Climate variability and aridity modulate the role of leaf shelters for arthropods: A global experiment.

Current climate change is disrupting biotic interactions and eroding biodiversity worldwide. However, species sensitive to aridity, high temperatures, and climate variability might find shelter in microclimatic refuges, such as leaf rolls built by arthropods. To explore how the importance of leaf shelters for terrestrial arthropods changes with latitude, elevation, and climate, we conducted a distributed experiment comparing arthropods in leaf rolls versus control leaves across 52 sites along an 11,790 km latitudinal gradient. We then probed the impact of short- versus long-term climatic impacts on roll use, by comparing the relative impact of conditions during the experiment versus average, baseline conditions at the site. Leaf shelters supported larger organisms and higher arthropod biomass and species diversity than non-rolled control leaves. However, the magnitude of the leaf rolls' effect differed between long- and short-term climate conditions, metrics (species richness, biomass, and body size), and trophic groups (predators vs. herbivores). The effect of leaf rolls on predator richness was influenced only by baseline climate, increasing in magnitude in regions experiencing increased long-term aridity, regardless of latitude, elevation, and weather during the experiment. This suggests that shelter use by predators may be innate, and thus, driven by natural selection. In contrast, the effect of leaf rolls on predator biomass and predator body size decreased with increasing temperature, and increased with increasing precipitation, respectively, during the experiment. The magnitude of shelter usage by herbivores increased with the abundance of predators and decreased with increasing temperature during the experiment. Taken together, these results highlight that leaf roll use may have both proximal and ultimate causes. Projected increases in climate variability and aridity are, therefore, likely to increase the importance of biotic refugia in mitigating the effects of climate change on species persistence.

Selection for niche differentiation in plant communities increases biodiversity effects.

In experimental plant communities, relationships between biodiversity and ecosystem functioning have been found to strengthen over time, a fact often attributed to increased resource complementarity between species in mixtures and negative plant-soil feedbacks in monocultures. Here we show that selection for niche differentiation between species can drive this increasing biodiversity effect. Growing 12 grassland species in test monocultures and mixtures, we found character displacement between species and increased biodiversity effects when plants had been selected over 8 years in species mixtures rather than in monocultures. When grown in mixtures, relative differences in height and specific leaf area between plant species selected in mixtures (mixture types) were greater than between species selected in monocultures (monoculture types). Furthermore, net biodiversity and complementarity effects were greater in mixtures of mixture types than in mixtures of monoculture types. Our study demonstrates a novel mechanism for the increase in biodiversity effects: selection for increased niche differentiation through character displacement. Selection in diverse mixtures may therefore increase species coexistence and ecosystem functioning in natural communities and may also allow increased mixture yields in agriculture or forestry. However, loss of biodiversity and prolonged selection of crops in monoculture may compromise this potential for selection in the longer term.

Towards an Integrative, Eco-Evolutionary Understanding of Ecological Novelty: Studying and Communicating Interlinked Effects of Global Change.

Global change has complex eco-evolutionary consequences for organisms and ecosystems, but related concepts (e.g., novel ecosystems) do not cover their full range. Here we propose an umbrella concept of "ecological novelty" comprising (1) a site-specific and (2) an organism-centered, eco-evolutionary perspective. Under this umbrella, complementary options for studying and communicating effects of global change on organisms, ecosystems, and landscapes can be included in a toolbox. This allows researchers to address ecological novelty from different perspectives, e.g., by defining it based on (a) categorical or continuous measures, (b) reference conditions related to sites or organisms, and (c) types of human activities. We suggest striving for a descriptive, non-normative usage of the term "ecological novelty" in science. Normative evaluations and decisions about conservation policies or management are important, but require additional societal processes and engagement with multiple stakeholders.

Integrating community assembly and biodiversity to better understand ecosystem function: the Community Assembly and the Functioning of Ecosystems (CAFE) approach.

The research of a generation of ecologists was catalysed by the recognition that the number and identity of species in communities influences the functioning of ecosystems. The relationship between biodiversity and ecosystem functioning (BEF) is most often examined by controlling species richness and randomising community composition. In natural systems, biodiversity changes are often part of a bigger community assembly dynamic. Therefore, focusing on community assembly and the functioning of ecosystems (CAFE), by integrating both species richness and composition through species gains, losses and changes in abundance, will better reveal how community changes affect ecosystem function. We synthesise the BEF and CAFE perspectives using an ecological application of the Price equation, which partitions the contributions of richness and composition to function. Using empirical examples, we show how the CAFE approach reveals important contributions of composition to function. These examples show how changes in species richness and composition driven by environmental perturbations can work in concert or antagonistically to influence ecosystem function. Considering how communities change in an integrative fashion, rather than focusing on one axis of community structure at a time, will improve our ability to anticipate and predict changes in ecosystem function.

Plant selection and soil legacy enhance long-term biodiversity effects.

Plant-plant and plant-soil interactions can help maintain plant diversity and ecosystem functions. Changes in these interactions may underlie experimentally observed increases in biodiversity effects over time via the selection of genotypes adapted to low or high plant diversity. Little is known, however, about such community-history effects and particularly the role of plant-soil interactions in this process. Soil-legacy effects may occur if co-evolved interactions with soil communities either positively or negatively modify plant biodiversity effects. We tested how plant selection and soil legacy influence biodiversity effects on productivity, and whether such effects increase the resistance of the communities to invasion by weeds. We used two plant selection treatments: parental plants growing in monoculture or in mixture over 8 yr in a grassland biodiversity experiment in the field, which we term monoculture types and mixture types. The two soil-legacy treatments used in this study were neutral soil inoculated with live or sterilized soil inocula collected from the same plots in the biodiversity experiment. For each of the four factorial combinations, seedlings of eight species were grown in monocultures or four-species mixtures in pots in an experimental garden over 15 weeks. Soil legacy (live inoculum) strongly increased biodiversity complementarity effects for communities of mixture types, and to a significantly weaker extent for communities of monoculture types. This may be attributed to negative plant-soil feedbacks suffered by mixture types in monocultures, whereas monoculture types had positive plant-soil feedbacks, in both monocultures and mixtures. Monocultures of mixture types were most strongly invaded by weeds, presumably due to increased pathogen susceptibility, reduced biomass, and altered plant-soil interactions of mixture types. These results show that biodiversity effects in experimental grassland communities can be modified by the evolution of positive vs. negative plant-soil feedbacks of plant monoculture vs. mixture types.

Plant selection and soil legacy enhance long-term biodiversity effects.

Plant-plant and plant-soil interactions can help maintain plant diversity and ecosystem functions. Changes in these interactions may underlie experimentally observed increases in biodiversity effects over time via the selection of genotypes adapted to low or high plant diversity. Little is known, however, about such community-history effects and particularly the role of plant-soil interactions in this process. Soil-legacy effects may occur if co-evolved interactions with soil communities either positively or negatively modify plant biodiversity effects. We tested how plant selection and soil legacy influence biodiversity effects on productivity, and whether such effects increase the resistance of the communities to invasion by weeds. We used two plant selection treatments: parental plants growing in monoculture or in mixture over 8 yr in a grassland biodiversity experiment in the field, which we term monoculture types and mixture types. The two soil-legacy treatments used in this study were neutral soil inoculated with live or sterilized soil inocula collected from the same plots in the biodiversity experiment. For each of the four factorial combinations, seedlings of eight species were grown in monocultures or four-species mixtures in pots in an experimental garden over 15 weeks. Soil legacy (live inoculum) strongly increased biodiversity complementarity effects for communities of mixture types, and to a significantly weaker extent for communities of monoculture types. This may be attributed to negative plant-soil feedbacks suffered by mixture types in monocultures, whereas monoculture types had positive plant-soil feedbacks, in both monocultures and mixtures. Monocultures of mixture types were most strongly invaded by weeds, presumably due to increased pathogen susceptibility, reduced biomass, and altered plant-soil interactions of mixture types. These results show that biodiversity effects in experimental grassland communities can be modified by the evolution of positive vs. negative plant-soil feedbacks of plant monoculture vs. mixture types.

Effects of management on aquatic tree-hole communities in temperate forests are mediated by detritus amount and water chemistry.

Arthropod communities in water-filled tree holes may be sensitive to impacts of forest management, for example via changes in environmental conditions such as resource input. We hypothesized that increasing forest management intensity (ForMI) negatively affects arthropod abundance and richness and shifts community composition and trophic structure of tree hole communities. We predicted that this shift is caused by reduced habitat and resource availability at the forest stand scale as well as reduced tree hole size, detritus amount and changed water chemistry at the tree holes scale. We mapped 910 water-filled tree holes in two regions in Germany and studied 199 tree hole inhabiting arthropod communities. We found that increasing ForMI indeed significantly reduced arthropod abundance and richness in water-filled tree holes. The most important indirect effects of management intensity on tree hole community structure were the reduced amounts of detritus for the tree hole inhabiting organisms and changed water chemistry at the tree hole scale, both of which seem to act as a habitat filter. Although habitat availability at the forest stand scale decreased with increasing management intensity, this unexpectedly increased local arthropod abundance in individual tree holes. However, regional species richness in tree holes significantly decreased with increasing management intensity, most likely due to decreased habitat diversity. We did not find that the management-driven increase in plant diversity at the forest stand scale affected communities of individual tree holes, for example via resource availability for adults. Our results suggest that management of temperate forests has to target a number of factors at different scales to conserve diverse arthropod communities in water-filled tree holes.

Bottom-up effects of plant diversity on multitrophic interactions in a biodiversity experiment.

Biodiversity is rapidly declining, and this may negatively affect ecosystem processes, including economically important ecosystem services. Previous studies have shown that biodiversity has positive effects on organisms and processes across trophic levels. However, only a few studies have so far incorporated an explicit food-web perspective. In an eight-year biodiversity experiment, we studied an unprecedented range of above- and below-ground organisms and multitrophic interactions. A multitrophic data set originating from a single long-term experiment allows mechanistic insights that would not be gained from meta-analysis of different experiments. Here we show that plant diversity effects dampen with increasing trophic level and degree of omnivory. This was true both for abundance and species richness of organisms. Furthermore, we present comprehensive above-ground/below-ground biodiversity food webs. Both above ground and below ground, herbivores responded more strongly to changes in plant diversity than did carnivores or omnivores. Density and richness of carnivorous taxa was independent of vegetation structure. Below-ground responses to plant diversity were consistently weaker than above-ground responses. Responses to increasing plant diversity were generally positive, but were negative for biological invasion, pathogen infestation and hyperparasitism. Our results suggest that plant diversity has strong bottom-up effects on multitrophic interaction networks, with particularly strong effects on lower trophic levels. Effects on higher trophic levels are indirectly mediated through bottom-up trophic cascades.

Dominant predators mediate the impact of habitat size on trophic structure in bromeliad invertebrate communities.

Local habitat size has been shown to influence colonization and extinction processes of species in patchy environments. However, species differ in body size, mobility, and trophic level, and may not respond in the same way to habitat size. Thus far, we have a limited understanding of how habitat size influences the structure of multitrophic communities and to what extent the effects may be generalizable over a broad geographic range. Here, we used water-filled bromeliads of different sizes as a natural model system to examine the effects of habitat size on the trophic structure of their inhabiting invertebrate communities. We collected composition and biomass data from 651 bromeliad communities from eight sites across Central and South America differing in environmental conditions, species pools, and the presence of large-bodied odonate predators. We found that trophic structure in the communities changed dramatically with changes in habitat (bromeliad) size. Detritivore : resource ratios showed a consistent negative relationship with habitat size across sites. In contrast, changes in predator: detritivore (prey) ratios depended on the presence of odonates as dominant predators in the regional pool. At sites without odonates, predator: detritivore biomass ratios decreased with increasing habitat size. At sites with odonates, we found odonates to be more frequently present in large than in small bromeliads, and predator: detritivore biomass ratios increased with increasing habitat size to the point where some trophic pyramids became inverted. Our results show that the distribution of biomass amongst food-web levels depends strongly on habitat size, largely irrespective of geographic differences in environmental conditions or detritivore species compositions. However, the presence of large-bodied predators in the regional species pool may fundamentally alter this relationship between habitat size and trophic structure. We conclude that taking into account the response and multitrophic effects of dominant, mobile species may be critical when predicting changes in community structure along a habitat-size gradient.

A comparison of the strength of biodiversity effects across multiple functions.

In order to predict which ecosystem functions are most at risk from biodiversity loss, meta-analyses have generalised results from biodiversity experiments over different sites and ecosystem types. In contrast, comparing the strength of biodiversity effects across a large number of ecosystem processes measured in a single experiment permits more direct comparisons. Here, we present an analysis of 418 separate measures of 38 ecosystem processes. Overall, 45 % of processes were significantly affected by plant species richness, suggesting that, while diversity affects a large number of processes not all respond to biodiversity. We therefore compared the strength of plant diversity effects between different categories of ecosystem processes, grouping processes according to the year of measurement, their biogeochemical cycle, trophic level and compartment (above- or belowground) and according to whether they were measures of biodiversity or other ecosystem processes, biotic or abiotic and static or dynamic. Overall, and for several individual processes, we found that biodiversity effects became stronger over time. Measures of the carbon cycle were also affected more strongly by plant species richness than were the measures associated with the nitrogen cycle. Further, we found greater plant species richness effects on measures of biodiversity than on other processes. The differential effects of plant diversity on the various types of ecosystem processes indicate that future research and political effort should shift from a general debate about whether biodiversity loss impairs ecosystem functions to focussing on the specific functions of interest and ways to preserve them individually or in combination.

Aquatic islands in the sky: 100 years of research on water-filled tree holes.

Water-filled tree holes are unique ecosystems that may occur high up in tree crowns and are essentially aquatic islands in the sky. Insect larvae, mesofauna, and other organisms colonize the waterbodies and feed on the accumulating detritus. Water-filled tree holes are not only important habitats for these species but have been used as model systems in ecology. Here, we review more than 100 years of research on tree-hole inhabiting organisms and show that most studies focus on selected or even single species (most of which are mosquitoes), whereas only few studies examine groups other than insects, especially in the tropics. Using a vote counting of results and a meta-analysis of community studies, we show that the effects of tree-hole size and resources on abundance and richness were investigated most frequently. Both were found to have a positive effect, but effect sizes were modulated by site-specific environmental variables such as temperature or precipitation. We also show that parameters such as the height of the tree holes above ground, tree-hole density, predation, and detritus type can be important drivers of organism abundance or richness but are less often tested. We identify several important research gaps and potential avenues for future research. Specifically, future studies should investigate the structure, functions, and temporal dynamics of tree-hole food webs and their cross-system interactions, for example, with terrestrial predators that act as a connection to their terrestrial surroundings in meta-ecosystems. Global observational or experimental tree-hole studies could contribute pivotal information on spatial variation of community structure and environmental drivers of community assembly. With a better understanding of these unique aquatic habitats in terrestrial ecosystems, natural and artificial tree holes can not only serve as model systems for addressing fundamental ecological questions but also serve as indicator systems of the impacts of environmental change on ecosystems.

Nonlinear effects of environmental drivers shape macroinvertebrate biodiversity in an agricultural pondscape.

Agriculture is a leading cause of biodiversity loss and significantly impacts freshwater biodiversity through many stressors acting locally and on the landscape scale. The individual effects of these numerous stressors are often difficult to disentangle and quantify, as they might have nonlinear impacts on biodiversity. Within agroecosystems, ponds are biodiversity hotspots providing habitat for many freshwater species and resting or feeding places for terrestrial organisms. Ponds are strongly influenced by their terrestrial surroundings, and understanding the determinants of biodiversity in agricultural landscapes remains difficult but crucial for improving conservation policies and actions. We aimed to identify the main effects of environmental and spatial variables on α-, ß-, and γ-diversities of macroinvertebrate communities inhabiting ponds (n = 42) in an agricultural landscape in the Northeast Germany, and to quantify the respective roles of taxonomic turnover and nestedness in the pondscape. We disentangled the nonlinear effects of a wide range of environmental and spatial variables on macroinvertebrate α- and ß-biodiversity. Our results show that α-diversity is impaired by eutrophication (phosphate and nitrogen) and that overshaded ponds support impoverished macroinvertebrate biota. The share of arable land in the ponds' surroundings decreases ß-diversity (i.e., dissimilarity in community), while ß-diversity is higher in shallower ponds. Moreover, we found that ß-diversity is mainly driven by taxonomic turnover and that ponds embedded in arable fields support local and regional diversity. Our findings highlight the importance of such ponds for supporting biodiversity, identify the main stressors related to human activities (eutrophication), and emphasize the need for a large number of ponds in the landscape to conserve biodiversity. Small freshwater systems in agricultural landscapes challenge us to compromise between human demands and nature conservation worldwide. Identifying and quantifying the effects of environmental variables on biodiversity inhabiting those ecosystems can help address threats impacting freshwater life with more effective management of pondscapes.

Pathways for cross-boundary effects of biodiversity on ecosystem functioning.

The biodiversity-ecosystem functioning concept asserts that processes in ecosystems are markedly influenced by species richness and other facets of biodiversity. However, biodiversity-ecosystem functioning studies have been largely restricted to single ecosystems, ignoring the importance of functional links - such as the exchange of matter, energy, and organisms - between coupled ecosystems. Here we present a basic concept and outline three pathways of cross-boundary biodiversity effects on ecosystem processes and propose an agenda to assess such effects, focusing on terrestrial-aquatic linkages to illustrate the case. This cross-boundary perspective of biodiversity-ecosystem functioning relationships presents a promising frontier for biodiversity and ecosystem science with repercussions for the conservation, restoration, and management of biodiversity and ecosystems from local to landscape scales.

First Nationwide Monitoring Program for the Detection of Potentially Invasive Mosquito Species in Austria.

In Austria, only fragmented information on the occurrence of alien and potentially invasive mosquito species exists. The aim of this study is a nationwide overview on the situation of those mosquitoes in Austria. Using a nationwide uniform protocol for the first time, mosquito eggs were sampled with ovitraps at 45 locations in Austria at weekly intervals from May to October 2020. The sampled eggs were counted and the species were identified by genetic analysis. The Asian tiger mosquito Aedes albopictus was found at two sites, once in Tyrol, where this species has been reported before, and for the first time in the province of Lower Austria, at a motorway rest stop. The Asian bush mosquito Aedes japonicus was widespread in Austria. It was found in all provinces and was the most abundant species in the ovitraps by far. Aedes japonicus was more abundant in the South than in the North and more eggs were found in habitats with artificial surfaces than in (semi-) natural areas. Further, the number of Ae. japonicus eggs increased with higher ambient temperature and decreased with higher wind speed. The results of this study will contribute to a better estimation of the risk of mosquito-borne disease in Austria and will be a useful baseline for a future documentation of changes in the distribution of those species.

Grassland biodiversity.

Grasslands comprise one of Earth's dominant biomes, accounting for up to 40% of its terrestrial area (Figure 1). The fundamental components of grassland habitats are grasses and grass-like plants, but diverse assemblages of other plant life forms and diverse animal communities also contribute to grassland biodiversity. Grasses have evolved traits allowing them to cope with climatic extremes, specific soil conditions, fires, and herbivory, all of which sustain grasslands by limiting the establishment, survival, growth, and dominance of woody vegetation. Grasslands occur in almost all climatic zones, except the poles, extreme arid zones, and the highest mountains (Figure 1). Temperate grassland habitats include Eurasian steppes, North American prairies, the pampas lowlands of South America, and Patagonian steppe. Tropical and subtropical grasslands (savannas) occur mostly in Africa and Australia, but are also found in the north of South America, in the southern United States, South Asia, and Southeast Asia.

Littoral macroinvertebrate communities of alpine lakes along an elevational gradient (Hohe Tauern National Park, Austria).

Alpine lakes support unique communities which may respond with great sensitivity to climate change. Thus, an understanding of the drivers of the structure of communities inhabiting alpine lakes is important to predict potential changes in the future. To this end, we sampled benthic macroinvertebrate communities and measured environmental variables (water temperature, dissolved oxygen, conductivity, pH, nitrate, turbidity, blue-green algal phycocyanin, chlorophyll-a) as well as structural parameters (habitat type, lake size, maximum depth) in 28 lakes within Hohe Tauern National Park, Austria, between altitudes of 2,000 and 2,700 m a.s.l. The most abundant macroinvertebrate taxa that we found were Chironomidae and Oligochaeta. Individuals of Coleoptera, Diptera, Hemiptera, Plecoptera, Trichoptera, Tricladida, Trombidiformes, Veneroida were found across the lakes and determined to family level. Oligochaeta were not determined further. Generalized linear modeling and permanova were used to identify the impact of measured parameters on macroinvertebrate communities. We found that where rocky habitats dominated the lake littoral, total macroinvertebrate abundance and family richness were lower while the ratio of Ephemeroptera, Plecoptera and Trichoptera (EPT) was higher. Zoo- and phytoplankton densities were measured in a subset of lakes but were not closely associated with macroinvertebrate abundance or family richness. With increasing elevation, macroinvertebrate abundances in small and medium-sized lakes increased while they decreased in large lakes, with a clear shift in community composition (based on families). Our results show that habitat parameters (lake size, habitat type) have a major influence on benthic macroinvertebrate community structure whereas elevation itself did not show any significant effects on communities. However, even habitat parameters are likely to change under climate change scenarios (e.g. via increased erosion) and this may affect alpine lake macroinvertebrates.

Biology, chance, or history? The predictable reassembly of temperate grassland communities.

Many studies have examined invasion resistance in plant communities, but few have explored the mechanisms of invasion and how subsequent community reassembly affects community functioning. Using natural dispersal and deliberate seed addition into grassland communities with different compositional and richness histories, we show that invaders establish in a nonrandom manner due to negative effects of resident functional groups on invading species from the same functional group. Invaders hence complement communities with originally low richness levels. Consequently, communities converge toward similar levels of species richness, high functional richness, and evenness, but not always maximum productivity. Invasion processes are faster but qualitatively similar when the effect of chance, in the form of dispersal stochasticity, is reduced by seed addition. Thus, dispersal limitation may influence community assembly, but it does not override functionally predictable assembly mechanisms. Some of the most productive communities prior to invasion are unstable in the face of invasion, leading to decreased productivity following invasion. We suggest that invasion into such communities occurs possibly because a pathogen-free niche is available rather than a resource niche. Thus, pathogens in addition to resource niches may be important biological drivers of community assembly.

Effect of plant species loss on aphid-parasitoid communities.

1. The consequences of species loss on ecosystem functioning within a single trophic level have been extensively studied. However, the loss of basal species is likely to have profound impacts on the abundance, richness and ecosystem functioning of species at higher trophic levels. 2. Here, we used experimentally established plant communities with a species richness gradient to study the effects of plant species loss on a multi-trophic insect community in the field. We measured densities and species richness of aphids and parasitic wasps (primary, secondary and facultative tertiary parasitoids of aphids) that naturally colonized the grassland plots. 3. Furthermore, we calculated two ecosystem functions: aphid load (the number of aphid individuals per host plant biomass used as a proxy for herbivory) and parasitism rate. We used structural equation models to explore pathways of direct and indirect effects of plant species richness on higher trophic levels. 4. We found that the densities and richness of species at all trophic levels were influenced by changes in plant species richness. The effects were rarely direct, but instead mediated by the abundance and species richness of aphid host plants and subsequent trophic levels. 5. The herbivore and primary parasitoid levels were most directly affected by changes in plant species richness, with highest insect densities and species richness occurring at intermediate plant species richness. The densities and species richness of secondary parasitoids declined linearly with plant species richness owing to sparser resources, resulting in shorter food chains in communities with the highest plant species richness. 6. Aphid load was highest at intermediate plant species richness and negatively affected by both host plant biomass and host plant species richness. Parasitism rate was mostly affected indirectly via aphid density and overall only weakly negatively related to plant species richness. 7. Our results demonstrate that plant species richness can have strong cascading effects up to high trophic levels. However, their direction may differ at the lower and higher ends of the plant species richness spectrum, cautioning against simplifying consequences of plant species loss for specialist food webs that may become limited by sparse resources at high plant richness.

Direct and indirect effects of land-use intensity on plant communities across elevation in semi-natural grasslands.

Grassland biodiversity is vulnerable to land use change. How to best manage semi-natural grasslands for maintaining biodiversity is still unclear in many cases because land-use processes may depend on environmental conditions and the indirect effects of land-use on biodiversity mediated by altered abiotic and biotic factors are rarely considered. Here we evaluate the relative importance of the direct and indirect effects of grazing intensity on plant communities along an elevational gradient on a large topographic scale in the Eastern Carpathians in Ukraine. We sampled for two years 31 semi-natural grasslands exposed to cattle grazing. Within each grassland site we measured plant community properties such as the number of species, functional groups, and the proportion of species undesirable for grazing. In addition, we recorded cattle density (as a proxy for grazing intensity), soil properties (bare soil exposure, soil organic carbon, and soil pH) and densities of soil decomposers (earthworms and soil microorganisms). We used structural equation modelling to explore the direct and indirect effects of grazing intensity on plant communities along the elevation gradient. We found that cattle density decreased plant species and functional diversity but increased the proportion of undesirable species. Some of these effects were directly linked to grazing intensity (i.e., species richness), while others (i.e., functional diversity and proportion of undesirable species) were mediated via bare soil exposure. Although grazing intensity decreased with elevation, the effects of grazing on the plant community did not change along the elevation gradient. Generally, elevation had a strong positive direct effect on plant species richness as well as a negative indirect effect, mediated via altered soil acidity and decreased decomposer density. Our results indicate that plant diversity and composition are controlled by the complex interplay among grazing intensity and changing environmental conditions along an elevation gradient. Furthermore, we found lower soil pH, organic carbon and decomposer density with elevation, indicating that the effects of grazing on soil and related ecosystem functions and services in semi-natural grasslands may be more pronounced with elevation. This demonstrates that we need to account for environmental gradients when attempting to generalize effects of land-use intensity on biodiversity.

Direct and indirect effects of forest management on tree-hole inhabiting aquatic organisms and their functional traits.

Ecological communities in forests have been shown to be strongly affected by forest management but a detailed understanding of how different components of management affect insect communities directly and indirectly via environmental variables, how management influences functional trait diversity and composition, and whether these results can be transferred to other functional groups besides insects (e.g. bacteria or nematodes) is still missing. To address these questions we used water-filled tree holes, which provide habitats for insect larvae and other aquatic organisms in forests, as a model system. We mapped all water-filled tree holes in 75 forest plots (1 ha) under different management intensity in three regions of Germany. We measured structural and climatic conditions at different spatial scales, sampled insect communities in 123 tree holes and bacterial and nematode communities in a subset of these. We found that forest management in terms of harvesting intensity and the proportion of non-natural tree species (species not part of the natural vegetation at the sites) negatively affected tree-hole abundance. An increased proportion of non-natural tree species had a positive direct effect on insect richness and functional diversity in the tree holes. However, a structural equation model showed that increasing management intensity had negative indirect effects on insect abundance and richness, operating via environmental variables at stand and tree-hole scale. Functional diversity and trait composition of insect communities similarly responded to changes in management-related variables. In contrast to insects, bacterial and nematode richness were not directly impacted by forest management but by other environmental variables. Our results suggest that forest management may strongly alter insect communities of tree holes, while nematodes and bacteria seem less affected. Most effects in our study were indirect and negative, indicating that management has often complex consequences for forest communities that should be taken into account in forest management schemes.