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.

Agricultural land use and biodiversity in the Alps - How cultural tradition and socioeconomically motivated changes are shaping grassland biodiversity in the Swiss Alps

Alpine grasslands are ecosystems with a great diversity of plant species. However, little is known about other levels of biodiversity, such as landscape diversity, diversity of biological interactions of plants with herbivores or fungal pathogens, and genetic diversity. We therefore explored natural and anthropogenic determinants of grassland biodiversity at several levels of biological integration, from the genetic to the landscape level in the Swiss Alps. Differences between cultural traditions (Romanic, Germanic, and Walser) turned out to still affect land use diversity and thus landscape diversity. Increasing land use diversity, in turn, increased plant species diversity per village. However, recent land use changes have reduced this diversity. Within grassland parcels, plant species diversity was higher on unfertilized mown grasslands than on fertilized or grazed ones. Most individual plants were affected by herbivores and fungal leaf pathogens, reflecting that parcels harbored a great diversity of herbivores and pathogens. However, as plant damage by herbivores and pathogens was not severe, conserving these biological interactions among plants is hardly compromising agricultural goals. A common-garden experiment revealed genetic differentiation of the important fodder grass Poa alpina between mown and grazed sites, suggesting adaptation. Per-village genetic diversity of Poa alpina was greater in villages with higher land use diversity, analogous to the higher plant species diversity there. Overall, landscape diversity and biodiversity within grassland parcels are currently declining. As this contradicts the intention of Swiss law and international agreements, financial incentives need to be re-allocated and should focus on promoting high biodiversity at the local and the landscape level. At the same time, this will benefit landscape attractiveness for tourists and help preserve a precious cultural heritage in the Swiss Alps.

Sampling-Design Effects on Properties of Species-Area Relationships - A Case Study from Estonian Dry Grassland Communities

Despite widespread use of species-area relationships (SARs), dispute remains over the most representative SAR model. Using data of small-scale SARs of Estonian dry grassland communities, we address three questions: (1) Which model describes these SARs best when known artifacts are excluded? (2) How do deviating sampling procedures (marginal instead of central position of the smaller plots in relation to the largest plot; single values instead of average values; randomly located subplots instead of nested subplots) influence the properties of the SARs? (3) Are those effects likely to bias the selection of the best model? Our general dataset consisted of 16 series of nested-plots (1 cm(2)-100 m(2), any-part system), each of which comprised five series of subplots located in the four corners and the centre of the 100-m(2) plot. Data for the three pairs of compared sampling designs were generated from this dataset by subsampling. Five function types (power, quadratic power, logarithmic, Michaelis-Menten, Lomolino) were fitted with non-linear regression. In some of the communities, we found extremely high species densities (including bryophytes and lichens), namely up to eight species in 1 cm(2) and up to 140 species in 100 m(2), which appear to be the highest documented values on these scales. For SARs constructed from nested-plot average-value data, the regular power function generally was the best model, closely followed by the quadratic power function, while the logarithmic and Michaelis-Menten functions performed poorly throughout. However, the relative fit of the latter two models increased significantly relative to the respective best model when the single-value or random-sampling method was applied, however, the power function normally remained far superior. These results confirm the hypothesis that both single-value and random-sampling approaches cause artifacts by increasing stochasticity in the data, which can lead to the selection of inappropriate models.

The role of landuse and natural determinants for grassland vegetation composition in the Swiss Alps

The Alps provide a high habitat diversity for plant species, structured by broad- and fine-scale abiotic site conditions. In man-made grasslands, vegetation composition is additionally affected by the type of landuse. We recorded vegetation composition in 216 parcels of grassland in 12 municipalities representing an area of 170 x 70 km in the south-eastern part of the Swiss Alps. Each parcel was characterized by a combination of altitudinal level (valley, intermediate, alp). traditional landuse (mown. grazed), current management (mown, grazed, abandoned). and Fertilization (unfertilized, fertilized). For each parcel we also assessed the abiotic factors aspect, slope, pH value, and geographic coordinates, and for each municipality annual precipitation and its cultural tradition. We analysed vegetation composition using (i) variation partitioning in RDA. (ii) cover of graminoids. non-legume forbs, and legumes, and (iii) dominance and frequency of species. Species composition was determined by, in decreasing order of variation explained. landuse, broad-scale abiotic factors, fine-scale abiotic factors. and cultural tradition. Current socio-economically motivated landuse changes, such as grazing of unfertilized former meadows or their abandonment, strongly affect vegetation composition. In our study, the frequency of characteristic meadow species was significantly smaller in grazed and even smaller in abandoned parcels than in still mown ones, suggesting less severe consequences of grazing for vegetation composition than of abandonment. Therefore. low-intensity grazing and mowing every few years should be considered valuable conservation alternatives to abandonment. Furthermore. because each landuse type was characterized by different species. a high variety of landuse types should be promoted to preserve plant species diversity in Alpine grasslands. (C) 2007 Gesellschaft fur Okologie. Published by Elsevier GmbH. All rights reserved.

Species mobility and frequency dynamics in a Swiss limestone grassland

Community dynamics in a calcareous grassland (Mesobrometum) in Egerkingen (Jura mountains, Switzerland) were investigated for 53 non-woody species in 25 1-m2 plots over 6 years. 50 0.0 1-m2 subplots per plot were recorded. The derived variables were spatial frequency, temporal frequency, frequency fluctuation, turnover, and cumulative frequency (each species), and cumulative species richness (all species). Spectra for 53 species of all variables were different for the two investigated spatial scales (0.0 1 m2, 1 m2). The comparison with other investigations of similar grass lands showed that the behaviour of some species is specific for this type of vegetation in general (e.g. Achillea millefolium, Arrhenatherum elatius, Bromus erectus ), but most species behaved in a stand-specific way, i.e. they may play another (similar or completely different) role in another grassland stand. Six spatio-temporal patterns were defined across species. To understand community dynamics, not only the dynamics of mobility but also of frequency fluctuations and spatial distribution of the species are fundamental. In addition, the understanding of temporal behaviour of all species present should be included. Averages always hide important information of vegetation dynamics, as was shown by the present investigation.

Response of temperate grasslands at different altitudes to simulated summer drought differed but scaled with annual precipitation

Water is an important resource for plant life. Since climate scenarios for Switzerland predict an average reduction of 20% in summer precipitation until 2070, understanding ecosystem responses to water shortage, e.g. in terms of plant productivity, is of major concern. Thus, we tested the effects of simulated summer drought on three managed grasslands along an altitudinal gradient in Switzerland from 2005 to 2007, representing typical management intensities at the respective altitude. We assessed the effects of experimental drought on above- and below-ground productivity, stand structure (LAI and vegetation height) and resource use (carbon and water). Responses of community above-ground productivity to reduced precipitation input differed among the three sites but scaled positively with total annual precipitation at the sites (R2=0.85). Annual community above-ground biomass productivity was significantly reduced by summer drought at the alpine site receiving the least amount of annual precipitation, while no significant decrease (rather an increase) was observed at the pre-alpine site receiving highest precipitation amounts in all three years. At the lowland site (intermediate precipitation sums), biomass productivity significantly decreased in response to drought only in the third year, after showing increased abundance of a drought tolerant weed species in the second year. No significant change in below-ground biomass productivity was observed at any of the sites in response to simulated summer drought. However, vegetation carbon isotope ratios increased under drought conditions, indicating an increase in water use efficiency. We conclude that there is no general drought response of Swiss grasslands, but that sites with lower annual precipitation seem to be more vulnerable to summer drought than sites with higher annual precipitation, and thus site-specific adaptation of management strategies will be needed, especially in regions with low annual precipitation.

An economic assessment of drought effects on three grassland systems in Switzerland

This paper analyzes the economic impacts of summer drought on Swiss grassland production. We combine field trial data from drought experiments in three different grasslands in Switzerland with site-specific information on economic costs and benefits. The analysis focuses on the economic implications of drought effects on grassland yields as well as grassland composition. In agreement with earlier studies, we found rather heterogeneous yield effects of drought on Swiss grassland systems, with significantly reduced yields as a response to drought at the lowland and sub-alpine sites, but increased yields at the wetter pre-alpine site. Relative yield losses were highest at the sub-alpine site (with annual yield losses of up to 37 %). However, because income from grassland production at extensive sites relies to a large extent on ecological direct payments, even large yield losses had only limited implications in terms of relative profit reductions. In contrast, negative drought impacts at the most productive, intensively managed lowland site were dominant, with average annual drought-induced profit margin reductions of about 28 %. This is furthermore emphasized if analyzing the farm level perspective of drought impacts. Combining site-specific effects at the farm level, we found that in particular farms with high shares of lowland grassland sites suffer from summer droughts in terms of farm-level fodder production and profit margins. Moreover, our results showed that the higher competitiveness of weeds (broad-leaved dock) under drought conditions will require increasing attention on weed control measures in future grassland production systems. Taking into account that the risk of drought occurrence is expected to increase in the coming years, additional instruments to cope with drought risks in fodder production and finally farmers’ income have to be developed.

Does organic grassland farming benefit plant and arthropod diversity at the expense of yield and soil fertility?

Organic management is one of the most popular strategies to reduce negative environmental impacts of intensive agriculture. However, little is known about benefits for biodiversity and potential worsening of yield under organic grasslands management across different grassland types, i.e. meadow, pasture and mown pasture. Therefore, we studied the diversity of vascular plants and foliage-living arthropods (Coleoptera, Araneae, Heteroptera, Auchenorrhyncha), yield, fodder quality, soil phosphorus concentrations and land-use intensity of organic and conventional grasslands across three study regions in Germany. Furthermore, all variables were related to the time since conversion to organic management in order to assess temporal developments reaching up to 18 years. Arthropod diversity was significantly higher under organic than conventional management, although this was not the case for Araneae, Heteroptera and Auchenorrhyncha when analyzed separately. On the contrary, arthropod abundance, vascular plant diversity and also yield and fodder quality did not considerably differ between organic and conventional grasslands. Analyses did not reveal differences in the effect of organic management among grassland types. None of the recorded abiotic and biotic parameters showed a significant trend with time since transition to organic management, except soil organic phosphorus concentrations which decreased with time. This implies that permanent grasslands respond slower and probably weaker to organic management than crop fields do. However, as land-use intensity and inorganic soil phosphorus concentrations were significantly lower in organic grasslands, overcoming seed and dispersal limitation by re-introducing plant species might be needed to exploit the full ecological potential of organic grassland management. We conclude that although organic management did not automatically increase the diversity of all studied taxa, it is a reasonable and useful way to support agro-biodiversity.

Community mean traits as additional indicators to monitor effects of land-use intensity on grassland plant diversity

Semi-natural grasslands, biodiversity hotspots in Central-Europe, suffer from the cessation of traditional land-use. Amount and intensity of these changes challenge current monitoring frameworks typically based on classic indicators such as selected target species or diversity indices. Indicators based on plant functional traits provide an interesting extension since they reflect ecological strategies at individual and ecological processes at community levels. They typically show convergent responses to gradients of land-use intensity over scales and regions, are more directly related to environmental drivers than diversity components themselves and enable detecting directional changes in whole community dynamics. However, probably due to their labor- and cost intensive assessment in the field, they have been rarely applied as indicators so far. Here we suggest overcoming these limitations by calculating indicators with plant traits derived from online accessible databases. Aiming to provide a minimal trait set to monitor effects of land-use intensification on plant diversity we investigated relationships between 12 community mean traits, 2 diversity indices and 6 predictors of land-use intensity within grassland communities of 3 different regions in Germany (part of the German ‘Biodiversity Exploratory’ research network). By standardization of traits and diversity measures, use of null models and linear mixed models we confirmed (i) strong links between functional community composition and plant diversity, (ii) that traits are closely related to land-use intensity, and (iii) that functional indicators are equally, or even more sensitive to land-use intensity than traditional diversity indices. The deduced trait set consisted of 5 traits, i.e., specific leaf area (SLA), leaf dry matter content (LDMC), seed release height, leaf distribution, and onset of flowering. These database derived traits enable the early detection of changes in community structure indicative for future diversity loss. As an addition to current monitoring measures they allow to better link environmental drivers to processes controlling community dynamics.

Determinants of plant establishment success in a multispecies introduction experiment with native and alien species

Determinants of plant establishment and invasion are a key issue in ecology and evolution. Although establishment success varies substantially among species, the importance of species traits and extrinsic factors as determinants of establishment in existing communities has remained difficult to prove in observational studies because they can be confounded and mask each other. Therefore, we conducted a large multispecies field experiment to disentangle the relative importance of extrinsic factors vs. species characteristics for the establishment success of plants in grasslands. We introduced 48 alien and 45 native plant species at different seed numbers into multiple grassland sites with or without experimental soil disturbance and related their establishment success to species traits assessed in five independent multispecies greenhouse experiments. High propagule pressure and high seed mass were the most important factors increasing establishment success in the very beginning of the experiment. However, after 3 y, propagule pressure became less important, and species traits related to biotic interactions (including herbivore resistance and responses to shading and competition) became the most important drivers of success or failure. The relative importance of different traits was environment-dependent and changed over time. Our approach of combining a multispecies introduction experiment in the field with trait data from independent multispecies experiments in the greenhouse allowed us to detect the relative importance of species traits for early establishment and provided evidence that species traits—fine-tuned by environmental factors—determine success or failure of alien and native plants in temperate grasslands.