypothesis that Indigenous custody of the land is compatible with the maintenance of intact forests. Here we combined traditional knowledge, phytolith analysis, remote sensing, and tree inventories to study old-growth forests in Panama's Darién. Phytoliths served to elucidate historical vegetation, remote sensing revealed the current and past Indigenous footprints while tree stature and identity characterised the forest. Until now there has been very little to no human impact within these forests and current Indigenous footprint is both small and stable. Large trees accounted for 13% of trees in the plots that we established. For over half of the species, the measured tree height was taller than previously published maximum heights, leading us to conclude that these forests are a truly exceptional ecological refugium. Noting that the local communities are not rewarded for their custody of these exceptional forests we call to revisit the Good Practice Guidance for Land Use Land Use Change and Forestry to include intact forest land. In the context of sub-optimal carbon finance options, we also propose matching as a methodology that could prove additionality of forest conservation initiatives in climate mitigation portfolios.

Abstract 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

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.

Here, Schuldt et al. collate data from two long-term grassland and forest biodiversity experiments to ask how plant diversity facets affect the diversity of higher trophic levels. The results show that positive effects of plant diversity on consumer diversity are mediated by plant structural and functional diversity, and vary across ecosystems and trophic levels.

Reforestation in sloping terrain is an important measure for soil erosion control and sustainable watershed management. The mechanical stability of such reforested stands, however, can be low due to a strong asymmetric shape of tree crowns. We investigated how neighbourhood tree species richness, neighbourhood pressure, tree height, and slope inclination affect crown asymmetry in a large-scale plantation biodiversity-ecosystem functioning experiment in subtropical China (BEF-China) over eight years. We took the advantage of terrestrial laser scanning (TLS) measurements, which provide non-destructive, high-resolution data of tree structure without altering tree interactions. Neighbourhood species richness significantly reduced crown asymmetry, and this effect became stronger at steeper slopes. Our results suggest that tree diversity promotes the mechanical stability of forest stands in sloping terrain and highlight the importance of TLS-data for a comprehensive understanding of the role of tree diversity in modulating crown interactions in mixed-species forest plantations.

Biodiversity change can impact ecosystem functioning, though this is primarily studied at lower trophic levels. Here, Schuldt et al. find that biodiversity components other than tree species richness are particularly important, and higher trophic level diversity plays a role in multifunctionality.

Though biodiversity is expected to be important in productivity in tree communities, there is little empirical evidence of this at local scales. Here, Fichtner et al. show that higher neighbourhood species richness increased tree growth, explaining over half of the variation in community productivity.

The effects of atmospheric nitrogen (N) deposition on ecosystem functioning largely depend on the retention of N in different ecosystem compartments, but accumulation and partitioning processes have rarely been quantified in long-term field experiments. In the present study we analysed for the first time decadal-scale flows and allocation patterns of N in a heathland ecosystem that has been subject to airborne N inputs over decades. Using a long-term 15N tracer experiment, we quantified N retention and flows to and between ecosystem compartments (above-ground/below-ground vascular biomass, moss layer, soil horizons, leachate). After 9 years, about 60% of the added 15N-tracer remained in the N cycle of the ecosystem. The moss layer proved to be a crucial link between incoming N and its allocation to different ecosystem compartments (in terms of a short-term capture, but long-term release function). However, about 50% of the 15N captured and released by the moss layer was not compensated for by a corresponding increase in recovery rates in any other compartment, probably due to denitrification losses from the moss layer in the case of water saturation after rain events. The O-horizon proved to be the most important long-term sink for added 15N, as reflected by an increase in recovery rates from 18 to 40% within 8 years. Less than 2.1% of 15N were recovered in the podzol-B-horizon, suggesting that only negligible amounts of N were withdrawn from the N cycle of the ecosystem. Moreover, 15N recovery was low in the dwarf shrub above-ground biomass (

Understanding how trees respond to global change drivers is central to predict changes in forest structure and functions. Although there is evidence on the mode of nitrogen (N) and drought (D) effects on tree growth, our understanding of the interplay of these factors is still limited. Simultaneously, as mixtures are expected to be less sensitive to global change as compared to monocultures, we aimed to investigate the combined effects of N addition and D on the productivity of three tree species (Fagus sylvatica, Quercus petraea, Pseudotsuga menziesii) in relation to functional diverse species mixtures using data from a four-year field experiment in Northwest Germany. Here we show that species mixing can mitigate the negative effects of combined N fertilization and D events, but the community response is mainly driven by the combination of certain traits rather than the tree species richness of a community. For beech, we found that negative effects of D on growth rates were amplified by N fertilization (i.e. combined treatment effects were non-additive), while for oak and fir, the simultaneous effects of N and D were additive. Beech and oak were identified as most sensitive to combined N+D effects with a strong size-dependency observed for beech, suggesting that the negative impact of N+D becomes stronger with time as beech grows larger. As a consequence, the net biodiversity effect declined at the community level, which can be mainly assigned to a distinct loss of complementarity in beech-oak mixtures. This pattern, however, was not evident in the other species-mixtures, indicating that neighborhood composition (i.e. trait combination), but not tree species richness mediated the relationship between tree diversity and treatment effects on tree growth. Our findings point to the importance of the qualitative role (‘trait portfolio’) that biodiversity play in determining resistance of diverse tree communities to environmental changes. As such, they provide further understanding for adaptive management strategies in the context of global change.

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.

Soil erosion is a key threat to many ecosystems, especially in subtropical China where high erosion rates occur. While the mechanisms that induce soil erosion on agricultural land are well understood, soil erosion processes in forests have rarely been studied. Throughfall kinetic energy (TKE) is influenced in manifold ways and often determined by the tree's leaf and architectural traits. We investigated the role of species identity in mono-specific stands on TKE by asking to what extent TKE is species-specific and which leaf and architectural traits account for variation in TKE. We measured TKE of 11 different tree species planted in monocultures in a biodiversity-ecosystem-functioning experiment in subtropical China, using sand-filled splash cups during five natural rainfall events in summer 2013. In addition, 14 leaf and tree architectural traits were measured and linked to TKE. Our results showed that TKE was highly species-specific. Highest TKE was found below Choerospondias axillaris and Sapindus saponaria, while Schima superba showed lowest TKE. These species-specific effects were mediated by leaf habit, leaf area (LA), leaf pinnation, leaf margin, stem diameter at ground level (GD), crown base height (CBH), tree height, number of branches and leaf area index (LAI) as biotic factors and throughfall as abiotic factor. Among these, leaf habit, tree height and LA showed the highest effect sizes on TKE and can be considered as major drivers of TKE. TKE was positively influenced by LA, GD, CBH, tree height, LAI, and throughfall amount while it was negatively influenced by the number of branches. TKE was lower in evergreen, simple leaved and dentate leaved than in deciduous, pinnated or entire leaved species. Our results clearly showed that soil erosion in forest plantations can be mitigated by the appropriate choice of tree species.

The role of competition in tree communities is increasingly well understood, while little is known about the patterns and mechanisms of the interplay between above- and belowground competition in tree communities. This knowledge, however, is crucial for a better understanding of community dynamics and developing adaptive near-natural management strategies. We assessed neighbourhood interactions in an unmanaged old-growth European beech (Fagus sylvatica) forest by quantifying variation in the intensity of above- (shading) and belowground competition (crowding) among dominant and co-dominant canopy beech trees during tree maturation. Shading had on average a much larger impact on radial growth than crowding and the sensitivity to changes in competitive conditions was lowest for crowding effects. We found that each mode of competition reduced the effect of the other. Increasing crowding reduced the negative effect of shading, and at high levels of shading, crowding actually had a facilitative effect and increased growth. Our study demonstrates that complementarity in above- and belowground processes enable F. sylvatica to alter resource acquisition strategies, thus optimising tree radial growth. As a result, competition seemed to become less important in stands with a high growing stock and tree communities with a long continuity of anthropogenic undisturbed population dynamics. We suggest that growth rates do not exclusively depend on the density of potential competitors at the intraspecific level, but on the conspecific aggregation of large-diameter trees and their functional role for regulating biotic filtering processes. This finding highlights the potential importance of the rarely examined relationship between the spatial aggregation pattern of large-diameter trees and the outcome of neighbourhood interactions, which may be central to community dynamics and the related forest ecosystem services.

Future climates are likely to include extreme events, which in turn have great impacts on ecological systems. In this study, we investigated possible effects that could mitigate stem breakage caused by a rare and extreme ice storm in a Chinese subtropical forest across a gradient of forest diversity. We used Bayesian modeling to correct stem breakage for tree size and variance components analysis to quantify the influence of taxon, leaf and wood functional traits, and stand level properties on the probability of stem breakage. We show that the taxon explained four times more variance in individual stem breakage than did stand level properties; trees with higher specific leaf area (SLA) were less susceptible to breakage. However, a large part of the variation at the taxon scale remained unexplained, implying that unmeasured or undefined traits could be used to predict damage caused by ice storms. When aggregated at the plot level, functional diversity and wood density increased after the ice storm. We suggest that for the adaption of forest management to climate change, much can still be learned from looking at functional traits at the taxon level.

There is ample evidence that continuously existing forests and afforestations on previously agricultural land differ with regard to ecosystem functions and services such as carbon sequestration, nutrient cycling and biodiversity. However, no studies have so far been conducted on possible long-term (>100 years) impacts on tree growth caused by differences in the ecological continuity of forest stands. In the present study we analysed the variation in tree-ring width of sessile oak (Quercus petraea (Matt.) Liebl.) trees (mean age 115-136 years) due to different land-use histories (continuously existing forests, afforestations both on arable land and on heathland). We also analysed the relation of growth patterns to soil nutrient stores and to climatic parameters (temperature, precipitation). Tree rings formed between 1896 and 2005 were widest in trees afforested on arable land. This can be attributed to higher nitrogen and phosphorous availability and indicates that former fertilisation may continue to affect the nutritional status of forest soils for more than one century after those activities have ceased. Moreover, these trees responded more strongly to environmental changes - as shown by a higher mean sensitivity of the tree-ring widths - than trees of continuously existing forests. However, the impact of climatic parameters on the variability in tree-ring width was generally small, but trees on former arable land showed the highest susceptibility to annually changing climatic conditions. We assume that incompletely developed humus horizons as well as differences in the edaphon are responsible for the more sensitive response of oak trees of recent forests (former arable land and former heathland) to variation in environmental conditions. We conclude that forests characterised by a long ecological continuity may be better adapted to global change than recent forest ecosystems.

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. [ABSTRACT FROM AUTHOR]