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IntroductionIn production forests, management can have cascading effects on biodiversity and ecosystem services. Acoustic diversity reflects the diversity of vocalizing animals and has also considerable recreational value for human well-being, but the relationship between acoustic diversity and forest management remains largely unexploredMethodWe recorded acoustic diversity on forest plots along a gradient of silvicultural management intensity (SMI) in three regions of Germany. We explored the diurnal and seasonal temporal dynamics in acoustic diversity index (ADI) from March to July using generalized additive mixed models (GAMMs). We further investigated the interrelation between acoustic diversity and silvicultural management intensity, forest structural diversity, as well as tree diversity, bird species richness and abundance using structural equation modeling (SEM).ResultsSilvicultural management intensity had significant effects on the temporal dynamics of ADI in May and June from dawn till dusk, but variance explained by SMI was low. We confirmed our hypothesis that ADI was reduced by SMI due to its cascading effects on forest structural diversity and bird species richness and abundance.DiscussionAcoustic diversity indices can provide valuable insights into how forest management affects the acoustic activity of soniferous communities. We discuss how this can indicate both changes in species diversity as well as their vocal activity. We further address potential implications for forest management.

Abstract Plant diversity effects on community productivity often increase over time. Whether the strengthening of diversity effects is caused by temporal shifts in species-level overyielding (i.e., higher species-level productivity in diverse communities compared with monocultures) remains unclear. Here, using data from 65 grassland and forest biodiversity experiments, we show that the temporal strength of diversity effects at the community scale is underpinned by temporal changes in the species that yield. These temporal trends of species-level overyielding are shaped by plant ecological strategies, which can be quantitatively delimited by functional traits. In grasslands, the temporal strengthening of biodiversity effects on community productivity was associated with increasing biomass overyielding of resource-conservative species increasing over time, and with overyielding of species characterized by fast resource acquisition either decreasing or increasing. In forests, temporal trends in species overyielding differ when considering above- versus belowground resource acquisition strategies. Overyielding in stem growth decreased for species with high light capture capacity but increased for those with high soil resource acquisition capacity. Our results imply that a diversity of species with different, and potentially complementary, ecological strategies is beneficial for maintaining community productivity over time in both grassland and forest ecosystems.

IntroductionForests managed for timber production can also be managed for biodiversity conservation by retaining structures typical of old-growth forests, which provide heterogenous structures for forest-dwelling species, including birds. Ecoacoustic monitoring of forest birds is now a well-studied field, however the extent to which acoustic indices can reflect bird assemblage responses to stands of lower or higher structural heterogeneity is disputed. MethodIn this study, we acoustically surveyed 75 plots over two years in mature, previously managed forests and computed acoustic indices from the recordings. We first identified an acoustic index that significantly correlated with bird richness over more than one spring season. Next, we tested the response of bird richness to individual forest structural elements using linear regressions. We then repeated this analysis but combined the individual structural elements into one structural composition variable using an NMDS and gam smooth overlay, to compare the effect of individual forest structures versus overall forest heterogeneity on bird richness. We then repeated this analysis using our selected acoustic index, the Normalized Difference Soundscape Index (NDSI), to see if it followed the same patterns as bird richness in response to individual and collective forest structural elements.ResultsOur results showed that plots with high bird richness were also associated with high NDSI values, and high values of both variables occurred in plots with high structural heterogeneity (tree species richness, tree size variability, and snag height variability) and low amounts of standing and lying deadwood. DiscussionOur findings suggest that once an acoustic index can be identified as a robust correlate of bird richness, it can serve as a proxy for the response of birds to differences in forest structural heterogeneity within a managed forest context. It is therefore possible for forest managers interested in conserving or increasing bird richness in their production forests to identify a robust acoustic index to track the response of bird richness to management decisions over time.

Biodiversity-ecosystem functioning relationships may change over time. Here, Wagg et al. show that richness-productivity and richness stability relationships grow stronger over time in an experimental grassland community, and shed light on the ecological mechanisms.

Measurement(s) plant trait Technology Type(s) various Factor Type(s) none Sample Characteristic - Organism Tracheophyta Sample Characteristic - Environment natural environment Sample Characteristic - Location global

Using experimental communities of grassland species, this study shows that drought-exposure history can accelerate recovery from subsequent drought through increased niche complementarity between species. This transgenerational effect may enhance the sustainability of biodiversity and ecosystem functioning in a future with more frequent droughts.

Abstract Background Positive effects of plant species richness on community biomass in biodiversity experiments are often stronger than those from observational field studies. This may be because experiments are initiated with randomly assembled species compositions whereas field communities have experienced filtering. Methods We compared aboveground biomass production of randomly assembled communities of 2–16 species (controls) with experimentally filtered communities from which subordinate species were removed, resulting in removal communities of 1–8 species. Results Removal communities had (1) 12.6% higher biomass than control communities from which they were derived, that is, with double species richness and (2) 32.0% higher biomass than control communities of equal richness. These differences were maintained along the richness gradient. The increased productivity of removal communities was paralleled by increased species evenness and complementarity. Conclusions Result (1) indicates that subordinate species can reduce community biomass production, suggesting a possible explanation for why the most diverse field communities sometimes do not have the highest productivity. Result (2) suggests that if a community of S species has been derived by filtering from a pool of 2S randomly chosen species it is more productive than a community derived from a pool of S randomly chosen species without filtering.

Investigating changes in belowground functional plant traits is an important step toward a better understanding of vegetation dynamics during primary succession. However, in alpine glacier forelands, we still lack an accurate assessment of plant rooting patterns. In this study, we established two proglacial chronosequences with contrasting bedrocks to investigate changes in rooting patterns and biomass allocation with terrain age. We extracted soil cores up to 1 m depth and measured root traits every 10 cm of each drilled core. Furthermore, we sampled aboveground biomass determining the contributions of functional groups to total aboveground biomass. We found that root traits associated with the root economics spectrum varied significantly along the chronosequences. Vertical root distribution coefficients revealed that early successional communities had more evenly distributed root systems compared to late successional communities. Biomass allocation showed diverging patterns. We found evidence for both the isometric allocation and optimal partitioning hypotheses. In addition, we observed a significant correlation between rooting parameters and plant community composition, suggesting that the dominance of distinct plant functional groups was one important factor explaining the observed rooting patterns. Our results shed light on the often neglected belowground compartments during plant succession and contribute to a better understanding of hillslope functioning.

Tick-borne diseases represent the majority of vector-borne human diseases in Europe, with Ixodes ricinus, mostly present in forests, as the main vector. Studies show that vertebrate hosts diversification would decrease the prevalence of these pathogens. However, it is not well known whether habitat diversity can have similar impact on ticks and their infection rates. We measured the presence and abundance of different stages of I. ricinus, and the prevalence of associated pathogens in a large-scale forest experiment in which we manipulated tree diversity and moisture level. We showed that larval abundance was influenced by tree species identity, with larvae being more present in pine plots than in oak plots, while nymph abundance increased with canopy tree density. The proportion of Borrelia burgdorferi s.l.-infected nymphs decreased with increasing tree diversity. Our findings suggest that tree overstorey composition, structure and diversity, can affect tick abundance and pathogen prevalence. They support the idea that forest habitats may have “diluting” or “amplifying” effects on tick-borne diseases with direct relevance for human health.

Abstract Biodiversity is a major driver of numerous ecosystem functions. However, consequences of changes in forest biodiversity remain difficult to predict because of limited knowledge about how tree diversity influences ecosystem functions. Litter decomposition is a key process affecting nutrient cycling, productivity, and carbon storage and can be influenced by plant biodiversity. Leaf litter species composition, environmental conditions, and the detritivore community are main components of the decomposition process, but their complex interactions are poorly understood. In this study, we tested the effect of tree functional diversity (FD) on litter decomposition in a field experiment manipulating tree diversity and partitioned the effects of litter physiochemical diversity and the detritivore community. We used litterbags with different mesh sizes to separate the effects of microorganisms and microfauna, mesofauna, and macrofauna and monitored soil fauna using pitfall traps and earthworm extractions. We hypothesized that higher tree litter FD accelerates litter decomposition due to the availability of complementary food components and higher activity of detritivores. Although we did not find direct effects of tree FD on litter decomposition, we identified key litter traits and macrodetritivores that explained part of the process. Litter mass loss was found to decrease with an increase in leaf litter carbon:nitrogen ratio. Moreover, litter mass loss increased with an increasing density of epigeic earthworms, with most pronounced effects in litterbags with a smaller mesh size, indicating indirect effects. Higher litter FD and litter nutrient content were found to increase the density of surface‐dwelling macrofauna and epigeic earthworm biomass. Based on structural equation modeling, we conclude that tree FD has a weak positive effect on soil surface litter decomposition by increasing the density of epigeic earthworms and that litter nitrogen‐related traits play a central role in tree composition effects on soil fauna and decomposition.

Abstract Background In contrast with the negligible contribution of the forest understorey to the total aboveground phytobiomass of a forest, its share in annual litter production and nutrient cycling may be more important. Whether and how this functional role of the understorey differs across forest types and depends upon overstorey characteristics remains to be investigated. Methods We sampled 209 plots of the FunDivEUROPE Exploratory Platform, a network of study plots covering local gradients of tree diversity spread over six contrasting forest types in Europe. To estimate the relative contribution of the understorey to carbon and nutrient cycling, we sampled non-lignified aboveground understorey biomass and overstorey leaf litterfall in all plots. Understorey samples were analysed for C, N and P concentrations, overstorey leaf litterfall for C and N concentrations. We additionally quantified a set of overstorey attributes, including species richness, proportion of evergreen species, light availability (representing crown density) and litter quality, and investigated whether they drive the understorey’s contribution to carbon and nutrient cycling. Results and conclusions Overstorey litter production and nutrient stocks in litterfall clearly exceeded the contribution of the understorey for all forest types, and the share of the understorey was higher in forests at the extremes of the climatic gradient. In most of the investigated forest types, it was mainly light availability that determined the contribution of the understorey to yearly carbon and nutrient cycling. Overstorey species richness did not affect the contribution of the understorey to carbon and nutrient cycling in any of the investigated forest types.

Abstract Retention forestry, which retains a portion of the original stand at the time of harvesting to maintain continuity of structural and compositional diversity, has been originally developed to mitigate the impacts of clear‐cutting. Retention of habitat trees and deadwood has since become common practice also in continuous‐cover forests of Central Europe. While the use of retention in these forests is plausible, the evidence base for its application is lacking, trade‐offs have not been quantified, it is not clear what support it receives from forest owners and other stakeholders and how it is best integrated into forest management practices. The Research Training Group ConFoBi (Conservation of Forest Biodiversity in Multiple‐use Landscapes of Central Europe) focusses on the effectiveness of retention forestry, combining ecological studies on forest biodiversity with social and economic studies of biodiversity conservation across multiple spatial scales. The aim of ConFoBi is to assess whether and how structural retention measures are appropriate for the conservation of forest biodiversity in uneven‐aged and selectively harvested continuous‐cover forests of temperate Europe. The study design is based on a pool of 135 plots (1 ha) distributed along gradients of forest connectivity and structure. The main objectives are (a) to investigate the effects of structural elements and landscape context on multiple taxa, including different trophic and functional groups, to evaluate the effectiveness of retention practices for biodiversity conservation; (b) to analyze how forest biodiversity conservation is perceived and practiced, and what costs and benefits it creates; and (c) to identify how biodiversity conservation can be effectively integrated in multi‐functional forest management. ConFoBi will quantify retention levels required across the landscape, as well as the socio‐economic prerequisites for their implementation by forest owners and managers. ConFoBi's research results will provide an evidence base for integrating biodiversity conservation into forest management in temperate forests.

Higher plant diversity in agricultural settings is often associated with lower biomass yield and with lower forage quality. Here, Schaub et al. show positive effects of plant diversity on biomass yield, quality-adjusted yield and revenues in semi-natural grassland across a range of management intensities.

In this ecoacoustic study we used the setting of a tropical tree diversity planted forest to analyze temporal patterns in the composition of soundscapes and to test the effects of tree species richness on associated biodiversity measured as acoustic diversity. The analysis of soundscapes offers easy, rapid and sustainable methods when assessing biodiversity. During the last years the quantification of regional or global acoustic variability in sounds and the analysis of different soundscapes has been evolving into an important tool for biodiversity conservation, especially since case studies confirmed a relationship between land-use management, forest structure and acoustic diversity. Here we analyzed soundscapes from two seasons (dry and rainy season) and aurally inspected a subset of audio recordings to describe temporal patterns in soundscape composition. Several acoustic indices were calculated and we performed a correlation analysis and a non-metric multidimensional scaling analysis to identify acoustic indices that: (i) were complementary to each other and such represented different aspects of the local soundscapes and (ii) related most strongly to differences in acoustic composition among tree species richness, season and day phase. Thus, we chose “High Frequency Cover,” “Bioacoustic Index,” and “Events Per Second” to test the hypothesis that acoustic diversity increases with increasing tree species richness. Monocultures differed significantly from polycultures during night recordings, with respect to High Frequency Cover. This index covers sounds above 8 kHz and thus represents part of the orthopteran community. We conclude that increasing tree species richness in a young tropical forest plantation had positive effects on the vocalizing communities. The strongest effects were found for acoustic activity of the orthopteran community. In contrast to birds, orthopterans have smaller home ranges, and are therefore important indicator species for small scale environmental conditions.

Abstract One of the most important drivers for the coexistence of plant species is the resource heterogeneity of a certain environment, and several studies in different ecosystems have supported this resource heterogeneity–diversity hypothesis. However, to date, only a few studies have measured heterogeneity of light and soil resources below forest canopies to investigate their influence on understory plant species richness. Here, we aim to determine (1) the influence of forest stand structural complexity on the heterogeneity of light and soil resources below the forest canopy and (2) whether heterogeneity of resources increases understory plant species richness. Measures of stand structural complexity were obtained through inventories and remote sensing techniques in 135 1‐ha study plots of temperate forests, established along a gradient of forest structural complexity. We measured light intensity and soil chemical properties on six 25 m² subplots on each of these 135 plots and surveyed understory vegetation. We calculated the coefficient of variation of light and soil parameters to obtain measures of resource heterogeneity and determined understory plant species richness at plot level. Spatial heterogeneity of light and of soil pH increased with higher stand structural complexity, although heterogeneity of soil pH did not increase in conditions of generally high levels of light availability. Increasing light heterogeneity was also associated with increasing understory plant species richness. However, light heterogeneity had no such effects in conditions where soil resource heterogeneity (variation in soil C:N ratios) was low. Our results support the resource heterogeneity–diversity hypothesis for temperate forest understory at the stand scale. Our results also highlight the importance of interaction effects between the heterogeneity of both light and soil resources in determining plant species richness.

Abstract As of 2020, the world has an estimated 290 million ha of planted forests and this number is continuously increasing. Of these, 131 million ha are monospecific planted forests under intensive management. Although monospecific planted forests are important in providing timber, they harbor less biodiversity and are potentially more susceptible to disturbances than natural or diverse planted forests. Here, we point out the increasing scientific evidence for increased resilience and ecosystem service provision of functionally and species diverse planted forests (hereafter referred to as diverse planted forests) compared to monospecific ones. Furthermore, we propose five concrete steps to foster the adoption of diverse planted forests: (1) improve awareness of benefits and practical options of diverse planted forests among land‐owners, managers, and investors; (2) incentivize tree species diversity in public funding of afforestation and programs to diversify current maladapted planted forests of low diversity; (3) develop new wood‐based products that can be derived from many different tree species not yet in use; (4) invest in research to assess landscape benefits of diverse planted forests for functional connectivity and resilience to global‐change threats; and (5) improve the evidence base on diverse planted forests, in particular in currently under‐represented regions, where new options could be tested.

Litter decomposition is a key process for carbon and nutrient cycling in terrestrial ecosystems and is mainly controlled by environmental conditions, substrate quantity and quality as well as microbial community abundance and composition. In particular, the effects of climate and atmospheric nitrogen (N) deposition on litter decomposition and its temporal dynamics are of significant importance, since their effects might change over the course of the decomposition process. Within the TeaComposition initiative, we incubated Green and Rooibos teas at 524 sites across nine biomes. We assessed how macroclimate and atmospheric inorganic N deposition under current and predicted scenarios (RCP 2.6, RCP 8.5) might affect litter mass loss measured after 3 and 12 months. Our study shows that the early to mid-term mass loss at the global scale was affected predominantly by litter quality (explaining 73% and 62% of the total variance after 3 and 12 months, respectively) followed by climate and N deposition. The effects of climate were not litter-specific and became increasingly significant as decomposition progressed, with MAP explaining 2% and MAT 4% of the variation after 12 months of incubation. The effect of N deposition was litter-specific, and significant only for 12-month decomposition of Rooibos tea at the global scale. However, in the temperate biome where atmospheric N deposition rates are relatively high, the 12-month mass loss of Green and Rooibos teas decreased significantly with increasing N deposition, explaining 9.5% and 1.1% of the variance, respectively. The expected changes in macroclimate and N deposition at the global scale by the end of this century are estimated to increase the 12-month mass loss of easily decomposable litter by 1.1–3.5% and of the more stable substrates by 3.8–10.6%, relative to current mass loss. In contrast, expected changes in atmospheric N deposition will decrease the mid-term mass loss of high-quality litter by 1.4–2.2% and that of low-quality litter by 0.9–1.5% in the temperate biome. Our results suggest that projected increases in N deposition may have the capacity to dampen the climate-driven increases in litter decomposition depending on the biome and decomposition stage of substrate.

Abstract For decades, ecologists have investigated the effects of tree species diversity on tree productivity at different scales and with different approaches ranging from observational to experimental study designs. Using data from five European national forest inventories (16,773 plots), six tree species diversity experiments (584 plots), and six networks of comparative plots (169 plots), we tested whether tree species growth responses to species mixing are consistent and therefore transferrable between those different research approaches. Our results confirm the general positive effect of tree species mixing on species growth (16% on average) but we found no consistency in species‐specific responses to mixing between any of the three approaches, even after restricting comparisons to only those plots that shared similar mixtures compositions and forest types. These findings highlight the necessity to consider results from different research approaches when selecting species mixtures that should maximize positive forest biodiversity and functioning relationships.

Scientific knowledge in the field of ecology is increasingly enriched by data acquired by the general public participating in citizen science (CS) programs. Yet, doubts remain about the reliability of such data, in particular when acquired by schoolchildren. We built upon an ongoing CS program, 'Oak Bodyguards', to assess the ability of schoolchildren to accurately estimate the strength of biotic interactions in terrestrial ecosystems. We used standardized protocols to estimate attack rates on artificial caterpillars and insect herbivory on oak leaves. We compared estimates made by schoolchildren with estimates made by professional scientists who had been trained in predation and herbivory assessments (henceforth, trained scientists), and trained scientists’ estimates with those made by professional scientists with or without expertise (untrained) in predation or herbivory assessment. Compared with trained scientists, both schoolchildren and untrained professional scientists overestimated attack rates, but assessments made by the latter were more consistent. Schoolchildren tended to overestimate insect herbivory, as did untrained professional scientists. Raw data acquired by schoolchildren participating in CS programs therefore require several quality checks by trained professional scientists before being used. However, such data are of no less value than data collected by untrained professional scientists. CS with schoolchildren can be a valuable tool for carrying out ecological research, provided that the data itself is acquired by professional scientists from material collected by citizens.

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

Abstract Background The tick Ixodes ricinus has considerable impact on the health of humans and other terrestrial animals because it transmits several tick-borne pathogens (TBPs) such as B. burgdorferi (sensu lato), which causes Lyme borreliosis (LB). Small forest patches of agricultural landscapes provide many ecosystem services and also the disservice of LB risk. Biotic interactions and environmental filtering shape tick host communities distinctively between specific regions of Europe, which makes evaluating the dilution effect hypothesis and its influence across various scales challenging. Latitude, macroclimate, landscape and habitat properties drive both hosts and ticks and are comparable metrics across Europe. Therefore, we instead assess these environmental drivers as indicators and determine their respective roles for the prevalence of B. burgdorferi in I. ricinus. Methods We sampled I. ricinus and measured environmental properties of macroclimate, landscape and habitat quality of forest patches in agricultural landscapes along a European macroclimatic gradient. We used linear mixed models to determine significant drivers and their relative importance for nymphal and adult B. burgdorferi prevalence. We suggest a new prevalence index, which is pool-size independent. Results During summer months, our prevalence index varied between 0 and 0.4 per forest patch, indicating a low to moderate disservice. Habitat properties exerted a fourfold larger influence on B. burgdorferi prevalence than macroclimate and landscape properties combined. Increasingly available ecotone habitat of focal forest patches diluted and edge density at landscape scale amplified B. burgdorferi prevalence. Indicators of habitat attractiveness for tick hosts (food resources and shelter) were the most important predictors within habitat patches. More diverse and abundant macro- and microhabitat had a diluting effect, as it presumably diversifies the niches for tick-hosts and decreases the probability of contact between ticks and their hosts and hence the transmission likelihood. Conclusions Diluting effects of more diverse habitat patches would pose another reason to maintain or restore high biodiversity in forest patches of rural landscapes. We suggest classifying habitat patches by their regulating services as dilution and amplification habitat, which predominantly either decrease or increase B. burgdorferi prevalence at local and landscape scale and hence LB risk. Particular emphasis on promoting LB-diluting properties should be put on the management of those habitats that are frequently used by humans. In the light of these findings, climate change may be of little concern for LB risk at local scales, but this should be evaluated further.

Abstract Background The castor bean tick (Ixodes ricinus) transmits infectious diseases such as Lyme borreliosis, which constitutes an important ecosystem disservice. Despite many local studies, a comprehensive understanding of the key drivers of tick abundance at the continental scale is still lacking. We analyze a large set of environmental factors as potential drivers of I. ricinus abundance. Our multi-scale study was carried out in deciduous forest fragments dispersed within two contrasting rural landscapes of eight regions, along a macroclimatic gradient stretching from southern France to central Sweden and Estonia. We surveyed the abundance of I. ricinus, plant community composition, forest structure and soil properties and compiled data on landscape structure, macroclimate and habitat properties. We used linear mixed models to analyze patterns and derived the relative importance of the significant drivers. Results Many drivers had, on their own, either a moderate or small explanatory value for the abundance of I. ricinus, but combined they explained a substantial part of variation. This emphasizes the complex ecology of I. ricinus and the relevance of environmental factors for tick abundance. Macroclimate only explained a small fraction of variation, while properties of macro- and microhabitat, which buffer macroclimate, had a considerable impact on tick abundance. The amount of forest and the composition of the surrounding rural landscape were additionally important drivers of tick abundance. Functional (dispersules) and structural (density of tree and shrub layers) properties of the habitat patch played an important role. Various diversity metrics had only a small relative importance. Ontogenetic tick stages showed pronounced differences in their response. The abundance of nymphs and adults is explained by the preceding stage with a positive relationship, indicating a cumulative effect of drivers. Conclusions Our findings suggest that the ecosystem disservices of tick-borne diseases, via the abundance of ticks, strongly depends on habitat properties and thus on how humans manage ecosystems from the scale of the microhabitat to the landscape. This study stresses the need to further evaluate the interaction between climate change and ecosystem management on I. ricinus abundance.

Abstract Models predict that vertical gradients of foliar nitrogen (N) allocation, increasing from bottom to top of plant canopies, emerge as a plastic response to optimise N utilisation for carbon assimilation. While this mechanism has been well documented in monocultures, its relevance for mixed stands of varying species richness remains poorly understood. We used 21 naturally assembled grassland communities to analyse the gradients of N in the canopy using N allocation coefficients (K N ) estimated from the distribution of N per foliar surface area (KN-F) and ground surface area (KN-G). We tested whether: 1) increasing plant species richness leads to more pronounced N gradients as indicated by higher K N -values, 2) K N is a good predictor of instantaneous net ecosystem CO2 exchange and 3) functional diversity of leaf N concentration as estimated by Rao’s Q quadratic diversity metric is a good proxy of K N . Our results show a negative (for KN-G) or no relationship (for KN-F) between species richness and canopy N distribution, but emphasize a link (positive relationship) between more foliar N per ground surface area in the upper layers of the canopy (i.e. under higher KN-G) and ecosystem CO2 uptake. Rao’s Q was not a good proxy for either K N .

Abstract Background Species richness affects processes and functions in many ecosystems. Since management of temperate grasslands is directly affecting species composition and richness, it can indirectly govern how systems respond to fluctuations in environmental conditions. Our aim in this study was to investigate whether species richness in managed grasslands can buffer the effects of drought and warming manipulations and hence increase the resistance to climate change. We established 45 plots in three regions across Germany, each with three different management regimes (pasture, meadow and mown pasture). We manipulated spring warming using open-top chambers and summer drought using rain-out shelters for 4 weeks. Results Measurements of species richness, above- and below-ground biomass and soil carbon and nitrogen concentrations showed significant but inconsistent differences among regions, managements and manipulations. We detected a three-way interaction between species richness, management and region, indicating that our study design was sensitive enough to detect even intricate effects. Conclusions We could not detect a pervasive effect of species richness on biomass differences between treatments and controls, indicating that a combination of spring warming and summer drought effects on grassland systems are not consistently moderated by species richness. We attribute this to the relatively high number of species even at low richness levels, which already provides the complementarity required for positive biodiversity–ecosystem functioning relationships. A review of the literature also indicates that climate manipulations largely fail to show richness-buffering, while natural experiments do, suggesting that such manipulations are milder than reality or incur treatment artefacts.

How biodiversity is linked to multiple ecosystem functions is not fully understood. Here, the authors show that a new mechanism, which they term the 'jack-of-all-trades' effect, best explains patterns of tree diversity and ecosystem multifunctionality in European forests.

Abstract Human‐caused declines in biodiversity have stimulated intensive research on the consequences of biodiversity loss for ecosystem services and policy initiatives to preserve the functioning of ecosystems. Short‐term biodiversity experiments have documented positive effects of plant species richness on many ecosystem functions, and longer‐term studies indicate, for some ecosystem functions, that biodiversity effects can become stronger over time. Theoretically, a biodiversity effect can strengthen over time by an increasing performance of high‐diversity communities, by a decreasing performance of low‐diversity communities, or a combination of both processes. Which of these two mechanisms prevail, and whether the increase in the biodiversity effect over time is a general property of many functions remains currently unclear. These questions are an important knowledge gap as a continuing decline in the performance of low‐diversity communities would indicate an ecosystem‐service debt resulting from delayed effects of species loss on ecosystem functioning. Conversely, an increased performance of high‐diversity communities over time would indicate that the benefits of biodiversity are generally underestimated in short‐term studies. Analyzing 50 ecosystem variables over 11 years in the world's largest grassland biodiversity experiment, we show that overall plant diversity effects strengthened over time. Strengthening biodiversity effects were independent of the considered compartment (above‐ or belowground), organizational level (ecosystem variables associated with the abiotic habitat, primary producers, or higher trophic levels such as herbivores and pollinators), and variable type (measurements of pools or rates). We found evidence that biodiversity effects strengthened because of both a progressive decrease in functioning in species‐poor and a progressive increase in functioning in species‐rich communities. Our findings provide evidence that negative feedback effects at low biodiversity are as important for biodiversity effects as complementarity among species at high biodiversity. Finally, our results indicate that a current loss of species will result in a future impairment of ecosystem functioning, potentially decades beyond the moment of species extinction.

Tree diversity in forests is an important driver of ecological processes including herbivory. Empirical evidence suggests both negative and positive effects of tree diversity on herbivory, which can be, respectively, attributed to associational resistance or associational susceptibility. Tree diversity experiments allow testing for associational effects, but evidence regarding which pattern predominates is mixed. Furthermore, it is unknown if herbivory on tree species of native vs. exotic origin is influenced by changing tree diversity in a similar way, or if exotic tree species escape natural enemies, resulting in lower damage that is unrelated to tree diversity. To address these questions, we established a young tree diversity experiment in temperate southwestern Germany that uses high planting density (49 trees per plot; plot size 13 m2). The species pool consists of six congeneric species pairs of European and North American origin (12 species in total) planted in monocultures and mixtures (1, 2, 4, 6 species). We assessed leaf damage by leaf-chewing insects on more than 5,000 saplings of six broadleaved tree species. Plot-level tree species richness increased leaf damage, which more than doubled from monocultures to six-species mixtures, strongly supporting associational susceptibility. However, leaf damage among congeneric native and exotic species pairs was similar. There were marked differences in patterns of leaf damage across tree genera, and only the genera likely having a predominately generalist herbivore community showed associational susceptibility, irrespective of the geographical origin of a tree species. In conclusion, an increase in tree species richness in young temperate forests may result in associational susceptibility to feeding by generalist herbivores.

Effects of tree species diversity on decomposition can operate via a multitude of mechanism, including alterations of microclimate by the forest canopy. Studying such effects in natural settings is complicated by the fact that topography also affects microclimate and thus decomposition, so that effects of diversity are more difficult to isolate. Here, we quantified decomposition rates of standard litter in young subtropical forest stands, separating effects of canopy tree species richness and topography, and quantifying their direct and micro-climate-mediated components. Our litterbag study was carried out at two experimental sites of a biodiversity-ecosystem functioning field experiment in south-east China (BEF-China). The field sites display strong topographical heterogeneity and were planted with tree communities ranging from monocultures to mixtures of 24 native subtropical tree species. Litter bags filled with senescent leaves of three native tree species were placed from Nov. 2011 to Oct. 2012 on 134 plots along the tree species diversity gradient. Topographic features were measured for all and microclimate in a subset of plots. Stand species richness, topography and microclimate explained important fractions of the variations in litter decomposition rates, with diversity and topographic effects in part mediated by microclimatic changes. Tree stands were 2-3 years old, but nevertheless tree species diversity explained more variation (54.3%) in decomposition than topography (7.7%). Tree species richness slowed litter decomposition, an effect that slightly depended on litter species identity. A large part of the variance in decomposition was explained by tree species composition, with the presence of three tree species playing a significant role. Microclimate explained 31.4% of the variance in decomposition, and was related to lower soil moisture. Within this microclimate effect, species diversity (without composition) explained 8.9% and topography 34.4% of variance. Topography mainly affected diurnal temperature amplitudes by varying incident solar radiation.

BackgroundLoss of biodiversity and increased nutrient inputs are two of the most crucial anthropogenic factors driving ecosystem change. Although both received considerable attention in previous studies, information on their interactive effects on ecosystem functioning is scarce. In particular, little is known on how soil biota and their functions are affected by combined changes in plant diversity and fertilization.Methodology/principal findingsWe investigated the effects of plant diversity, functional community composition, and fertilization on the biomass and respiration of soil microbial communities in a long-term biodiversity experiment in semi-natural grassland (Jena Experiment). Plant species richness enhanced microbial basal respiration and microbial biomass, but did not significantly affect microbial specific respiration. In contrast, the presence of legumes and fertilization significantly decreased microbial specific respiration, without altering microbial biomass. The effect of legumes was superimposed by fertilization as indicated by a significant interaction between the presence of legumes and fertilization. Further, changes in microbial stoichiometry (C-to-N ratio) and specific respiration suggest the presence of legumes to reduce N limitation of soil microorganisms and to modify microbial C use efficiency.Conclusions/significanceOur study highlights the role of plant species and functional group diversity as well as interactions between plant community composition and fertilizer application for soil microbial functions. Our results suggest soil microbial stoichiometry to be a powerful indicator of microbial functioning under N limited conditions. Although our results support the notion that plant diversity and fertilizer application independently affect microbial functioning, legume effects on microbial N limitation were superimposed by fertilization, indicating significant interactions between the functional composition of plant communities and nutrient inputs for soil processes.

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.

The degree to which biodiversity may promote the stability of grasslands in the light of climatic variability, such as prolonged summer drought, has attracted considerable interest. Studies so far yielded inconsistent results and in addition, the effect of different grassland management practices on their response to drought remains an open question. We experimentally combined the manipulation of prolonged summer drought (sheltered vs. unsheltered sites), plant species loss (6 levels of 60 down to 1 species) and management intensity (4 levels varying in mowing frequency and amount of fertilizer application). Stability was measured as resistance and resilience of aboveground biomass production in grasslands against decreased summer precipitation, where resistance is the difference between drought treatments directly after drought induction and resilience is the difference between drought treatments in spring of the following year. We hypothesized that (i) management intensification amplifies biomass decrease under drought, (ii) resistance decreases with increasing species richness and with management intensification and (iii) resilience increases with increasing species richness and with management intensification.We found that resistance and resilience of grasslands to summer drought are highly dependent on management intensity and partly on species richness. Frequent mowing reduced the resistance of grasslands against drought and increasing species richness decreased resistance in one of our two study years. Resilience was positively related to species richness only under the highest management treatment. We conclude that low mowing frequency is more important for high resistance against drought than species richness. Nevertheless, species richness increased aboveground productivity in all management treatments both under drought and ambient conditions and should therefore be maintained under future climates.

The accelerating rate of change in biodiversity patterns, mediated by ever increasing human pressures and global warming, demands a better understanding of the relationship between the structure of biological communities and ecosystem functioning (BEF). Recent investigations suggest that the functional structure of communities, i.e. the composition and diversity of functional traits, is the main driver of ecological processes. However, the predictive power of BEF research is still low, the integration of all components of functional community structure as predictors is still lacking, and the multifunctionality of ecosystems (i.e. rates of multiple processes) must be considered. Here, using a multiple-processes framework from grassland biodiversity experiments, we show that functional identity of species and functional divergence among species, rather than species diversity per se, together promote the level of ecosystem multifunctionality with a predictive power of 80%. Our results suggest that primary productivity and decomposition rates, two key ecosystem processes upon which the global carbon cycle depends, are primarily sustained by specialist species, i.e. those that hold specialized combinations of traits and perform particular functions. Contrary to studies focusing on single ecosystem functions and considering species richness as the sole measure of biodiversity, we found a linear and non-saturating effect of the functional structure of communities on ecosystem multifunctionality. Thus, sustaining multiple ecological processes would require focusing on trait dominance and on the degree of community specialization, even in species-rich assemblages.

The diversity-stability hypothesis states that current losses of biodiversity can impair the ability of an ecosystem to dampen the effect of environmental perturbations on its functioning. Using data from a long-term and comprehensive biodiversity experiment, we quantified the temporal stability of 42 variables characterizing twelve ecological functions in managed grassland plots varying in plant species richness. We demonstrate that diversity increases stability i) across trophic levels (producer, consumer), ii) at both the system (community, ecosystem) and the component levels (population, functional group, phylogenetic clade), and iii) primarily for aboveground rather than belowground processes. Temporal synchronization across studied variables was mostly unaffected with increasing species richness. This study provides the strongest empirical support so far that diversity promotes stability across different ecological functions and levels of ecosystem organization in grasslands.