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Nutrient cycling in terrestrial ecosystems is affected by various factors such as plant diversity and insect herbivory. While several studies suggest insect herbivory to depend on plant diversity, their interacting effect on nutrient cycling is unclear. In a greenhouse experiment with grassland microcosms of one to six plant species of two functional groups (grasses and legumes), we tested the influence of plant species richness (diversity) and functional composition on plant community biomass production, insect foliar herbivory, soil microbial biomass, and nutrient concentrations in throughfall. To manipulate herbivory, zero, three or six generalist grasshoppers ( Chorthippus parallelus) were added to the plant communities. Increasing plant species richness increased shoot biomass and grasshopper performance, without significantly affecting root biomass or insect herbivory. Plant functional composition affected all of these parameters, e.g. legume communities showed the highest shoot biomass, the lowest grasshopper performance and suffered the least herbivory. Nutrient concentrations (dissolved mineral N, PO4-P, SO4-S) and pH in throughfall increased with herbivory. PO4-P and pH increases were positively affected by plant diversity, especially under high herbivore pressure. Plant functional composition affected several throughfall variables, sometimes fully explaining diversity effects. Increasing plant diversity tended to increase soil microbial biomass, but only under high herbivore pressure. Faeces quantities strongly correlated with changes in pH and PO4-P; frass may therefore be an important driver of throughfall pH and a main source of PO4-P released from living plants. Our results indicate that insect herbivory may significantly influence fast nutrient cycling processes in natural communities, particularly so in managed grasslands. [ABSTRACT FROM AUTHOR]

As a rule, communities consist of few abundant and many rare species, which is reflected in the characteristic shape of species abundance distributions (SADs). The processes that shape these SADs have been a longstanding problem for ecological research. Although many studies found strong negative effects of increasing land-use intensity on diversity, few reports consider land-use effects on SADs., Arthropods (insects and spiders) were sampled on 142 grassland plots in three regions in Germany, which were managed with different modes (mowing, fertilization and/or grazing) and intensities of land use. We analysed the effect of land use on three parameters characterizing the shape of SADs: abundance decay rate (the steepness of the rank abundance curve, represented by the niche-preemption model parameter), dominance (Berger-Parker dominance) and rarity (Fisher's alpha). Furthermore, we tested the core-satellite hypothesis by comparing the species' rank within the SAD to their distribution over the land-use gradient., When data on Araneae, Cicadina, Coleoptera, Heteroptera and Orthoptera were combined, abundance decay rate increased with combined land-use intensity (including all modes). Among the single land-use modes, increasing fertilization and grazing intensity increased the decay rate of all taxa, while increasing mowing frequency significantly affected the decay rate only in interaction with fertilization. Results of single taxa differed in their details, but all significant interaction effects included fertilization intensity. Dominance generally increased with increasing fertilization and rarity decreased with increasing grazing or mowing intensity, despite small differences among taxa and regions. The majority of species found on <10% of the plots per region were generally rare (<10 individuals), which is in accordance with the core-satellite hypothesis. We found significant differences in the rarity and dominance of species between plots of low and high intensity for all three land-use modes and for the combined land-use intensity., We conclude that effects of land-use intensity on SADs lead to a stronger dominance of the most abundant species. Furthermore, species which have restricted distributions are more likely to also be rare species in the local SAD and therefore are at high risk of being lost under intensive land use. [ABSTRACT FROM AUTHOR]

Anthropogenic changes in biodiversity and atmospheric temperature significantly influence ecosystem processes. However, little is known about potential interactive effects of plant diversity and warming on essential ecosystem properties, such as soil microbial functions and element cycling. We studied the effects of orthogonal manipulations of plant diversity (one, four, and 16 species) and warming (ambient, +1.5°C, and +3°C) on soil microbial biomass, respiration, growth after nutrient additions, and activities of extracellular enzymes in 2011 and 2012 in the BAC (biodiversity and climate) perennial grassland experiment site at Cedar Creek, Minnesota, USA. Focal enzymes are involved in essential biogeochemical processes of the carbon, nitrogen, and phosphorus cycles. Soil microbial biomass and some enzyme activities involved in the C and N cycle increased significantly with increasing plant diversity in both years. In addition, 16-species mixtures buffered warming induced reductions in topsoil water content. We found no interactive effects of plant diversity and warming on soil microbial biomass and growth rates. However, the activity of several enzymes (1,4-ß-glucosidase, 1,4-ß-N-acetylglucosaminidase, phosphatase, peroxidase) depended on interactions between plant diversity and warming with elevated activities of enzymes involved in the C, N, and P cycles at both high plant diversity and high warming levels. Increasing plant diversity consistently decreased microbial biomass-specific enzyme activities and altered soil microbial growth responses to nutrient additions, indicating that plant diversity changed nutrient limitations and/or microbial community composition. In contrast to our expectations, higher plant diversity only buffered temperature effects on soil water content, but not on microbial functions. Temperature effects on some soil enzymes were greatest at high plant diversity. In total, our results suggest that the fundamental temperature ranges of soil microbial communities may be sufficiently broad to buffer their functioning against changes in temperature and that plant diversity may be a dominant control of soil microbial processes in a changing world. [ABSTRACT FROM AUTHOR]

Biodiversity experiments generally report a positive effect of plant biodiversity on aboveground biomass (overyielding), which typically increases with time. Various studies also found overyielding for belowground plant biomass, but this has never been measured over time. Also, potential underlying mechanisms have remained unclear. Differentiation in rooting patterns among plant species and plant functional groups has been proposed as a main driver of the observed biodiversity effect on belowground biomass, leading to more efficient belowground resource use with increasing diversity, but so far there is little evidence to support this. We analyzed standing root biomass and its distribution over the soil profile, along a 1-16 species richness gradient over eight years in the Jena Experiment in Germany, and compared belowground to aboveground overyielding. In our long-term dataset, total root biomass increased with increasing species richness but this effect was only apparent after four years. The increasingly positive relationship between species richness and root biomass, explaining 12% of overall variation and up to 28% in the last year of our study, was mainly due to decreasing root biomass at low diversity over time. Functional group composition strongly affected total standing root biomass, explaining 44% of variation, with grasses and legumes having strong overall positive and negative effects, respectively. Functional group richness or interactions between functional group presences did not strongly contribute to overyielding. We found no support for the hypothesis that vertical root differentiation increases with species richness, with functional group richness or composition. Other explanations, such as stronger negative plant-soil feedbacks in low-diverse plant communities on standing root biomass and vertical distribution should be considered. [ABSTRACT FROM AUTHOR]

Many studies have examined whether communities are structured by random or deterministic processes, and both are likely to play a role, but relatively few studies have attempted to quantify the degree of randomness in species composition. We quantified, for the first time, the degree of randomness in forest bird communities based on an analysis of spatial autocorrelation in three regions of Germany. The compositional dissimilarity between pairs of forest patches was regressed against the distance between them. We then calculated the y-intercept of the curve, i.e. the ‘nugget’, which represents the compositional dissimilarity at zero spatial distance. We therefore assume, following similar work on plant communities, that this represents the degree of randomness in species composition. We then analysed how the degree of randomness in community composition varied over time and with forest management intensity, which we expected to reduce the importance of random processes by increasing the strength of environmental drivers. We found that a high portion of the bird community composition could be explained by chance (overall mean of 0.63), implying that most of the variation in local bird community composition is driven by stochastic processes. Forest management intensity did not consistently affect the mean degree of randomness in community composition, perhaps because the bird communities were relatively insensitive to management intensity. We found a high temporal variation in the degree of randomness, which may indicate temporal variation in assembly processes and in the importance of key environmental drivers. We conclude that the degree of randomness in community composition should be considered in bird community studies, and the high values we find may indicate that bird community composition is relatively hard to predict at the regional scale. [ABSTRACT FROM AUTHOR]

Loss of plant diversity influences essential ecosystem processes as aboveground productivity, and can have cascading effects on the arthropod communities in adjacent trophic levels. However, few studies have examined how those changes in arthropod communities can have additional impacts on ecosystem processes caused by them (e.g. pollination, bioturbation, predation, decomposition, herbivory). Therefore, including arthropod effects in predictions of the impact of plant diversity loss on such ecosystem processes is an important but little studied piece of information. In a grassland biodiversity experiment, we addressed this gap by assessing aboveground decomposer and herbivore communities and linking their abundance and diversity to rates of decomposition and herbivory. Path analyses showed that increasing plant diversity led to higher abundance and diversity of decomposing arthropods through higher plant biomass. Higher species richness of decomposers, in turn, enhanced decomposition. Similarly, species-rich plant communities hosted a higher abundance and diversity of herbivores through elevated plant biomass and C:N ratio, leading to higher herbivory rates. Integrating trophic interactions into the study of biodiversity effects is required to understand the multiple pathways by which biodiversity affects ecosystem functioning. [ABSTRACT FROM AUTHOR]

Intensive land use is a driving force for biodiversity decline in many ecosystems. In semi-natural grasslands, land-use activities such as mowing, grazing and fertilization affect the diversity of plants and arthropods, but the combined effects of different drivers and the chain of effects are largely unknown. In this study we used structural equation modelling to analyse how the arthropod communities in managed grasslands respond to land use and whether these responses are mediated through changes in resource diversity or resource quantity (biomass). Plants were considered resources for herbivores which themselves were considered resources for predators. Plant and arthropod (herbivores and predators) communities were sampled on 141 meadows, pastures and mown pastures within three regions in Germany in 2008 and 2009. Increasing land-use intensity generally increased plant biomass and decreased plant diversity, mainly through increasing fertilization. Herbivore diversity decreased together with plant diversity but showed no response to changes in plant biomass. Hence, land-use effects on herbivore diversity were mediated through resource diversity rather than quantity. Land-use effects on predator diversity were mediated by both herbivore diversity (resource diversity) and herbivore quantity (herbivore biomass), but indirect effects through resource quantity were stronger. Our findings highlight the importance of assessing both direct and indirect effects of land-use intensity and mode on different trophic levels. In addition to the overall effects, there were subtle differences between the different regions, pointing to the importance of regional land-use specificities. Our study underlines the commonly observed strong effect of grassland land use on biodiversity. It also highlights that mechanistic approaches help us to understand how different land-use modes affect biodiversity. [ABSTRACT FROM AUTHOR]

Forest management not only affects biodiversity but also might alter ecosystem processes mediated by the organisms, i.e. herbivory the removal of plant biomass by plant-eating insects and other arthropod groups. Aiming at revealing general relationships between forest management and herbivory we investigated aboveground arthropod herbivory in 105 plots dominated by European beech in three different regions in Germany in the sun-exposed canopy of mature beech trees and on beech saplings in the understorey. We separately assessed damage by different guilds of herbivores, i.e. chewing, sucking and scraping herbivores, gall-forming insects and mites, and leaf-mining insects. We asked whether herbivory differs among different forest management regimes (unmanaged, uneven-aged managed, even-aged managed) and among age-classes within even-aged forests. We further tested for consistency of relationships between regions, strata and herbivore guilds. On average, almost 80% of beech leaves showed herbivory damage, and about 6% of leaf area was consumed. Chewing damage was most common, whereas leaf sucking and scraping damage were very rare. Damage was generally greater in the canopy than in the understorey, in particular for chewing and scraping damage, and the occurrence of mines. There was little difference in herbivory among differently managed forests and the effects of management on damage differed among regions, strata and damage types. Covariates such as wood volume, tree density and plant diversity weakly influenced herbivory, and effects differed between herbivory types. We conclude that despite of the relatively low number of species attacking beech; arthropod herbivory on beech is generally high. We further conclude that responses of herbivory to forest management are multifaceted and environmental factors such as forest structure variables affecting in particular microclimatic conditions are more likely to explain the variability in herbivory among beech forest plots. [ABSTRACT FROM AUTHOR]

Forestry in Europe changed the tree species composition and reduced dead-wood amount and heterogeneity, and therefore negatively affected saproxylic diversity. Efficient conservation requires knowledge about the importance of the relevant diversity drivers across taxa. We examined the relative importance of space vs. host for saproxylic diversity at a spatial extend of 600 km in Germany. Further, we disentangled effects of among regions, forest stands, host clades, and tree species on saproxylic diversity. This allows inferences for spatial- and host tree-related conservation strategies. Beetle, fungal sporocarp, molecular-derived fungal, and bacterial communities were studied in a large nested dead-wood experiment comprising 11 tree species. We used multiplicative diversity partitioning to assess the diversity of rare, typical, and dominant species. The beta-diversity of beetles and fungal sporocarps was equally explained by space and host, but that of molecular fungi and bacteria mainly by the host. Across taxa, beta-diversity was higher among forest stands than among regions. However, for beetles and fungal sporocarps, differences among regions were also important. Host tree clade and host tree species were important for beetle and host clade for fungal sporocarp beta-diversity. Host tree species was more important than host clade for the beta-diversity of molecular fungi and bacteria. The divergent response of different taxa to space and host calls into question the use of a simple spatially-centered or host-centered strategy. Instead, a high dead-wood tree species diversity on a broad spatial coverage at the national scale in temperate European forests is necessary to maintain rare and abundant species. • Relative importance of space versus host across saproxylic taxa is unknown. • Beetle diversity is driven by space and host, microbial diversity mainly by the host. • Different fungal diversity measures yielded different results. • Conservation requires dead-wood enrichment with different clades and tree species. • Dead-wood enrichment must be established in different forest types across regions. [ABSTRACT FROM AUTHOR]

Biodiverse communities have been shown to sustain high levels of multifunctionality and thus a loss of species likely negatively impacts ecosystem functions. For most taxa, however, roles of individual species are poorly known. Rare species, often most likely to go extinct, may have unique traits and functional roles. Alternatively, rare species may be functionally redundant, such that their loss would not disrupt ecosystem functions. We quantified the functional role of rare species by using captures of wood-living (saproxylic) beetle species, combined with recent databases of morphological and ecological traits, from three regions in central and northern Europe. Using a rarity index based on species' local abundance, geographic range, and habitat breadth, we used local and regional species removal simulations to examine the contributions of both the rarest and the most common beetle species to three measures of community functional structure: functional richness, functional specialization, and functional originality. Both regionally and locally, all three measures declined more rapidly when rare species were removed than under common (or random) species removal scenarios. These consistent patterns across scales and among forest types give evidence that rare species provide unique functional contributions, and their loss may disproportionately impact ecosystem functions. This implies that conservation measures targeting rare and endangered species, such as preserving intact forests with dead wood and mature trees, can provide broader ecosystem-level benefits. Experimental research linking functional structure to ecosystem processes should be prioritized to increase understanding of the functional consequences of species loss and to develop more effective conservation strategies. [ABSTRACT FROM AUTHOR]

Plant-herbivore interactions are influenced by host plant quality which in turn is affected by plant growth conditions. Competition is the major biotic and nutrient availability a major abiotic component of a plant’s growth environment. Yet, surprisingly few studies have investigated impacts of competition and nutrient availability on herbivore performance and reciprocal herbivore effects on plants. We studied growth of the specialist aphid, Macrosiphoniella tanacetaria, and its host plant tansy, Tanacetum vulgare, under experimental addition of inorganic and organic fertilizer crossed with competition by goldenrod, Solidago canadensis. Because of evidence that competition by goldenrod is mediated by allelopathic compounds, we also added a treatment with activated carbon. Results showed that fertilization increased, and competition with goldenrod decreased, plant biomass, but this was likely mediated by resource competition. There was no evidence from the activated carbon treatment that allelopathy played a role which instead had a fertilizing effect. Aphid performance increased with higher plant biomass and depended on plant growth conditions, with fertilization and AC increasing, and plant competition decreasing aphid numbers. Feedbacks of aphids on plant performance interacted with plant growth conditions in complex ways depending on the relative magnitude of the effects on plant biomass and aphid numbers. In the basic fertilization treatment, tansy plants profited from increased nutrient availability by accumulating more biomass than they lost due to an increased number of aphids under fertilization. When adding additional fertilizer, aphid numbers increased so high that tansy plants suffered and showed reduced biomass compared with controls without aphids. Thus, the ecological cost of an infestation with aphids depends on the balance of effects of growth conditions on plant and herbivore performance. These results emphasize the importance to investigate both perspectives in plant herbivore interactions and characterize the effects of growth conditions on plant and herbivore performance and their respective feedbacks. [ABSTRACT FROM AUTHOR]

Background: Infiltration is a key process in determining the water balance, but so far effects of earthworms, soil texture, plant species diversity and their interaction on infiltration capacity have not been studied. Methodology/Principal Findings: We measured infiltration capacity in subplots with ambient and reduced earthworm density nested in plots of different plant species (1, 4, and 16 species) and plant functional group richness and composition (1 to 4 groups; legumes, grasses, small herbs, tall herbs). In summer, earthworm presence significantly increased infiltration, whereas in fall effects of grasses and legumes on infiltration were due to plant-mediated changes in earthworm biomass. Effects of grasses and legumes on infiltration even reversed effects of texture. We propose two pathways: (i) direct, probably by modifying the pore spectrum and (ii) indirect, by enhancing or suppressing earthworm biomass, which in turn influenced infiltration capacity due to change in burrowing activity of earthworms. Conclusions/Significance: Overall, the results suggest that spatial and temporal variations in soil hydraulic properties can be explained by biotic processes, especially the presence of certain plant functional groups affecting earthworm biomass, while soil texture had no significant effect. Therefore biotic parameters should be taken into account in hydrological applications. [ABSTRACT FROM AUTHOR]

Highlights: [•] We tested if spatial diversity patterns correlate among different organismic groups. [•] Generally, β-diversity revealed to contribute much more to γ-diversity than α-diversity. [•] β-Diversity correlated more strongly between trophic levels than α-diversity. [•] More strongly connected trophic levels revealed not generally stronger relationships. [•] Heterogeneity of beech forests and differences among trophic guilds must be considered in conservation strategies. [Copyright &y& Elsevier]

Highlights: [•] Organic management is able to promote arthropod biodiversity in grasslands. [•] No trend towards higher plant diversity was found in organic grasslands. [•] Organic farming does not bring severe economical losses in yield or soil fertility. [•] No differences in the effect of organic farming among grassland types were found. [ABSTRACT FROM AUTHOR]

Numerous studies have reported positive effects of species richness on plant community productivity. Such biodiversity effects are usually quantified by comparing the performance of plant mixtures with reference monocultures. However, several mechanisms, such as the lack of resource complementarity and facilitation or the accumulation of detrimental agents, suggest that monocultures are more likely than mixtures to deteriorate over time. Increasing biodiversity effects over time could therefore result from declining monocultures instead of reflecting increases in the functioning of mixtures. Commonly, the latter is assumed when positive trends in biodiversity effects occur. Here, we analysed the performance of 60 grassland species growing in monocultures and mixtures over 9 years in a biodiversity experiment to clarify whether their temporal biomass dynamics differed and whether a potential decline of monocultures contributed significantly to the positive net biodiversity effect observed. Surprisingly, individual species’ populations produced, on average, significantly more biomass per unit area when growing in monoculture than when growing in mixture. Over time, productivity of species decreased at a rate that was, on average, slightly more negative in monocultures than in mixtures. The mean net biodiversity effect across all mixtures was continuously positive and ranged between 64–217 g per m2. Short-term increases in the mean net biodiversity effect were only partly due to deteriorating monocultures and were strongly affected by particular species gaining dominance in mixtures in the respective years. We conclude that our species performed, on average, comparably in monocultures and mixtures; monoculture populations being slightly more productive than mixture populations but this trend decreased over time. This suggested that negative feedbacks had not yet affected monocultures strongly but could potentially become more evident in the future. Positive biodiversity effects on aboveground productivity were heavily driven by a small, but changing, set of species that behaved differently from the average species. [ABSTRACT FROM AUTHOR]

Recent studies assessing the role of biological diversity for ecosystem functioning indicate that the diversity of functional traits and the evolutionary history of species in a community, not the number of taxonomic units, ultimately drives the biodiversity-ecosystem-function relationship. Here, we simultaneously assessed the importance of plant functional trait and phylogenetic diversity as predictors of major trophic groups of soil biota (abundance and diversity), six years from the onset of a grassland biodiversity experiment. Plant functional and phylogenetic diversity were generally better predictors of soil biota than the traditionally used species or functional group richness. Functional diversity was a reliable predictor for most biota, with the exception of soil microorganisms, which were better predicted by phylogenetic diversity. These results provide empirical support for the idea that the diversity of plant functional traits and the diversity of evolutionary lineages in a community are important for maintaining higher abundances and diversity of soil communities. [ABSTRACT FROM AUTHOR]

1: Given the predictions of increased drought probabilities under various climate change scenarios, there have been numerous experimental field studies simulating drought using transparent roofs in different ecosystems and regions. Such roofs may, however, have unknown side effects, called artifacts, on the measured variables potentially confounding the experimental results. A roofed control allows the quantification of potential artifacts, which is lacking in most experiments. 2: We conducted a drought experiment in experimental grasslands to study artifacts of transparent roofs and the resulting effects of artifacts on ecosystems relative to drought on three response variables (aboveground biomass, litter decomposition and plant metabolite profiles). We established three drought treatments, using (1) transparent roofs to exclude rainfall, (2) an unroofed control treatment receiving natural rainfall and (3) a roofed control, nested in the drought treatment but with rain water reapplied according to ambient conditions. 3: Roofs had a slight impact on air (+0.14°C during night) and soil temperatures (−0.45°C on warm days, +0.25°C on cold nights), while photosynthetically active radiation was decreased significantly (−16%). Aboveground plant community biomass was reduced in the drought treatment (−41%), but there was no significant difference between the roofed and unroofed control, i.e., there were no measurable roof artifact effects. 4: Compared to the unroofed control, litter decomposition was decreased significantly both in the drought treatment (−26%) and in the roofed control treatment (−18%), suggesting artifact effects of the transparent roofs. Moreover, aboveground metabolite profiles in the model plant species Medicago x varia were different from the unroofed control in both the drought and roofed control treatments, and roof artifact effects were of comparable magnitude as drought effects. 5: Our results stress the need for roofed control treatments when using transparent roofs for studying drought effects, because roofs can cause significant side effects. [ABSTRACT FROM AUTHOR]

Understanding how land use and management practices affect biodiversity is essential for developing effective restoration and conservation strategies. Here, we used ant communities to evaluate the effects of historical land uses (former agriculture or tree plantation) and current management (grazing, burning, mowing, and fertilizing) in subtropical grasslands. We also examined how ant species richness and composition are affected through changes in resource diversity and habitat structure. We classified 80 study sites in southern Brazil into two groups: permanent and secondary grasslands. Permanent grasslands presented high, medium or low management intensities that had never been under other land use. Secondary grasslands are former grasslands that were used for agriculture or pine plantations in the past, followed by passive restoration. We sampled ant communities with D-Vac suction in three multiplots per site in 2013–2015. We found that secondary grasslands had lower ant species diversity and different community composition than permanent grasslands. Ant diversity did not differ among permanent grasslands, but composition was markedly different, especially when comparing medium and high management intensities with low. Additionally, ant richness in secondary grasslands was mainly mediated by a decreased resource diversity (i.e., plant species richness) and grass cover, while burning and mowing management increased ant richness by increasing resource diversity. Our results indicate that passive recovery of grasslands in early successional stages after other land uses leads to incomplete recovery of ant communities, as important resource and structure-related drivers of ant species remain altered. In contrast, traditional management practices promote ant diversity. • Historical and current land use shape ant communities in subtropical grasslands. • Secondary grasslands have reduced ant diversity and altered species composition. • Management intensity did not affect ant richness but drive species composition. • Resource diversity and grass cover loss are the main drivers of ant community changes. • Ants can be used as indicators in conservation and restoration monitoring in subtropical grasslands. [ABSTRACT FROM AUTHOR]

Plant functional traits affect the capacity of herbivores to find, choose, and consume plants. However, in a community composed of different plant species, it is unclear what proportion of herbivory on a focal plant is explained by its own traits and which is explained by the characteristics of the surrounding vegetation (i.e., nonadditive effects). Moreover, nonadditive effects could be positive or negative, and it is not known if they are related to community properties such as diversity. To quantify nonadditive effects, we developed four different additive models based on monoculture herbivory rates or plant traits and combined them with measurements of standing invertebrate herbivore damage along an experimental plant diversity gradient ranging from monocultures to 60-species mixtures. In all four models, positive nonadditive effects were detected, i.e., herbivory levels were higher in polycultures than what was expected from monoculture data, and these effects contributed up to 25% of the observed variance in herbivory. Importantly, the nonadditive effects, which were defined as the deviance of the models' predictions from the observed herbivory, were positively correlated with the communities' plant species richness. Consequently, interspecific interactions appear to have an important impact on the levels of herbivory of a community. Identifying those community properties that capture the effects of these interactions is a next important challenge for our understanding of how the environment interacts with plant traits to drive levels of herbivory. [ABSTRACT FROM AUTHOR]

With the aim of wood production with negligible negative effects on biodiversity and ecosystem processes, a silvicultural practice of selective logging with natural regeneration has been implemented in European beech forests (Fagus sylvatica) during the last decades. Despite this near-to-nature strategy, species richness of various taxa is lower in these forests than in unmanaged forests. To develop guidelines to minimize the fundamental weaknesses in the current practice, we linked functional traits of saproxylic beetle species to ecosystem characteristics. We used continental-scale data from 8 European countries and regional-scale data from a large forest in southern Germany and forest-stand variables that represented a gradient of intensity of forest use to evaluate the effect of current near-to-nature management strategies on the functional diversity of saproxylic beetles. Forest-stand variables did not have a statistically significant effect on overall functional diversity, but they did significantly affect community mean and diversity of single functional traits. As the amount of dead wood increased the composition of assemblages shifted toward dominance of larger species and species preferring dead wood of large diameter and in advanced stages of decay. The mean amount of dead wood across plots in which most species occurred was from 20 to 60 m3/ha. Species occurring in plots with mean dead wood >60 m3/ha were consistently those inhabiting dead wood of large diameter and in advanced stages of decay. On the basis of our results, to make current wood-production practices in beech forests throughout Europe more conservation oriented (i.e., promoting biodiversity and ecosystem functioning), we recommend increasing the amount of dead wood to >20 m3/ha; not removing dead wood of large diameter (50 cm) and allowing more dead wood in advanced stages of decomposition to develop; and designating strict forest reserves, with their exceptionally high amounts of dead wood, that would serve as refuges for and sources of saproxylic habitat specialists. Efectos Actuales del Manejo Casi Natural de Bosques sobre la Composición de Atributos Funcionales de Escarabajos Saproxílicos en Bosques de Haya [ABSTRACT FROM AUTHOR]

In the course of the biodiversity-ecosystem functioning debate, the issue of multifunctionality of species communities has recently become a major focus. Elemental stoichiometry is related to a variety of processes reflecting multiple plant responses to the biotic and abiotic environment. It can thus be expected that the diversity of a plant assemblage alters community level plant tissue chemistry. We explored elemental stoichiometry in aboveground plant tissue (ratios of carbon, nitrogen, phosphorus, and potassium) and its relationship to plant diversity in a 5-year study in a large grassland biodiversity experiment (Jena Experiment). Species richness and functional group richness affected community stoichiometry, especially by increasing C:P and N:P ratios. The primacy of either species or functional group richness effects depended on the sequence of testing these terms, indicating that both aspects of richness were congruent and complementary to expected strong effects of legume presence and grass presence on plant chemical composition. Legumes and grasses had antagonistic effects on C:N (−27.7% in the presence of legumes, +32.7% in the presence of grasses). In addition to diversity effects on mean ratios, higher species richness consistently decreased the variance of chemical composition for all elemental ratios. The diversity effects on plant stoichiometry has several non-exclusive explanations: The reduction in variance can reflect a statistical averaging effect of species with different chemical composition or a optimization of nutrient uptake at high diversity, leading to converging ratios at high diversity. The shifts in mean ratios potentially reflect higher allocation to stem tissue as plants grew taller at higher richness. By showing a first link between plant diversity and stoichiometry in a multiyear experiment, our results indicate that losing plant species from grassland ecosystems will lead to less reliable chemical composition of forage for herbivorous consumers and belowground litter input. [ABSTRACT FROM AUTHOR]

The importance of competition between similar species in driving community assembly is much debated. Recently, phylogenetic patterns in species composition have been investigated to help resolve this question: phylogenetic clustering is taken to imply environmental filtering, and phylogenetic overdispersion to indicate limiting similarity between species. We used experimental plant communities with random species compositions and initially even abundance distributions to examine the development of phylogenetic pattern in species abundance distributions. Where composition was held constant by weeding, abundance distributions became overdispersed through time, but only in communities that contained distantly related clades, some with several species (i.e., a mix of closely and distantly related species). Phylogenetic pattern in composition therefore constrained the development of overdispersed abundance distributions, and this might indicate limiting similarity between close relatives and facilitation/complementarity between distant relatives. Comparing the phylogenetic patterns in these communities with those expected from the monoculture abundances of the constituent species revealed that interspecific competition caused the phylogenetic patterns. Opening experimental communities to colonization by all species in the species pool led to convergence in phylogenetic diversity. At convergence, communities were composed of several distantly related but species-rich clades and had overdispersed abundance distributions. This suggests that limiting similarity processes determine which species dominate a community but not which species occur in a community. Crucially, as our study was carried out in experimental communities, we could rule out local evolutionary or dispersal explanations for the patterns and identify ecological processes as the driving force, underlining the advantages of studying these processes in experimental communities. Our results show that phylogenetic relations between species provide a good guide to understanding community structure and add a new perspective to the evidence that niche complementarity is critical in driving community assembly. [ABSTRACT FROM AUTHOR]

Invertebrate herbivores can impact plant performance and plant communities. Conversely, plants can affect the ability of herbivores to find, choose, and consume them through their functional traits. While single plant traits have been related to rates of herbivory, most often involving single herbivore-plant pairs, much less is known about which suite of plant traits is important for determining herbivory for a pool of plant species interacting with a natural herbivore community. In this study we measured aboveground herbivore damage on 51 herbaceous species growing in monocultures of a grassland biodiversity experiment and collected 42 different plant traits representing four trait groups: physiological, morphological, phenological, and herbivore related. Using the method of random forests and multiple regression, we identified seven traits that are important predictors of herbivore damage (leaf nitrogen and lignin concentration, number of coleopteran and hemipteran herbivores potentially feeding on the plants, leaf life span, stem growth form, and root architecture); leaf nitrogen and lignin concentration were the two most important predictors. The final model accounted for 63% of the variation in herbivore damage. Traits from all four trait groups were selected, showing that a variety of plant characteristics can be statistically important when assessing folivory, including root traits. Our results emphasize that it is necessary to use a multivariate approach for identifying traits affecting complex ecological processes such as herbivory. [ABSTRACT FROM AUTHOR]

In soil, Acidobacteria constitute on average 20% of all bacteria, are highly diverse, and are physiologically active in situ. However, their individual functions and interactions with higher taxa in soil are still unknown. Here, potential effects of land use, soil properties, plant diversity, and soil nanofauna on acidobacterial community composition were studied by cultivation-independent methods in grassland and forest soils from three different regions in Germany. The analysis of l6S rRNA gene clone libraries representing all studied soils revealed that grassland soils were dominated by subgroup Gp6 and forest soils by subgroup GpI Acidobacteria. The analysis of a large number of sites (n = 57) by 6S rRNA gene fingerprinting methods (terminal restriction fragment length polymorphism (T-RFLP] and denaturing gradient gel electrophoresis [DGGE]) showed that Acidobactersa diversities differed between grassland and forest soils but also among the three different regions. Edaphic properties, such as PH, organic carbon, total nitrogen, C/N ratio, phosphorus, nitrate, ammonium, soil moisture, soil temperature, and soil respiration, had an impact on community composition as assessed by fingerprinting. However, interrelations with environmental parameters among subgroup terminal restriction fragments (T-RPs) differed significantly, e.g., different Gpl T-RF5 correlated positively or negatively with nitrogen content. Novel significant correlations of Acidobacteria subpopulations (i.e., individual populations within subgroups) with soil nanofauna and vascular plant diversity were revealed only by analysis of clone sequences. Thus, for detecting novel interrelations of environmental parameters with Acidobacteria, individual populations within subgroups have to be considered. [ABSTRACT FROM AUTHOR]

Abstract: European farmland bird populations have decreased dramatically in recent decades and agricultural intensification has been identified as the main cause contributing to these declines. Identifying which specific intensification pressures are driving those population trends seems vital for bird conservation in European farmland. We investigated the response of ground-nesting farmland birds to the multivariate process of agricultural intensification in six European countries covering a bio-geographical and intensification gradient. Supported by PCA analysis, two groups of factors, related to field management and landscape modification, were considered, seeking to discriminate the relative importance of the effects of these main intensification components. Variance partition analysis showed that landscape factors accounted for most of the variation of ground-nesting farmland bird individual and breeding pair densities, as well as Skylark (i.e. our single model species) individual densities. In the case of Skylark breeders, field factors were found to be more important to explain their density. Our results suggest that in general farmland bird densities as well as Skylark densities are higher in simple landscapes dominated by agriculture, but with smaller fields and more different crops on the farms. In addition, high yields were negatively related to bird densities. We conclude that while management actions aimed at farmland bird conservation taken at landscape level may exert a strong positive effect on overall bird densities, those taken at field level are also relevant, particularly for breeders and, therefore, may potentially influence the persistence of these species’ populations. [Copyright &y& Elsevier]

Abstract: Agricultural intensification is a major driver of global environmental change. Disentangling the relative impact of losses in plant species richness and intensified management on higher trophic level organisms is important for conservation recommendations. We established different management regimes along an experimental gradient of plant species richness within “The Jena-Experiment” in Germany and quantified herbivory as well as grasshoppers and pollinators. Herbivory, grasshopper density and species richness and frequency of flower visiting pollinators were recorded two times in each of four subplots of altogether 80 plots differing in plant species richness. Each of the four subplots was subject to four different levels of fertilizer application and mowing to simulate very low, low, high and very high land use intensity. Fertilization and mowing significantly affected plant–herbivore interactions but plant species richness had no discernible effect. Grasshoppers were most abundant at high intensity subplots (3.1 individuals perm2) and least abundant on very low intensity subplots (1.3 individuals perm2). Leaf damage caused by herbivores was highest in even the very high intensity subplots (3.7%) with four mowing events per year and high amounts of fertilizer application and lowest on subplots with a low management intensity (2.4%) comprising two mowing events per year but no fertilizer application. In contrast, pollinators benefited most from lower management intensities, with only one or two mowing events and no fertilizer addition. In addition, higher numbers of flowering plant species and increased blossom cover was associated with enhanced pollinator species richness and flower visitation. Our results indicate that even in grasslands with high plant species richness, mowing and fertilization are more important drivers of herbivory and flower visitation by pollinators. Management with no more than two cuts per year and without fertilizer application in our grasslands balanced the ecosystem functions of increased pollination and decreased herbivory. [Copyright &y& Elsevier]

1. Aphid natural enemies include not only predators and parasitoids but also pathogens, of which fungi are the most studied for biological control. While wing formation in aphids is induced by abiotic conditions, it is also affected by biotic interactions with their arthropod natural enemies. Wing induction via interactions with arthropod natural enemies is mediated by the increase in their physical contact when alarmed (pseudo-crowding). Pathogenic fungi do not trigger this alarm behaviour in aphids and, therefore, no pseudo-crowding occurs. 2. We hypothesise that, while pathogenic fungi will stimulate maternally induced wing formation, the mechanism is different and is influenced by pathogen specificity. We tested this hypothesis using two entomopathogenic fungi, Pandora neoaphidis and Beauveria bassiana, an aphid specialist and a generalist respectively, on the pea aphid, Acyrthosiphon pisum Harris. 3. We first demonstrate that pea aphids infected with either pathogen and maintained in groups on broad bean plants produced a higher proportion of winged morphs than uninfected control aphids. We then show that, when maintained in isolation, aphids infected with either pathogen also produced higher proportions of winged offspring than control aphids. There was no difference between P. neoaphidis and B. bassiana in their effects on wing induction in either experiment. 4. Unlike the effect of predators and parasitoids on pea aphid wing induction, the effect of pathogens is independent of physical contact with other aphids, suggesting that physiological cues induce wing formation in infected aphids. It is possible that aphids benefit from wing induction by escaping infected patches whilst pathogens may benefit through dispersion. Possible mechanisms of wing induction are discussed. [ABSTRACT FROM AUTHOR]

Seed dispersal by ants (myrmecochory) is widespread, and seed adaptations to myrmecochory are common, especially in the form of fatty appendices (elaiosomes). In a recent study, slugs were identified as seed dispersers of myrmecochores in a central European beech forest. Here we used 105 beech forest sites to test whether myrmecochore presence and abundance is related to ant or gastropod abundance and whether experimentally exposed seeds are removed by gastropods. Myrmecochorous plant cover was positively related to gastropod abundance but was negatively related to ant abundance. Gastropods were responsible for most seed removal and elaiosome damage, whereas insects (and rodents) played minor roles. These gastropod effects on seeds were independent of region or forest management. We suggest that terrestrial gastropods can generally act as seed dispersers of myrmecochorous plants and even substitute myrmecochory, especially where ants are absent or uncommon. [ABSTRACT FROM AUTHOR]

Two tansy-feeding aphids, Macrosiphoniella tanacetaria (MA) and Metopeurum fuscoviride (ME), were studied at a small spatial scale in and around Jena (< 80 km2) using polymorphic microsatellite markers. Both species were found in approximately 60% of sites formerly known to harbour the aphids, although, generally when they did occur, they occurred singly (MA ∼50%; ME ∼60%) and rarely together on the same plant at the same time (approximately 10%) and then usually only in the early part of the growing season. This difference may be a result of quasi-apparent competition effects elicited by ants farming ME aphids, and preferentially actively eliminating or disturbing MA aphids. In terms of population genetics, both aphids showed extreme genetic heterogeneity within a metapopulation structure, with ME more than MA (i.e. higher FST values, approximately 0.4 versus 0.15, respectively), and limited levels of interpopulation gene flow. Subpopulations often deviated from Hardy-Weinberg equilibrium and showed linkage disequilibria, as expected in animals with extended parthenogenetic reproduction, and had positive FIS values for most large samples, suggesting inbreeding, and possibly philopatry, certainly in ME. Hierarchical analysis (allele range and number per locus, analysis of molecular variance and FST) strongly suggested that the plant rather than site governs the level of genetic variation. Bayesian clustering analysis revealed that both species had heterogeneous historical genetic patterning, with K (number of subgroups) in the range 3-7. Evidence is also provided from isolation-by-distance and private allele analyses indicating that, in MA, the presence of winged autumn males, absent in ME where males are wingless, influences comparative population genetic structuring, such that ME subpopulations are comparatively more inbred and genetically differentiated than MA subpopulations. Lastly, additional spatial arrangement (ALLELES-IN-SPACE) analysis showed that, in both species, certain subpopulations were genetically isolated from the remainder, probably as a result of geographical barriers, including intervening buildings and woods. As such, the biology of these tansy aphids living in semi-natural habitats is very different from many pest aphid species examined within agro-ecosystems and infesting ephemeral crops. This is because the former appear to be much more reluctant to fly and hence show contrastingly much higher levels of interpopulation divergence, even at small spatial scales as investigated in the present study. Indeed, the number of genotypic clusters found for tansy aphids using Bayesian approaches is similar to that globally for the major pest, the peach-potato aphid, Myzus persicae. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104, 838-865. [ABSTRACT FROM AUTHOR]

Abstract: Plant diversity was shown to influence the N cycle, but plant diversity effects on other nutrients remain unclear. We tested whether plant species richness or the presence/absence of particular functional plant groups influences P partitioning among differently extractable pools in soil, P concentrations in soil solution, and exploitation of P resources (i.e. the proportion of total bioavailable P in plants and soil that was stored in aboveground biomass) by the plant community in a 5-year biodiversity experiment in grassland. The experimental grassland site established in 2002 had 82 plots with different combinations of numbers of species (1, 2, 4, 8, 16, 60) and functional groups (grasses, small non-leguminous herbs, tall non-leguminous herbs, legumes). In 2007, we determined P partitioning (Hedley) in soil of all experimental plots. We sampled plant community biomass and continuously extracted soil solution with suction plates from March 2003 to February 2007 and determined PO4-P concentrations in all samples. The presence of legumes increased aboveground P storage in plants and decreased labile Pi concentrations in soil because of their higher demands for P associated with N2 fixation. During cold periods, readily plant-available PO4-P concentrations in soil solution increased in legume-containing mixtures likely caused by leaching from P-rich residues. We found a consistently positive effect of plant species richness on P exploitation by the plant community which was independent of the presence of particular plant functional groups. With proceeding time after establishment, plant species richness increasingly contributed to the explanation of the variance in P exploitation. Therefore, plant strategies to efficiently acquire P seem to become increasingly important in these grasslands. We conclude that diverse plant communities are better prepared than less diverse mixtures to respond to P limitation induced by continuously high atmospheric N deposition. [Copyright &y& Elsevier]

Summary 1. While positive effects of biodiversity on temporal stability of communities have been demonstrated in theoretical and empirical studies, diversity-stability relationships at the population level remain poorly understood. 2. We investigated temporal variability of plant populations in experimental grassland plots of varying species richness (1, 2, 4, 8, 16-60 species), functional group richness and composition (presence/absence of legumes × grasses × small herbs × tall herbs) in a long-term biodiversity experiment from 2003 to 2009 ('Jena Experiment'). 3. Average population stability, defined as the reciprocal of the coefficient of variation of above-ground biomass production over time, differed largely between species but was generally higher in grasses and small herbs than in legumes and tall herbs. Furthermore, population stability was positively related to a species' proportional contribution to community biomass. Thus, an increasing number of subordinate species explained lower average population stabilities at higher diversity levels. 4. A negative covariance (CV) across all species-richness levels suggested negatively correlated species dynamics. Species belonging to different functional groups fluctuated asynchronously, while species dynamics within functional groups were more synchronous. Community-wide species synchrony decreased with increasing species richness, and temporal stability at the community level increased. 5. Synthesis: Our results suggest that diversity-stability relationships are driven by fluctuations in the population biomass of individual species which are less synchronized in more diverse than in less diverse mixtures and monocultures. Dominant plant species tend to be more stabilized than subdominant species, independently of community species richness. However, asynchrony of population dynamics outweighs decreasing population stability with increasing species richness, resulting in higher temporal stability at the plant community level. [ABSTRACT FROM AUTHOR]

Forest canopies are an important part of forest ecosystems and comprise the majority of arthropod diversity. As tree crowns provide various microhabitats, a suitable trapping standard for canopy-dwelling arthropods is required. However, vertical trap position is often not standardized. We developed six vertical placement strategies for flight-interception traps based on information on tree height and crown base obtained from forest inventory data. Strategies differed in how crown base and height of trees were weighted in the calculations of trap height. Forest inventory data from almost 1600 plots in three regions of Germany. Were used to calculate the theoretical suitability of the strategies. The strategy to place traps at the canopy centre was predicted to result in very few trees for which placement would not be possible because trees are too short or crown base is too high, and this strategy by definition resulted in the lowest deviation of trap position from the canopy centre. The strategy Centre was tested in the field by installing three flight-interception traps each in 150 deciduous or coniferous stands of various age structures. On average, 53.8% of the traps were installed at exactly the desired height and for 86.9% of the traps the difference was <10% from the desired height. The strategy worked less well in thickets and pole woods. Overall, our results show that (1) forest inventory data are highly suitable to derive a priori trap placement strategies, even though these data are generally collected in only a small circle in each forest stand, (2) placing traps in the canopy centre largely avoids the problem of not being able to place traps at the desired height in the field, and (3) calculations of trap height should ideally be based on plot-specific data, but data from many stands of one forest type also provide a reasonable fit. [ABSTRACT FROM AUTHOR]

Organic farming in Europe has been shown to enhance biodiversity locally, but potential interactions with the surrounding landscape and the potential effects on ecosystem services are less well known. In cereal fields on 153 farms in five European regions, we examined how the species richness and abundance of wild plants, ground beetles and breeding birds, and the biological control potential of the area, were affected by organic and conventional farming, and how these effects were modified by landscape complexity (percentage of arable crops within 1000 m of the study plots). Information on biodiversity was gathered from vegetation plots, pitfall traps and by bird territory mapping. The biological control potential was measured as the percentage of glued, live aphids removed from plastic labels exposed in cereal fields for 24 h. Predation on aphids was highest in organic fields in complex landscapes, and declined with increasing landscape homogeneity. The biological control potential in conventional fields was not affected by landscape complexity, and in homogenous landscapes it was higher in conventional fields than in organic fields, as indicated by an interaction between farming practice and landscape complexity. A simplification of the landscape, from 20% to 100% arable land, reduced plant species richness by about 16% and cover by 14% in organic fields, and 33% and 5·5% in conventional fields. For birds, landscape simplification reduced species richness and abundance by 34% and 32% in organic fields and by 45·5% and 39% in conventional fields. Ground beetles were more abundant in simple landscapes, but were unaffected by farming practice. Synthesis and applications. This Europe-wide study shows that organic farming enhanced the biodiversity of plants and birds in all landscapes, but only improved the potential for biological control in heterogeneous landscapes. These mixed results stress the importance of taking both local management and regional landscape complexity into consideration when developing future agri-environment schemes, and suggest that local-regional interactions may affect other ecosystem services and functions. This study also shows that it is not enough to design and monitor agri-environment schemes on the basis of biodiversity, but that ecosystem services should be considered too. [ABSTRACT FROM AUTHOR]

In insect species characterized by inbreeding, limited dispersal, and a metapopulation structure, high genetic differentiation and reduced genetic diversity within local populations are expected. Using the model system Lysiphlebus hirticornis Mackauer, a specialist parasitoid of the tansy aphid, Metopeurum fuscoviride Stroyan (Hemiptera: Aphididae), we examined within-site temporal population dynamics and genetics, including molecular variation at the tansy plant level. Aphid-parasitoid dynamics were surveyed and parasitoids sampled from 72 tansy plants at 11 sites in and around Jena, Germany, over one growing season. Thereafter, parasitoid samples were genotyped at 11 polymorphic microsatellite loci. Colonization, extinction, and recolonization events occurred during the season. Allele numbers and identities were highly variable over time. When samples from all sites were pooled, allele number over all loci showed a decreasing trend with time. At the level of sites, temporal changes in genetic diversity were more variable. Analysis of molecular variance revealed that samples at the plant level explained the highest variance compared to at site level. We conclude that the genetic structuring of this insect is very fine grained (i.e. at the tansy plant level) and the temporal genetic diversity is explained by a combination of extinction and recolonization events, as well as inbreeding. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, , 737-749. [ABSTRACT FROM AUTHOR]

Abstract: Bird abundance, species richness and diversity of bird communities were analysed on 59 plots during breeding season and 61 plots during winter (500m×500m around winter wheat fields) in two contrasting regions of Germany differing in land-use history (West vs. East), landscape structural complexity (40–100% arable land) and farming practice (organic vs. conventional). Abundance, species richness and diversity were higher in West Germany, presumably due to the higher overall habitat-type diversity, and decreased with decreasing landscape complexity, this being marked for farmland and forest birds. In contrast, abundances of openland bird species were higher in East Germany during both seasons and increased with decreasing landscape complexity. During breeding season (but not in winter) organic farming enhanced the species richness of all bird groups, presumably due to more and diverse food resources. Farmland and forest bird species in agricultural landscapes therefore appear to be mainly enhanced by landscape complexity owing to the availability of nesting and sheltering places in non-crop habitats, while openland bird species require high proportions of arable land. [Copyright &y& Elsevier]

Recent theoretical studies suggest that the stability of ecosystem processes is not governed by diversity per se, but by multitrophic interactions in complex communities. However, experimental evidence supporting this assumption is scarce. We investigated the impact of plant diversity and the presence of above- and below-ground invertebrates on the stability of plant community productivity in space and time, as well as the interrelationship between both stability measures in experimental grassland communities. We sampled above-ground plant biomass on subplots with manipulated above- and below-ground invertebrate densities of a grassland biodiversity experiment (Jena Experiment) 1, 4 and 6 years after the establishment of the treatments to investigate temporal stability. Moreover, we harvested spatial replicates at the last sampling date to explore spatial stability. The coefficient of variation of spatial and temporal replicates served as a proxy for ecosystem stability. Both spatial and temporal stability increased to a similar extent with plant diversity. Moreover, there was a positive correlation between spatial and temporal stability, and elevated plant density might be a crucial factor governing the stability of diverse plant communities. Above-ground insects generally increased temporal stability, whereas impacts of both earthworms and above-ground insects depended on plant species richness and the presence of grasses. These results suggest that inconsistent results of previous studies on the diversity-stability relationship have in part been due to neglecting higher trophic-level interactions governing ecosystem stability. Changes in plant species diversity in one trophic level are thus unlikely to mirror changes in multitrophic interrelationships. Our results suggest that both above- and below-ground invertebrates decouple the relationship between spatial and temporal stability of plant community productivity by differently affecting the homogenizing mechanisms of plants in diverse plant communities. Synthesis. Species extinctions and accompanying changes in multitrophic interactions are likely to result not only in alterations in the magnitude of ecosystem functions but also in its variability complicating the assessment and prediction of consequences of current biodiversity loss. [ABSTRACT FROM AUTHOR]

Background: One of the most significant consequences of contemporary global change is the rapid decline of biodiversity in many ecosystems. Knowledge of the consequences of biodiversity loss in terrestrial ecosystems is largely restricted to single ecosystem functions. Impacts of key plant functional groups on soil biota are considered to be more important than those of plant diversity; however, current knowledge mainly relies on short-term experiments. Methodology/Principal Findings: We studied changes in the impacts of plant diversity and presence of key functional groups on soil biota by investigating the performance of soil microorganisms and soil fauna two, four and six years after the establishment of model grasslands. The results indicate that temporal changes of plant community effects depend on the trophic affiliation of soil animals: plant diversity effects on decomposers only occurred after six years, changed little in herbivores, but occurred in predators after two years. The results suggest that plant diversity, in terms of species and functional group richness, is the most important plant community property affecting soil biota, exceeding the relevance of plant above- and belowground productivity and the presence of key plant functional groups, i.e. grasses and legumes, with the relevance of the latter decreasing in time. Conclusions/Significance: Plant diversity effects on biota are not only due to the presence of key plant functional groups or plant productivity highlighting the importance of diverse and high-quality plant derived resources, and supporting the validity of the singular hypothesis for soil biota. Our results demonstrate that in the long term plant diversity essentially drives the performance of soil biota questioning the paradigm that belowground communities are not affected by plant diversity and reinforcing the importance of biodiversity for ecosystem functioning. [ABSTRACT FROM AUTHOR]

Abstract: This study examined the effects of agricultural intensity, various farming practices, landscape composition and vegetation cover on the abundance and species richness of wintering farmland birds, assessed simultaneously across seven European regions. The abundance and species richness of wintering farmland birds were negatively affected by agricultural intensity. The effects of yield and farm type were interlinked. Of the 10 farming practices assessed, mechanical weeding and the amount of organic fertilizer applied negatively affected farmland birds, presumably due to reduced food availability on arable fields. Positive effects of organic farming on farmland birds proved to be limited to simplified landscapes. More farmland birds were observed in areas with more stubble, pasture and green manure crops. Species richness was higher in areas with more pasture. The results of this study show that farm management, vegetation cover and landscape composition all influence wintering farmland birds. Heterogeneous landscapes comprising arable crops as well as grasslands support most species of farmland birds in winter. The effectiveness of organic farming and agri-environment schemes depends on landscape composition. Therefore, different agri-environment schemes should be designed for different landscape types. [ABSTRACT FROM AUTHOR]

In order to reduce parasite-induced mortality, hosts may be involved in mutualistic interactions in which the partner contributes to resistance against the parasite. The pea aphid, Acyrthosiphon pisum Harris (Hemiptera: Aphididae), harbours secondary bacterial endosymbionts, some of which have been reported to confer resistance against aphid parasitoids. Although this resistance often results in death of the developing parasitoid larvae, some parasitoid individuals succeed in developing into adults. Whether these individuals suffer from fitness reduction compared to parasitoids developing in pea aphid clones without symbionts has not been tested so far. Using 30 pea aphid clones that differed in their endosymbiont complement, we studied the effects of these endosymbionts on aphid resistance against the parasitoid Aphidius ervi Haliday (Hymenoptera: Braconidae: Aphidiinae), host–parasitoid physiological interactions, and fitness of emerging adult parasitoids. The number of symbiont species in an aphid clone was positively correlated with a number of resistance measurements but there were also clear symbiont-specific effects on the host–parasitoid interaction. As in previous studies, pea aphid clones infected with Hamiltonella defensa Moran et al. showed resistance against the parasitoid. In addition, pea aphid clones infected with Regiella insecticola Moran et al. and co-infections of H. defensa– Spiroplasma, R. insecticola– Spiroplasma, and R. insecticola– H. defensa showed reduced levels of parasitism and mummification. Parasitoids emerging from symbiont-infected aphid clones often had a longer developmental time and reduced mass. The number of teratocytes was generally lower when parasitoids oviposited in aphid clones with a symbiont complement. Interestingly, unparasitized aphids infected with Serratia symbiotica Moran et al. and R. insecticola had a higher fecundity than unparasitized aphids of uninfected pea aphid clones. We conclude that in addition to conferring resistance, pea aphid symbionts also negatively affect parasitoids that successfully hatch from aphid mummies. Because of the link between aphid resistance and the number of teratocytes, the mechanism underlying resistance by symbiont infection may involve interference with teratocyte development. [ABSTRACT FROM AUTHOR]

Abstract: A major issue in current ecological research is the effect of biodiversity on ecosystem functioning. Although several studies reported a positive diversity – productivity relationship, the role of soil animals has been largely neglected. Nematodes are among the most widespread and important herbivores causing substantial yield losses in agriculture; however, impacts of nematodes on the diversity – productivity relationship in semi-natural plant communities have not been investigated until today. In the framework of the Jena Experiment (Thuringia, Germany) we established control and nematicide treated subplots to manipulate nematode densities on plots varying in plant species (1–16) and functional group richness (1–4). We explored the interacting effects of nematicide application and plant diversity on the main trophic groups of nematodes and on aboveground plant productivity. Nematicide application reduced the number of nematodes significantly, particularly that of plant feeders and predators. The negative impact of nematicide application on plant and bacterial feeders depended however on the diversity of the plant community. Total plant shoot biomass tended to decrease in the presence of ambient nematode densities. In detail, nematode effects varied however with plant functional group identity by reducing only the shoot biomass of herbs significantly but not that of legumes. Furthermore, the shoot biomass of grasses tended to decrease in the presence of ambient nematode densities. In contrast to total shoot biomass, nematodes decreased grass shoot biomass only in high diverse but not in low diverse plant communities. Thus, the present study for the first time highlights that nematodes likely modify the community structure und functions of semi-natural plant communities by altering the competition between plant functional groups and by attenuating the diversity – productivity relationship. [Copyright &y& Elsevier]

The consequences of plant species loss are rarely assessed in a multi-trophic context and especially effects on life-history traits of organisms at higher trophic levels have remained largely unstudied. We used a grassland biodiversity experiment and measured the effects of two components of plant diversity, plant species richness and the presence of nitrogen-fixing legumes, on several life-history traits of naturally colonizing aphids and their primary and secondary parasitoids in the field. We found that, irrespective of aphid species identity, the proportion of winged aphid morphs decreased with increasing plant species richness, which was correlated with decreasing host plant biomass. Similarly, emergence proportions of parasitoids decreased with increasing plant species richness. Both, emergence proportions and proportions of female parasitoids were lower in plots with legumes, where host plants had increased nitrogen concentrations. This effect of legume presence could indicate that aphids were better defended against parasitoids in high-nitrogen environments. Body mass of emerged individuals of the two most abundant primary parasitoid species was, however, higher in plots with legumes, suggesting that once parasitoids could overcome aphid defenses, they could profit from larger or more nutritious hosts. Our study demonstrates that cascading effects of plant species loss on higher trophic levels such as aphids, parasitoids and secondary parasitoids begin with changed life-history traits of these insects. Thus, life-history traits of organisms at higher trophic levels may be useful indicators of bottom-up effects of plant diversity on the biodiversity of consumers. [ABSTRACT FROM AUTHOR]

In beech-dominated forests in Central Europe, many spring geophytes show adaptations to seed dispersal by ants (myrmecochory). Ants, however, can be rare in such moist forests. Motivated by observations of slug feeding on seeds we investigated the seed consumption of two plant species, Anemone nemorosa and Asarum europaeum, by slugs, in a series of experiments. In a seed predation experiment in a beech forest, we found that seed removal was strongly reduced when gastropods were excluded from the seed depots. The contribution of insects, including ants, and rodents to seed removal was relatively less but differed between May and July. In the laboratory, slug species, in particular Arion sp., consumed seeds of both plant species. Slugs either consumed the elaiosomes of seeds or swallowed seeds intact. Swallowed seeds were defecated undamaged and germinated as well as control seeds when buried overwinter, indicating the potential for seed dispersal by slugs. We also recovered seeds of myrmecochores in the faeces of several slugs caught in forests. In a slug release experiment in the forest, slugs moved up to 14.6 m (mean 4.4 m) in 15 h, which is the median gut passage time of seeds based on measurements made in the laboratory. We also found that when slug-defecated seeds were offered to rodents, these were less attractive than control seeds, suggesting that passage through the slug gut reduces seed predation risk. Our results demonstrate that slugs are significant consumers of elaiosomes or entire seeds of ant-dispersed plants and that they can function as seed dispersers of these plants. [ABSTRACT FROM AUTHOR]

The resistance of a plant community against herbivore attack may depend on plant species richness, with monocultures often much more severely affected than mixtures of plant species. Here, we used a plant–herbivore system to study the effects of selective herbivory on consumption resistance and recovery after herbivory in 81 experimental grassland plots. Communities were established from seed in 2002 and contained 1, 2, 4, 8, 16 or 60 plant species of 1, 2, 3 or 4 functional groups. In 2004, pairs of enclosure cages (1 m tall, 0.5 m diameter) were set up on all 81 plots. One randomly selected cage of each pair was stocked with 10 male and 10 female nymphs of the meadow grasshopper, Chorthippus parallelus. The grasshoppers fed for 2 months, and the vegetation was monitored over 1 year. Consumption resistance and recovery of vegetation were calculated as proportional changes in vegetation biomass. Overall, grasshopper herbivory averaged 6.8%. Herbivory resistance and recovery were influenced by plant functional group identity, but independent of plant species richness and number of functional groups. However, herbivory induced shifts in vegetation composition that depended on plant species richness. Grasshopper herbivory led to increases in herb cover at the expense of grasses. Herb cover increased more strongly in species-rich mixtures. We conclude that selective herbivory changes the functional composition of plant communities and that compositional changes due to selective herbivory depend on plant species richness. [ABSTRACT FROM AUTHOR]

Invertebrate herbivores are ubiquitous in most terrestrial ecosystems, and theory predicts that their impact on plant community biomass should depend on diversity and productivity of the associated plant communities. To elucidate general patterns in the relationship between invertebrate herbivory, plant diversity, and productivity, we carried out a long-term herbivore exclusion experiment at multiple grassland sites in a mountainous landscape of central Germany. Over a period of five years, we used above- and belowground insecticides as well as a molluscicide to manipulate invertebrate herbivory at 14 grassland sites, covering a wide range of plant species diversity (13-38 species/m²) and aboveground plant productivity (272-1125 g·m-2·yr-1), where plant species richness and productivity of the sites were not significantly correlated. Herbivore exclusion had significant effects on the plant communities: it decreased plant species richness and evenness, and it altered plant community composition. In particular, exclusion of belowground herbivores promoted grasses at the expense of herbs. In contrast to our expectation, herbivore effects on plant community biomass were not influenced by productivity. However, effect size of invertebrate herbivores was negatively correlated with plant diversity of the grasslands: the effect of herbivory on biomass tended to be negative at sites of high diversity and positive at sites of low diversity. In general, the effects of aboveground herbivores were relatively small as compared to belowground herbivores, which were important drivers of plant community composition. Our study is the first to show that variation in the effects of invertebrate herbivory on plant communities across a landscape is significantly influenced by plant species richness. [ABSTRACT FROM AUTHOR]

To understand the influence of plant quality on intraguild predation and consequently on the suppression of a shared prey population as well as on plant yield, the interactions between Aphis gossypii Glover (Hemiptera: Aphididae) (shared prey), Aphidoletes aphidimyza (Rondani) (Diptera: Cecidomyiidae) (intermediate predator), and Orius laevigatus Fieber (Hemiptera: Anthocoridae) (top predator) were investigated in 25-day experiments on cucumber, Cucumis sativus L. (Cucurbitaceae) at various N fertilization levels (90, 150, and 190 p.p.m.) in microcosm set-ups under greenhouse conditions. The final aphid population size was significantly affected by an interactive effect of N fertilization and predator application. Regardless of the N fertilization levels, O. laevigatus alone was more effective in aphid suppression than A. aphidimyza alone. In addition, the risk for aphids of being predated upon by both predators together was significantly reduced in the low and medium-N fertilization levels, whereas it was additive in the high-N fertilization treatment. The A. aphidimyza population was suppressed by O. laevigatus in both the 90 and 150 p.p.m. N treatments. However, there was no intraguild predation of O. laevigatus on A. aphidimyza at the 190 p.p.m. N level. Total plant yield depended on predator treatments and N fertilization levels, with the highest yield produced at the 150 p.p.m. N fertilization level in treatments with either O. laevigatus alone or with both predators together. Our results demonstrate that the weak asymmetric intraguild predation among A. aphidimyza and O. laevigatus does not influence the ability of both predators together to diminish bottom-up effects on aphid populations and the yield losses associated with aphid infestations. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

1. Generalist insect herbivores occupy a variety of habitats that differ in food plant composition. Dietary mixing has been proposed as a possibility for generalists to overcome nutritional deficiencies of single plant species, but only a few studies have investigated herbivore feeding and fitness for diets that resemble natural scenarios. We studied feeding behaviour, survival, and reproduction of the generalist grasshopper Chorthippus parallelus raised on food plants of four typical habitats. 2. Grasshopper diet consisted of grasses (92.5%), legumes (6.7%) and, in small quantities, other forbs (0.8%). Diet selection differed between the four food plant mixtures, and depended on grasshopper sex and developmental stage. There was no correlation between the relative abundance of plant species in the field and the fraction of these species in the grasshopper diet. 3. Grasshoppers survived on average for 40.4 ± 1.0 days before maturity, grew 106.8 mg until maturity moult, and females laid 4.1 ± 0.4 egg pods, each of which contained 8.5 ± 0.4 eggs. However, despite the differences in feeding behaviour, grasshopper fitness was the same in all of the four food plant mixtures. While the digestibility of ingested food was similar in the four different treatments, indices indicated differences in the conversion efficiency to body mass. 4. Our results show that C. parallelus is a plastic feeder with no fixed preferences in diet composition. The results emphasise that generalist herbivores can counteract putative quality deficiencies of single food plants by selective dietary mixing. [ABSTRACT FROM AUTHOR]

Theory predicts negative effects of increasing plant diversity on the abundance of specialist insect herbivores, but little is known about how plant diversity affects the performance and abundance of generalist insect herbivores. We studied oviposition rates and offspring numbers in females of the generalist grasshopper Chorthippus parallelus that were collected in 15 montane grasslands, in 2005 and 2007 along a gradient of plant species richness in central Germany. In addition to plant species richness, we determined evenness and plant community composition in the grasslands and measured aboveground plant biomass and other habitat variables such as leaf area index, vegetation height, and solar radiation. There was substantial variation among sites in grasshopper fecundity and the number of nymphs that hatched from the egg pods. Both fitness measures were positively influenced by plant species richness at the sites, while female fitness did not correlate with any of the other habitat parameters. Abundance of C. parallelus in the grasslands was positively correlated with plant species richness, plant community composition, and incident solar radiation of the sites. There were no phenological differences between grasshoppers from the different study sites. Our results suggest that decreasing biodiversity threatens the persistence not only of specialist, but also of generalist insect herbivores via a variety of mechanisms including a decrease in diversity of the generalists' food plants. [ABSTRACT FROM AUTHOR]

Abstract: Soil organic carbon (OC) stocks are highly variable in space. This study aimed at analysing regional soil OC stock variability with direct measurements of OC stocks at 89 sampling points distributed across a forested area (160km−2) of the Hainich region in Thuringia/Germany. We sampled complete soil profiles by horizons and depth increments. The major goal of this study was the analysis of regional OC stocks. Soils of our study area were classified as Cambisols, Luvisols and Stagnosols. The averaged total OC stocks of the solum ranged from 7.4kgm−2 in Luvisols to 10.3kgm−2 in Cambisols, but the analysis of variance revealed no significant differences among the soil groups. In contrast, significant differences in A and B horizon OC stocks of soil groups could be detected. This was not the case, when depth increments instead of soil horizons were studied. Our results implied that sampling by depth increments is favourable for the detection of regional OC stocks and that sampling by soil horizons should be used for the study of pedogenic processes which control OC stocks in soil. Differences in OC stocks among soil groups are more pronounced if horizons are considered, which shows that mixing of soil horizons by fixed depth increments destroy important pedogenetical information. A stepwise multiple regression exposed horizon thickness as the most important factor affecting OC stocks in soil horizons. This has major implications for detection of changes in carbon stock, since the variability of regional OC stocks is less controlled by OC concentration and bulk density rather than by soil horizon thickness. To detect differences smaller than 10% of regional OC stocks one would need more than 300 A horizon samples. If 0–10cm increments were sampled, less than 100 samples would be sufficient to detect the same difference. [Copyright &y& Elsevier]