The functioning and service provisioning of ecosystems in the face of anthropogenic environmental and biodiversity change is a cornerstone of ecological research. The last three decades of ...biodiversity–ecosystem functioning (BEF) research have provided compelling evidence for the significant positive role of biodiversity in the functioning of many ecosystems. Despite broad consensus of this relationship, the underlying ecological and evolutionary mechanisms have not been well understood. This complicates the transition from a description of patterns to a predictive science. The proposed Research Unit aims at filling this gap of knowledge by applying novel experimental and analytical approaches in one of the longest-running biodiversity experiments in the world: the Jena Experiment. The central aim of the Research Unit is to uncover the mechanisms that determine BEF relationships in the short- and in the long-term. Increasing BEF relationships with time in long-term experiments do not only call for a paradigm shift in the appreciation of the relevance of biodiversity change, they likely are key to understanding the mechanisms of BEF relationships in general. The subprojects of the proposed Research Unit fall into two tightly linked main categories with two research areas each that aim at exploring variation in community assembly processes and resulting differences in biotic interactions as determinants of the long-term BEF relationship. Subprojects under “Microbial community assembly” and “Assembly and functions of animal communities” mostly focus on plant diversity effects on the assembly of communities and their feedback effects on biotic interactions and ecosystem functions. Subprojects under “Mediators of plant-biotic interactions” and “Intraspecific diversity and micro-evolutionary changes” mostly focus on plant diversity effects on plant trait expression and micro-evolutionary adaptation, and subsequent feedback effects on biotic interactions and ecosystem functions. This unification of evolutionary and ecosystem processes requires collaboration across the proposed subprojects in targeted plant and soil history experiments using cutting-edge technology and will produce significant synergies and novel mechanistic insights into BEF relationships. The Research Unit of the Jena Experiment is uniquely positioned in this context by taking an interdisciplinary and integrative approach to capture whole-ecosystem responses to changes in biodiversity and to advance a vibrant research field.
Biodiversity-ecosystem function experiments test how species diversity influences fundamental ecosystem processes. Historically, arthropod driven functions, such as herbivory and pest-control, have ...been thought to be influenced by direct and indirect associations among species. Although a number of studies have evaluated how plant diversity affects arthropod communities and arthropod-mediated ecosystem processes, it remains unclear whether diversity effects on arthropods are sufficiently consistent over time such that observed responses can be adequately predicted by classical hypotheses based on associational effects. By combining existing results from a long-term grassland biodiversity experiment (Jena Experiment) with new analyses, we evaluate the consistency of consumer responses within and across taxonomic, trophic, and trait-based (i.e. vertical stratification) groupings, and we consider which changes in arthropod community composition are associated with changes in consumer-mediated ecosystem functions.
Overall, higher plant species richness supported more diverse and complex arthropod communities and this pattern was consistent across multiple years. Vegetation-associated arthropods responded more strongly to changes in plant species richness than ground-dwelling arthropods. Additionally, increases in plant species richness were associated with shifts in the species-abundance distributions for many, but not all taxa. For example, highly specialized consumers showed a decrease in dominance and an increase in the number of rare species with increasing plant species richness. Most ecosystem processes investigated responded to increases in plant species richness in the same way as the trophic group mediating the process, e.g. both herbivory and herbivore diversity increase with increasing plant species richness. In the Jena Experiment and other studies, inconsistencies between predictions based on classic hypotheses of associational effects and observed relationships between plant species richness and arthropod diversity likely reflect the influence of multi-trophic community dynamics and species functional trait distributions. Future research should focus on testing a broader array of mechanisms to unravel the biological processes underlying the biodiversity-ecosystem functioning relationships.
Understanding factors that maintain ecosystem stability is critical in the face of environmental change. Experiments simulating species loss from grassland have shown that losing biodiversity ...decreases ecosystem stability. However, as the originally sown experimental communities with reduced biodiversity develop, plant evolutionary processes or the assembly of interacting soil organisms may allow ecosystems to increase stability over time. We explored such effects in a long-term grassland biodiversity experiment with plant communities with either a history of co-occurrence (selected communities) or no such history (naïve communities) over a 4-yr period in which a major flood disturbance occurred. Comparing communities of identical species composition, we found that selected communities had temporally more stable biomass than naïve communities, especially at low species richness. Furthermore, selected communities showed greater biomass recovery after flooding, resulting in more stable post-flood productivity. In contrast to a previous study, the positive diversity–stability relationship was maintained after the flooding. Our results were consistent across three soil treatments simulating the presence or absence of co-selected microbial communities. We suggest that prolonged exposure of plant populations to a particular community context and abiotic site conditions can increase ecosystem temporal stability and resilience due to short-term evolution. A history of co-occurrence can in part compensate for species loss, as can high plant diversity in part compensate for the missing opportunity of such adaptive adjustments.
Recent studies have shown that the diversity of flowering plants can enhance pollinator richness and visitation frequency and thereby increase the resilience of pollination. It is assumed that flower ...traits explain these effects, but it is still unclear which flower traits are responsible, and knowing that, if pollinator richness and visitation frequency are more driven by mass‐ratio effects (mean trait values) or by trait diversity.
Here, we analyse a three‐year data set of pollinator observations collected in a European grassland plant diversity experiment (The Jena experiment). The data entail comprehensive flower trait measurements, including reward traits (nectar and pollen amount), morphological traits (height, symmetry, area, colour spectra) and chemical traits (nectar‐amino acid and nectar‐sugar concentration). We test if pollinator species richness and visitation frequency of flower communities depend on overall functional diversity combining all flower traits within a community, single trait diversities (within trait variation) and community‐weighted means of the single traits, using Bayesian inference.
Overall functional diversity did not affect pollinator species richness, but reduced visitation frequency. When looking at individual flower traits separately, we found that single trait diversity of flower reflectance and flower morphology were important predictors of pollinator visitation frequency. Moreover, independent of total flower abundance, community‐weighted means of flower height, area, reflectance, nectar‐sugar concentration and nectar‐amino acid concentration strongly affected both pollinator species richness and visitation frequency.
Our results, challenge the idea that functional diversity always positively affects ecosystem functions. Nonetheless, we demonstrate that both single trait diversity and mass‐ratio effects of flower traits play an important role for diverse and frequent flower visits, which underlines the functionality of flower traits for pollination services.
It remains unclear whether biodiversity buffers ecosystems against climate extremes, which are becoming increasingly frequent worldwide. Early results suggested that the ecosystem productivity of ...diverse grassland plant communities was more resistant, changing less during drought, and more resilient, recovering more quickly after drought, than that of depauperate communities. However, subsequent experimental tests produced mixed results. Here we use data from 46 experiments that manipulated grassland plant diversity to test whether biodiversity provides resistance during and resilience after climate events. We show that biodiversity increased ecosystem resistance for a broad range of climate events, including wet or dry, moderate or extreme, and brief or prolonged events. Across all studies and climate events, the productivity of low-diversity communities with one or two species changed by approximately 50% during climate events, whereas that of high-diversity communities with 16-32 species was more resistant, changing by only approximately 25%. By a year after each climate event, ecosystem productivity had often fully recovered, or overshot, normal levels of productivity in both high- and low-diversity communities, leading to no detectable dependence of ecosystem resilience on biodiversity. Our results suggest that biodiversity mainly stabilizes ecosystem productivity, and productivity-dependent ecosystem services, by increasing resistance to climate events. Anthropogenic environmental changes that drive biodiversity loss thus seem likely to decrease ecosystem stability, and restoration of biodiversity to increase it, mainly by changing the resistance of ecosystem productivity to climate events.
Identifying to what degree inherent characteristics of plant species and their variation in response to their environment regulate the temporal stability of plant populations is important to ...understand patterns of species coexistence and the stability of ecosystems. Longevity is a key characteristic of plant life history and an important component of demographic storage, but age is usually unknown for herbaceous species.
In a 12‐year‐old biodiversity experiment (Jena Experiment) comprising 80 grassland communities with six levels of plant species richness (1, 2, 4, 8, 16 and 60 species) and four levels of functional groups richness (1, 2, 3 and 4 functional groups), we studied populations of 38 dicotyledonous forb species (N = 1,683 plant individuals). The sampled individuals represented three plant functional groups (legumes, small herbs and tall herbs) and two different growth forms (species with long‐lived primary roots and clonal species with rhizomes/stolons). We assessed the age of plant individuals by means of growth ring analysis and related the age of plant populations to their temporal stability in terms of peak biomass production.
On average, plant species richness did not affect the mean age of the populations or the maximum age of individuals found in a population. Age of herbs with taproots increased and age of herbs with clonal growth decreased with increasing species richness, cancelling out each other when growth forms were analysed together. Mean population age was lowest for small herbs and highest for tall herbs, while legumes had an intermediate population age. Herbs with a taproot were on average older than herbs with a rhizome. Across all species‐richness levels, populations with older individuals were more stable in terms of biomass production over time.
Synthesis. Our study shows for the first time across multiple species that the longevity of forbs is affected by the diversity of the surrounding plant community, and that plant longevity as an important component of demographic storage increases the temporal stability of populations of grassland forb species.
Our study shows for the first time across multiple species that the longevity of forbs is affected by the diversity of the surrounding plant community, and that plant longevity as an important component of demographic storage increases the temporal stability of populations of grassland forb species.
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Changes to primary producer diversity can cascade up to consumers and affect ecosystem processes. Although the effect of producer diversity on higher trophic groups have been studied, these studies ...often quantify taxonomy‐based measures of biodiversity, like species richness, which do not necessarily reflect the functioning of these communities. In this study, we assess how plant species richness affects the functional composition and diversity of higher trophic levels and discuss how this might affect ecosystem processes, such as herbivory, predation and decomposition.
Based on six different consumer traits, we examined the functional composition of arthropod communities sampled in experimental plots that differed in plant species richness. The two components we focused on were functional variation in the consumer community structure (functional structure) and functional diversity, expressed as functional richness, evenness and divergence.
We found a consistent positive effect of plant species richness on the functional richness of herbivores, carnivores, and omnivores, but not decomposers, and contrasting patterns for functional evenness and divergence. Increasing plant species richness shifted the omnivore community to more predatory and less mobile species, and the herbivore community to more specialized and smaller species. This was accompanied by a shift towards more species occurring in the vegetation than in the ground layer.
Our study shows that plant species richness strongly affects the functional structure and diversity of aboveground arthropod communities. The observed shifts in body size (herbivores), specialization (herbivores), and feeding mode (omnivores) together with changes in the functional diversity may underlie previously observed increases in herbivory and predation in plant communities of higher diversity.
Plant diversity changes can impact the abundance, diversity, and functioning of species at higher trophic levels. We used an experimental gradient in grassland plant diversity ranging from 1 to 16 ...plant species to study multitrophic interactions among plants, cavity-nesting bees and wasps, and their natural enemies, and analysed brood cell density, insect diversity (species richness), and bee and wasp community similarity over two consecutive years. The bee and wasp communities were more similar among the high (16 species) diversity plots than among plots of the lower diversity levels (up to 8 species), and a more similar community of bees and wasps resulted in a more similar community of their parasitoids. Plant diversity, which was closely related to flower diversity, positively and indirectly affected bee diversity and the diversity of their parasitoids via increasing brood cell density of bees. Increasing plant diversity directly led to higher wasp diversity. Parasitism rates of bees and wasps (hosts) were not affected by plant diversity, but increased with the diversity of their respective parasitoids. Decreases in parasitism rates of bees arose from increasing brood cell density of bees (hosts), whereas decreasing parasitism rates of wasps arose from increasing wasp diversity (hosts). In conclusion, decreases in plant diversity propagated through different trophic levels: from plants to insect hosts to their parasitoids, decreasing density and diversity. The positive relationship between plant diversity and the community similarity of higher trophic levels indicates a community-stabilising effect of high plant diversity.
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.
1. Plants influence associated soil biotic communities that in turn can alter the performance of the subsequently growing plants. Although such "plant-soil feedbacks" (PSFs) are considered as ...important drivers of plant community assembly, past PSF studies have mainly addressed plant biomass production. However, plant performance is not only the production of biomass but comprises a sequence of different life stages: from seed germination over vegetative growth up to the production of a viable progeny. 2. Here, we assessed the effects of soil biotic communities that were previously conditioned for 3 years by a focal plant species monoculture or species mixtures on key plant life stages from germination and vegetative growth to flowering and the production of viable seeds. We used three common grassland herb species that were grown in a sterile substrate and inoculated with a sterile control soil or with living soils. Living soils were conditioned either by the focal species in monoculture or a four- or eight-species mixture that included the focal species to represent a decrease in the target plants' conspecific influence on the soil communities. 3. We show that the effect of soil biota changed from positive at the plants' juvenile life stages to neutral or negative at the plants' adult life stages and ultimately decreased plant fitness. A higher conspecific influence on the soil communities pronounced the positive effects at the juvenile life stage but also the negative effects at adult life stages. Further, we observed direct soil biotic effects on flower production and plant fitness that were not mediated by adult biomass production. This suggests that soil biotic effects may alter plant resource allocation and even may have transgenerational effects on plant fitness. 4. Synthesis. We conclude that there is no overarching effect of soil biota that remains consistent at all the life stages of a plant. Thus, our results highlight the importance to consider plant life stage and ultimately plant fitness especially when plant-soil interactions are used to explain plant community dynamics.
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