Recruitment limitation constrains local species richness and productivity in dry grassland

Species coexistence and local-scale species richness are limited by the availability of seeds and microsites for germination and establishment. We conducted a seed addition experiment in seminatural grassland at three sites in southern Switzerland and repeated the experiment in two successive years to evaluate various circumstances under which seed limitation and establishment success affect community functioning. A collection of 144000 seeds of 22 meadow species including grasses and forbs of local provenance was gathered, and seeds were individually sown in a density that resembled natural seed rain. The three communities were seed limited. Three years after sowing, single species varied in emergence (0–50%), survival (0–69%), and establishment rates (0–27%). One annual and 13 perennial species reached reproductive stage. Low establishment at one site and reduced growth at another site indicated stronger microsite limitation compared to the third site. Recruitment was influenced by differences in abiotic environmental conditions between sites (water availability, soil minerals) and by within-site differences in biotic interaction (competition). At the least water-limited site, sowing resulted in an increase in phytomass due to establishment of short-lived perennials in the second and third years after sowing. This increase persisted over the following two years due to establishment of longer-lived perennials. After sowing in a wetter year with higher phytomass, however, productivity did not increase, because higher intensity of competition in an early phase of establishment resulted in less vigorous plants later on. Due to the generally favorable weather conditions during this study, sowing year had a small effect on numbers of established individuals over all species. Recruitment limitation can thus constrain local-scale species richness and productivity, either by a lack of seeds or by reduced seedling growth, likely due to competition from the established vegetation.

Interacting effects of fertilization, mowing and grazing on plant species diversity of 1500 grasslands in Germany differ between regions

The relationship of different types of grassland use with plant species richness and composition (functional groups of herbs, legumes, and grasses) has so far been studied at small regional scales or comprising only few components of land use. We comprehensively studied the relationship between abandonment, fertilization, mowing intensity, and grazing by different livestock types on plant diversity and composition of 1514 grassland sites in three regions in North-East, Central and South-West Germany. We further considered environmental site conditions including soil type and topographical situation. Fertilized grasslands showed clearly reduced plant species diversity (−15% plant species richness, −0.1 Shannon diversity on fertilized grasslands plots of 16 m2) and changed composition (−3% proportion of herb species), grazing had the second largest effects and mowing the smallest ones. Among the grazed sites, the ones grazed by sheep had higher than average species richness (+27%), and the cattle grazed ones lower (−42%). Further, these general results were strongly modulated by interactions between the different components of land use and by regional context: land-use effects differed largely in size and sometimes even in direction between regions. This highlights the importance of comparing different regions and to involve a large number of plots when studying relationships between land use and plant diversity. Overall, our results show that great caution is necessary when extrapolating results and management recommendations to other regions.

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.

Weeds and endangered herbs have unforeseen dispersal helpers in the agri-environment: gastropods and earthworms

Agri-environmental schemes involving organic farming or set-aside management aim at promoting biodiversity and restoring ecosystem functioning in agrarian landscapes. Application of pesticides in these crop fields is strongly regulated facilitating the spread of weeds but also allowing for the establishment of endangered herbs and a variety of animals.Recent studies found gastropods and earthworms to be legitimate dispersers of seeds of wild plants. We assumed that both groups also playa significant role in the spread and establishment of wild plants within crop fields. Therefore, we are conducting a series of experiments in three different study systems on (1) the role of earthworms and gastropods as dispersers of rare herbs and weeds in an organic rye field in Germany, (2) the seed feeding behavior of gastropods of plants sown in fallow ground in Switzerland, and (3) weed dispersal in irrigated rice fields by golden apple snails in the Philippines.

Allelopathic effects of three plant invaders on germination of native species: a field study

The ability of some invasive plant species to produce biochemical compounds toxic to native species, called allelopathy, is thought to be one of the reasons for their success when introduced to a novel range, an idea known as the Novel Weapons Hypothesis. However, support for this hypothesis mainly comes from bioassays and experiments conducted under controlled environments, whereas field evidence is rare. In a field experiment, we investigated whether three plant species invasive in Europe, Solidago gigantea, Impatiens glandulifera and Erigeron annuus, inhibit the germination of native species through allelopathy more than an adjacent native plant community. At three sites for each invasive species, we compared the germination of native species that were sown on invaded and non-invaded plots. Half of these plots were amended with activated carbon to reduce the influence of potential allelopathic compounds. The germination of sown seeds and of seeds from the seedbank was monitored over a period of 9 weeks. Activated carbon generally enhanced seed germination. This effect was equally pronounced in invaded and adjacent non-invaded plots, indicating that invasive species do not suppress germination more than a native plant community. In addition, more seeds germinated from the seedbank on invaded than on non-invaded soil, probably due to previous suppression of germination by the invasive species. Our field study does not provide evidence for the Novel Weapons Hypothesis with respect to the germination success of natives. Instead, our results suggest that if invasive species release allelopathic compounds that suppress germination, they do so to a similar degree as the native plant community.

Biodiversity effects on nitrate concentrations in soil solution: a Bayesian model

Ecosystems are faced with high rates of species loss which has consequences for their functions and services. To assess the effects of plant species diversity on the nitrogen (N) cycle, we developed a model for monthly mean nitrate (NO3-N) concentrations in soil solution in 0-30 cm mineral soil depth using plant species and functional group richness and functional composition as drivers and assessing the effects of conversion of arable land to grassland, spatially heterogeneous soil properties, and climate. We used monthly mean NO3-N concentrations from 62 plots of a grassland plant diversity experiment from 2003 to 2006. Plant species richness (1-60) and functional group composition (1-4 functional groups: legumes, grasses, non-leguminous tall herbs, non-leguminous small herbs) were manipulated in a factorial design. Plant community composition, time since conversion from arable land to grassland, soil texture, and climate data (precipitation, soil moisture, air and soil temperature) were used to develop one general Bayesian multiple regression model for the 62 plots to allow an in-depth evaluation using the experimental design. The model simulated NO3-N concentrations with an overall Bayesian coefficient of determination of 0.48. The temporal course of NO3-N concentrations was simulated differently well for the individual plots with a maximum plot-specific Nash-Sutcliffe Efficiency of 0.57. The model shows that NO3-N concentrations decrease with species richness, but this relation reverses if more than approx. 25 % of legume species are included in the mixture. Presence of legumes increases and presence of grasses decreases NO3-N concentrations compared to mixtures containing only small and tall herbs. Altogether, our model shows that there is a strong influence of plant community composition on NO3-N concentrations.

Impacts of drought stress on water relations and carbon assimilation in grassland species at different altitudes

Grassland is an important ecosystem type which is not only used agriculturally, but also covers sites which cannot be used for other purposes, e.g. in very steep locations or above timberlines. Prolonged summer droughts in the mid-term future, as are predicted for Central Europe, are expected to have a major impact on such ecosystems. To address this topic, rainfall exclusion via shelters was performed on three grassland sites at different altitudes (393, 982 and 1978 m above sea level) in Switzerland. Diurnal drought treatment effects were studied at each study site on a completely sunny day towards the end of an 8–10 week shelter period. Ecophysiological parameters including gas exchange (An, gs and intrinsic WUE) and chlorophyll a fluorescence (Fv/Fm, ΦPSII and NPQ) were considered for several species. The lowland and the Alpine field site were more strongly affected by soil drought than the pre-Alpine site. At all sites, grasses showed different patterns of reductions in stomatal conductance under soil drought compared to legumes and forbs. In addition, grasses were significantly more affected by reductions in assimilation rates at all sites. Time courses of reductions in assimilation rates relative to controls differed between species at the Alpine site, as some species showed reduced assimilation rates at this site in the early morning. Thus, similar rainfall exclusion treatments can trigger different reactions in various species at different sites, which might not become obvious during mere midday measurements. Overall, results suggest strong impacts of prolonged summer drought on grassland net photosynthesis especially at the Alpine site and, within sites, for grasses

Plant diversity effects on the water balance of an experimental grassland

In the literature, contrasting effects of plant species richness on the soil water balance are reported. Our objective was to assess the effects of plant species and functional richness and functional identity on soil water contents and water fluxes in the experimental grassland of the Jena Experiment. The Jena Experiment comprises 86 plots on which plant species richness (0, 1, 2, 4, 8, 16, and 60) and functional group composition (zero to four functional groups: legumes, grasses, tall herbs, and small herbs) were manipulated in a factorial design. We recorded meteorological data and soil water contents of the 0·0–0·3 and 0·3–0·7 m soil layers and calculated actual evapotranspiration (ETa), downward flux (DF), and capillary rise with a soil water balance model for the period 2003–2007. Missing water contents were estimated with a Bayesian hierarchical model. Species richness decreased water contents in subsoil during wet soil conditions. Presence of tall herbs increased soil water contents in topsoil during dry conditions and decreased soil water contents in subsoil during wet conditions. Presence of grasses generally decreased water contents in topsoil, particularly during dry phases; increased ETa and decreased DF from topsoil; and decreased ETa from subsoil. Presence of legumes, in contrast, decreased ETa and increased DF from topsoil and increased ETa from subsoil. Species richness probably resulted in complementary water use. Specific functional groups likely affected the water balance via specific root traits (e.g. shallow dense roots of grasses and deep taproots of tall herbs) or specific shading intensity caused by functional group effects on vegetation cover. Copyright © 2013 John Wiley & Sons, Ltd.

Combined effect of soil bund with biological soil and water conservation measures in the northwestern Ethiopian highlands

Excessive runoff and soil erosion in the upper Blue Nile Basin poses a threat that has attracted the attention of the Ethiopian government because of the serious on-site effects in addition to downstream effects, such as the siltation of water harvesting structures and reservoirs. The objective of the study was to evaluate and recommend effective biophysical soil and water conservation measure(s) in the Debre Mewi watershed, about 30 km south of the Lake Tana. Six conservation measures were evaluated for their effects on runoff, soil loss, and forage yield using runoff plots. There was a significant difference between treatments for both runoff and soil loss. The four-year average annual soil loss in the different plots ranged from 26 to 71 t ha−1, and total runoff ranged from 180 to 302 mm, while annual rainfall varied between 854 mm in 2008 and 1247 mm in 2011. Soil bund combined with elephant grass had the lowest runoff and soil loss as compared to the other treatments, whereas the untreated control plot had the highest for both parameters. As an additional benefit, 2.8 and 0.7 t ha−1 year−1 of dried forage was obtained from elephant and local grasses, respectively. Furthermore, it was found that soil bund combined with Tephrosia increased soil organic matter by 13% compared to the control plot. Soil bund efficiency was significantly enhanced by combining them with biological measures and improved farmers’ perception of soil and water conservation measures.

Biotic and Abiotic Properties Mediating Plant Diversity Effects on Soil Microbial Communities in an Experimental Grassland

Plant diversity drives changes in the soil microbial community which may result in alterations in ecosystem functions. However, the governing factors between the composition of soil microbial communities and plant diversity are not well understood. We investigated the impact of plant diversity (plant species richness and functional group richness) and plant functional group identity on soil microbial biomass and soil microbial community structure in experimental grassland ecosystems. Total microbial biomass and community structure were determined by phospholipid fatty acid (PLFA) analysis. The diversity gradient covered 1, 2, 4, 8, 16 and 60 plant species and 1, 2, 3 and 4 plant functional groups (grasses, legumes, small herbs and tall herbs). In May 2007, soil samples were taken from experimental plots and from nearby fields and meadows. Beside soil texture, plant species richness was the main driver of soil microbial biomass. Structural equation modeling revealed that the positive plant diversity effect was mainly mediated by higher leaf area index resulting in higher soil moisture in the top soil layer. The fungal-to-bacterial biomass ratio was positively affected by plant functional group richness and negatively by the presence of legumes. Bacteria were more closely related to abiotic differences caused by plant diversity, while fungi were more affected by plant-derived organic matter inputs. We found diverse plant communities promoted faster transition of soil microbial communities typical for arable land towards grassland communities. Although some mechanisms underlying the plant diversity effect on soil microorganisms could be identified, future studies have to determine plant traits shaping soil microbial community structure. We suspect differences in root traits among different plant communities, such as root turnover rates and chemical composition of root exudates, to structure soil microbial communities.