Diversity Promotes Temporal Stability across Levels of Ecosystem Organization in Experimental Grasslands

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

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.

Sexual selection and speciation: the comparative evidence revisited

The spectacular diversity in sexually selected traits in the animal kingdom has inspired the hypothesis that sexual selection can promote species divergence. In recent years, several studies have attempted to test this idea by correlating species richness with estimates of sexual selection across phylogenies. These studies have yielded mixed results and it remains unclear whether the comparative evidence can be taken as generally supportive. Here, we conduct a meta-analysis of the comparative evidence and find a small but significant positive overall correlation between sexual selection and speciation rate. However, we also find that effect size estimates are influenced by methodological choices. Analyses that included deeper phylogenetic nodes yielded weaker correlations, and different proxies for sexual selection showed different relationships with species richness. We discuss the biological and methodological implications of these findings. We argue that progress requires more representative sampling and justification of chosen proxies for sexual selection and speciation rate, as well as more mechanistic approaches.

High Bee and Wasp Diversity in a Heterogeneous Tropical Farming System Compared to Protected Forest

It is a globally important challenge to meet increasing demands for resources and, at the same time, protect biodiversity and ecosystem services. Farming is usually regarded as a major threat to biodiversity due to its expansion into natural areas. We compared biodiversity of bees and wasps between heterogeneous small-scale farming areas and protected forest in northern coastal Belize, Central America. Malaise traps operated for three months during the transition from wet to dry season. Farming areas consisted of a mosaic of mixed crop types, open habitat, secondary forest, and agroforestry. Mean species richness per site (alpha diversity), as well as spatial and temporal community variation (beta diversity) of bees and wasps were equal or higher in farming areas compared to protected forest. The higher species richness and community variation in farmland was due to additional species that did not occur in the forest, whereas most species trapped in forest were also found in farming areas. The overall regional species richness (gamma diversity) increased by 70% with the inclusion of farming areas. Our results suggest that small-scale farming systems adjacent to protected forest may not only conserve, but even favour, biodiversity of some taxonomic groups. We can, however, not exclude possible declines of bee and wasp diversity in more intensified farmland or in landscapes completely covered by heterogeneous farming systems.

Does delaying the first mowing date benefit biodiversity in meadowland?

Background Meadows are regularly mown in order to provide fodder or litter for livestock and to prevent vegetation succession. However, the time of year at which meadows should be first mown in order to maximize biological diversity remains controversial and may vary with respect to context and focal taxa. We carried out a systematic review and meta-analysis on the effects of delaying the first mowing date upon plants and invertebrates in European meadowlands. Methods Following a CEE protocol, ISI Web of Science, Science Direct, JSTOR, Google and Google Scholar were searched. We recorded all studies that compared the species richness of plants, or the species richness or abundance of invertebrates, between grassland plots mown at a postponed date (treatment) vs plots mown earlier (control). In order to be included in the meta-analysis, compared plots had to be similar in all management respects, except the date of the first cut that was (mostly experimentally) manipulated. They were also to be located in the same meadow type. Meta-analyses applying Hedges’d statistic were performed. Results Plant species richness responded differently to the date to which mowing was postponed. Delaying mowing from spring to summer had a positive effect, while delaying either from spring to fall, or from early summer to later in the season had a negative effect. Invertebrates were expected to show a strong response to delayed mowing due to their dependence on sward structure, but only species richness showed a clearly significant positive response. Invertebrate abundance was positively influenced in only a few studies. Conclusions The present meta-analysis shows that in general delaying the first mowing date in European meadowlands has either positive or neutral effects on plant and invertebrate biodiversity (except for plant species richness when delaying from spring to fall or from early summer to later). Overall, there was also strong between-study heterogeneity, pointing to other major confounding factors, the elucidation of which requires further field experiments with both larger sample sizes and a distinction between taxon-specific and meadow-type-specific responses.

Impact of Land-Use Intensity and Productivity on Bryophyte Diversity in Agricultural Grasslands

While bryophytes greatly contribute to plant diversity of semi-natural grasslands, little is known about the relationships between land-use intensity, productivity, and bryophyte diversity in these habitats. We recorded vascular plant and bryophyte vegetation in 85 agricultural used grasslands in two regions in northern and central Germany and gathered information on land-use intensity. To assess grassland productivity, we harvested aboveground vascular plant biomass and analyzed nutrient concentrations of N, P, K, Ca and Mg. Further we calculated mean Ellenberg indicator values of vascular plant vegetation. We tested for effects of land-use intensity and productivity on total bryophyte species richness and on the species richness of acrocarpous (small & erect) and pleurocarpous (creeping, including liverworts) growth forms separately. Bryophyte species were found in almost all studied grasslands, but species richness differed considerably between study regions in northern Germany (2.8 species per 16 m2) and central Germany (6.4 species per 16 m2) due environmental differences as well as land-use history. Increased fertilizer application, coinciding with high mowing frequency, reduced bryophyte species richness significantly. Accordingly, productivity estimates such as plant biomass and nitrogen concentration were strongly negatively related to bryophyte species richness, although productivity decreased only pleurocarpous species. Ellenberg indicator values for nutrients proved to be useful indicators of species richness and productivity. In conclusion, bryophyte composition was strongly dependent on productivity, with smaller bryophytes that were likely negatively affected by greater competition for light. Intensive land-use, however, can also indirectly decrease bryophyte species richness by promoting grassland productivity. Thus, increasing productivity is likely to cause a loss of bryophyte species and a decrease in species diversity.

Dispersal and seed limitation affect diversity and productivity of montane grasslands

So far, seed limitation as a local process, and dispersal limitation as a regional process have been largely neglected in biodiversity-ecosystem functioning research. However, these processes can influence both local plant species diversity and ecosystem processes, such as biomass production. We added seeds of 60 species from the regional species pool to grassland communities at 20 montane grassland sites in Germany. In these sites, plant species diversity ranged from 10 to 34 species m(-2) and, before manipulation, diversity was not related to aboveground biomass, which ranged from 108 to 687 g m(-2). One year after seed addition, local plant species richness had increased on average by six species m(-2) (29%) compared with control plots, and this increase was highest in grasslands with intermediate productivity. The increased diversity after adding seeds was associated with an average increase of aboveground biomass of 36 g m(-2) (14.8%) compared with control plots. Thus, our results demonstrate that a positive relationship between changes in species richness and productivity, as previously reported from experimental plant communities, also holds for natural grassland ecosystems. Our results show that local plant communities are dispersal limited and a hump-shaped model appears to be the limiting outline of the natural diversity-productivity relationship. Hence, the effects of dispersal on local diversity can substantially affect the functioning of natural ecosystems.

A comparison of the strength of biodiversity effects across multiple functions

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

Seed availability in hay meadows: Land-use intensification promotes seed rain but not the persistent seed bank

Intensification of land use in semi-natural hay meadows has resulted in a decrease in species diversity. This is often thought to be caused by the reduced establishment of plant species due to high competition for light under conditions of increased productivity. Sowing experiments in grasslands have found reliable evidence that diversity can also be constrained by seed availability, implying that processes influencing the production and persistence of seeds may be important for the functioning of ecosystems. So far, the effects of land-use intensification on the seed rain and the persistence of seeds in the soil have been unclear. We selected six pairs of extensively managed (Festuco-Brometea) and intensively managed (Arrhenatheretalia) grassland with traditional late cutting regimes across Switzerland and covering an annual productivity gradient in the range 176–1211 gm−2. In each grassland community, we estimated seed rain and seed bank using eight pooled seed-trap or topsoil samples of 89 cm2 in each of six plots representing an area of c. 150 m2. The seed traps were established in spring 2010 and collected simultaneously with soil cores after an exposure of c. three months. We applied the emergence method in a cold frame over eight months to estimate density of viable seeds. With community productivity reflecting land-use intensification, the density and species richness in the seed rain increased, while mean seed size diminished and the proportions of persistent seeds and of species with persistent seeds in the topsoil declined. Stronger limitation of seeds in extensively managed semi-natural grasslands can explain the fact that such grasslands are not always richer in species than more intensively managed ones.

The impact of plant diversity and fertilization on fitness of a generalist grasshopper

In many environments land use intensification is likely to result in a decrease in species richness and in an increase in eutrophication. Although the importance of both factors for higher trophic levels such as insect herbivores is well documented, their impact has rarely been studied in combination. Herbivorous insects have a strong impact on the functioning of ecosystems and it is therefore important to understand how they are affected by eutrophication in high or low diversity environments. We used a grassland biodiversity experiment to investigate the combined effect of fertilization and plant diversity loss on the fitness of the generalist grasshopper Chorthippus parallelus by rearing grasshopper nymphs for four weeks in cages on unfertilized or fertilized (NPK) subplots across a species richness gradient from 1 to 60 plant species. Survival, the number of oothecae, body mass and the number of hatchlings were measured separately for each cage. Plant diversity had no effect on any of the grasshopper fitness measures, neither in unfertilized nor in fertilized plots. NPK-fertilization reduced grasshopper survival but increased body mass of males and reproductive success of the surviving females. Fertilization effects were not mediated by plant community structure, productivity or composition, suggesting that higher food plant quality was one of the main drivers. There was no interaction between plant diversity and fertilization on any of the measures. In conclusion, an increase in eutrophication, in both species-rich and species-poor grasslands, could lead to higher reproductive success and therefore higher abundances of herbivorous insects including insect pests, with fertilization effects dominating plant diversity effects.

Changes in biodiversity and vegetation composition in the central Swiss Alps during the transition from pristine forest to first farming

Aim: We investigate the response of vegetation composition and plant diversity to increasing land clearance, burning and agriculture at the Mesolithic–Neolithic transition (c. 6400–5000 bc) when first farming was introduced. Location: The Valais, a dry alpine valley in Switzerland. Methods: We combine high-resolution pollen, microscopic charcoal and sedimentological data to reconstruct past vegetation, fire and land use. Pollen evenness, rarefaction-based and accumulation-based palynological richness analyses were used to reconstruct past trends in plant diversity. Results: Our results show that from c. 5500 cal. yr bc, slash-and-burn activities created a more open landscape for agriculture, at the expense of Pinus and Betula forests. Land clearance by slash-and-burn promoted diverse grassland ecosystems, while on the long term it reduced woodland and forest diversity, affecting important tree species such as Ulmus and Tilia. Main conclusions: Understanding the resilience of Alpine ecosystems to past disturbance variability is relevant for future nature conservation plans. Our study suggests that forecasted land abandonment in the Alps will lead to pre-Neolithic conditions, with significant biodiversity losses in abandoned grassland ecosystems. Thus, management measures for biodiversity, such as ecological compensation areas, are needed in agricultural landscapes with a millennial history of human impact, such as the non-boreal European lowlands. Our study supports the hypothesis that species coexistence is maximized at an intermediate level of disturbances. For instance, species richness decreased when fire exceeded the quasi-natural variability observed during the Mesolithic times. Under a more natural disturbance regime, rather closed Pinus sylvestris and mixed oak forests would prevail.

Determining the long-term changes in biodiversity and provisioning services along a transect from Central Europe to the Mediterranean

Climate, land use and fire are strong determinants of plant diversity, potentially resulting in local extinctions, including rare endemic and economically valuable species. While climate and land use are decisive for vegetation composition and thus the species pool, fire disturbance can lead to landscape fragmentation, affecting the provisioning of important ecosystem services such as timber and raw natural resources. We use multi-proxy palaeoecological data with high taxonomic and temporal resolution across an environmental gradient to assess the long-term impact of major climate shifts, land use and fire disturbance on past vegetation openness and plant diversity (evenness and richness). Evenness of taxa is inferred by calculating the probability of interspecific encounter (PIE) of pollen and spores and species richness by palynological richness (PRI). To account for evenness distortions of PRI, we developed a new palaeodiversity measure, which is evenness-detrended palynological richness (DE-PRI). Reconstructed species richness increases from north to south regardless of time, mirroring the biodiversity increase across the gradient from temperate deciduous to subtropical evergreen vegetation. Climatic changes after the end of the last ice age contributed to biodiversity dynamics, usually by promoting species richness and evenness in response to warming. The data reveal that the promotion of diverse open-land ecosystems increased when human disturbance became determinant, while forests became less diverse. Our results imply that the today’s biodiversity has been shaped by anthropogenic forcing over the millennia. Future management strategies aiming at a successful conservation of biodiversity should therefore consider the millennia-lasting role of anthropogenic fire and human activities.

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.

Effects of forest management on the diversity of deadwood-inhabiting fungi in Central European forests

Land use and land use change affect deadwood amount, quality and associated biodiversity in forest ecosystems. Old growth or virgin forests, which are exceptionally rare in temperate Europe harbor more deadwood and associated fungal species than managed forests. Whether and how more recent abandonment of management, to reestablish more natural forests, affects deadwood amount and fungal diversity on deadwood is unknown. Our main aim was to compare deadwood amount, characteristics and deadwood inhabiting fungi in differently managed forest types typical for large areas of Central Europe. We sampled deadwood inhabiting fungi on 27 forest plots of 400 m2 each in three geographically distant regions in Germany. Three forest management types, namely managed coniferous, managed deciduous and unmanaged deciduous forests, were represented by nine plots each. In autumn 2008 we collected all fungal fruiting bodies on deadwood >7 cm of diameter. We found deadwood amounts and fungal species numbers in unmanaged forests to be lower than in managed forests, which we attributed to the lack of natural tree death during the short time since management abandonment of usually 10–30 years. However, rarefaction analysis among deadwood items in forest plots indicated a slightly higher species density in unmanaged forests, which may be the first signal of a positive effect on fungal species richness on deadwood after management was abandoned. Although the three study regions span a large geographical gradient, we did not detect differences in the fungal species composition or in deadwood amounts and patterns, which reflects the wide distribution of this group of organisms and points to consistent management procedures among study regions. A very clear composition difference however occurred between deciduous and coniferous wood showing species substrate specialization. We conclude that the amount of deadwood is the main driver of deadwood fungal species richness, and substrate diversity in terms of various decay degrees, deadwood tree species and deadwood size are also important. Thus, to promote species richness of deadwood fungi it is vital to enhance deadwood amounts and diversity

Organic vs. Conventional Grassland Management: Do N-15 and C-13 Isotopic Signatures of Hay and Soil Samples Differ?

Distinguishing organic and conventional products is a major issue of food security and authenticity. Previous studies successfully used stable isotopes to separate organic and conventional products, but up to now, this approach was not tested for organic grassland hay and soil. Moreover, isotopic abundances could be a powerful tool to elucidate differences in ecosystem functioning and driving mechanisms of element cycling in organic and conventional management systems. Here, we studied the delta N-15 and delta C-13 isotopic composition of soil and hay samples of 21 organic and 34 conventional grasslands in two German regions. We also used Delta delta N-15 (delta N-15 plant - delta N-15 soil) to characterize nitrogen dynamics. In order to detect temporal trends, isotopic abundances in organic grasslands were related to the time since certification. Furthermore, discriminant analysis was used to test whether the respective management type can be deduced from observed isotopic abundances. Isotopic analyses revealed no significant differences in delta C-13 in hay and delta C-13 in both soil and hay between management types, but showed that delta C-13 abundances were significantly lower in soil of organic compared to conventional grasslands. delta C-15 values implied that management types did not substantially differ in nitrogen cycling. Only delta C-13 in soil and hay showed significant negative relationships with the time since certification. Thus, our result suggest that organic grasslands suffered less from drought stress compared to conventional grasslands most likely due to a benefit of higher plant species richness, as previously shown by manipulative biodiversity experiments. Finally, it was possible to correctly classify about two third of the samples according to their management using isotopic abundances in soil and hay. However, as more than half of the organic samples were incorrectly classified, we infer that more research is needed to improve this approach before it can be efficiently used in practice.