Anne Ebeling

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

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

Biodiversity Increases the Resistance of Ecosystem Productivity to Climate Extremes

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.

Multitrophic effects of experimental changes in plant diversity on cavity-nesting bees, wasps, and their parasitoids

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.

Plant diversity effects on grassland productivity are robust to both nutrient enrichment and drought

Global change drivers are rapidly altering resource availability and biodiversity. While there is consensus that greater biodiversity increases the functioning of ecosystems, the extent to which biodiversity buffers ecosystem productivity in response to changes in resource availability remains unclear. We use data from 16 grassland experiments across North America and Europe that manipulated plant species richness and one of two essential resources—soil nutrients or water—to assess the direction and strength of the interaction between plant diversity and resource alteration on above-ground productivity and net biodiversity, complementarity, and selection effects. Despite strong increases in productivity with nutrient addition and decreases in productivity with drought, we found that resource alterations did not alter biodiversity–ecosystem functioning relationships. Our results suggest that these relationships are largely determined by increases in complementarity effects along plant species richness gradients. Although nutrient addition reduced complementarity effects at high diversity, this appears to be due to high biomass in monocultures under nutrient enrichment. Our results indicate that diversity and the complementarity of species are important regulators of grassland ecosystem productivity, regardless of changes in other drivers of ecosystem function.

Biodiversity effects on plant stoichiometry.

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.

Invertebrate herbivory increases along an experimental gradient of grassland plant diversity

Plant diversity is a key driver of ecosystem functioning best documented for its influence on plant productivity. The strength and direction of plant diversity effects on species interactions across trophic levels are less clear. For example, with respect to the interactions between herbivorous invertebrates and plants, a number of competing hypotheses have been proposed that predict either increasing or decreasing community herbivory with increasing plant species richness. We investigated foliar herbivory rates and consumed leaf biomass along an experimental grassland plant diversity gradient in year eight after establishment. The gradient ranged from one to 60 plant species and manipulated also functional group richness (from one to four functional groups—legumes, grasses, small herbs, and tall herbs) and plant community composition. Measurements in monocultures of each plant species showed that functional groups differed in the quantity and quality of herbivory damage they experienced, with legumes being more damaged than grasses or non-legume herbs. In mixed plant communities, herbivory increased with plant diversity and the presence of two key plant functional groups in mixtures had a positive (legumes) and a negative (grasses) effect on levels of herbivory. Further, plant community biomass had a strong positive impact on consumed leaf biomass, but little effect on herbivory rates. Our results contribute detailed data from a well-established biodiversity experiment to a growing body of evidence suggesting that an increase of herbivory with increasing plant diversity is the rule rather than an exception. Considering documented effects of herbivory on other ecosystem functions and the increase of herbivory with plant diversity, levels of herbivory damage might not only be a result, but also a trigger within the diversity–productivity relationship.

Flooding disturbances increase resource availability and productivity but reduce stability in diverse plant communities

The natural world is increasingly defined by change. Within the next 100 years, rising atmospheric CO₂ concentrations will continue to increase the frequency and magnitude of extreme weather events. Simultaneously, human activities are reducing global biodiversity, with current extinction rates at ~1,000 × what they were before human domination of Earths ecosystems. The co-occurrence of these trends may be of particular concern, as greater biological diversity could help ecosystems resist change during large perturbations. We use data from a 200-year flood event to show that when a disturbance is associated with an increase in resource availability, the opposite may occur. Flooding was associated with increases in productivity and decreases in stability, particularly in the highest diversity communities. Our results undermine the utility of the biodiversity-stability hypothesis during a large number of disturbances where resource availability increases. We propose a conceptual framework that can be widely applied during natural disturbances.

Plant Diversity Impacts Decomposition and Herbivory via Changes in Aboveground Arthropods

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.

Plant species richness and functional traits affect community stability after a flood event.

Climate change is expected to increase the frequency and magnitude of extreme weather events. It is therefore of major importance to identify the community attributes that confer stability in ecological communities during such events. In June 2013, a flood event affected a plant diversity experiment in Central Europe (Jena, Germany). We assessed the effects of plant species richness, functional diversity, flooding intensity and community means of functional traits on different measures of stability (resistance, resilience and raw biomass changes from pre-flood conditions). Surprisingly, plant species richness reduced community resistance in response to the flood. This was mostly because more diverse communities grew more immediately following the flood. Raw biomass increased over the previous year; this resulted in decreased absolute value measures of resistance. There was no clear response pattern for resilience. We found that functional traits drove these changes in raw biomass: communities with a high proportion of late-season, short-statured plants with dense, shallow roots and small leaves grew more following the flood. Late-growing species probably avoided the flood, whereas greater root length density might have allowed species to better access soil resources brought from the flood, thus growing more in the aftermath. We conclude that resource inputs following mild floods may favour the importance of traits related to resource acquisition and be less associated with flooding tolerance.

Plants are less negatively affected by flooding when growing in species‐rich plant communities

Flooding is expected to increase in frequency and severity in the future. The ecological consequences of flooding are the combined result of species-specific plant traits and ecological context. However, the majority of past flooding research has focused on individual model species under highly controlled conditions. An early summer flooding event in a grassland biodiversity experiment in Jena, Germany, provided the opportunity to assess flooding responses of 60 grassland species in monocultures and 16-species mixtures. We examined plant biomass, species-specific traits (plant height, specific leaf area (SLA), root aerenchyma, starch content) and soil porosity. We found that, on average, plant species were less negatively affected by the flood when grown in higher-diversity plots in July 2013. By September 2013, grasses were unaffected by the flood regardless of plant diversity, and legumes were severely negatively affected regardless of plant diversity. Plants with greater SLA and more root aerenchyma performed better in September. Soil porosity was higher in higher-diversity plots and had a positive effect on plant performance. As floods become more frequent and severe in the future, growing flood-sensitive plants in higher-diversity communities and in soil with greater soil aeration may attenuate the most negative effects of flooding.