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.

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.

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.

How do subordinate and dominant species in semi-natural mountain grasslands relate to productivity and land-use change?

Changes in agricultural practices of semi-natural mountain grasslands are expected to modify plant community structure and shift dominance patterns. Using vegetation surveys of 11 sites in semi-natural grasslands of the Swiss Jura and Swiss and French Alps, we determined the relative contribution of dominant, subordinate and transient plant species in grazed and abandoned communities and observed their changes along a gradient of productivity and in response to abandonment of pasturing. The results confirm the humpbacked diversity–productivity relationship in semi-natural grassland, which is due to the increase of subordinate species number at intermediate productivity levels. Grazed communities, at the lower or higher end of the species diversity gradient, suffered higher species loss after grazing abandonment. Species loss after abandonment of pasturing was mainly due to a higher reduction in the number of subordinate species, as a consequence of the increasing proportion of dominant species. When plant biodiversity maintenance is the aim, our results have direct implications for the way grasslands should be managed. Indeed, while intensification and abandonment have been accelerated since few decades, our findings in this multi-site analysis confirm the importance of maintaining intermediate levels of pasturing to preserve biodiversity.

Mean age of carbon in fine roots from temperate forests and grasslands with different management

Fine roots are the most dynamic portion of a plant's root system and a major source of soil organic matter. By altering plant species diversity and composition, soil conditions and nutrient availability, and consequently belowground allocation and dynamics of root carbon (C) inputs, land-use and management changes may influence organic C storage in terrestrial ecosystems. In three German regions, we measured fine root radiocarbon (14C) content to estimate the mean time since C in root tissues was fixed from the atmosphere in 54 grassland and forest plots with different management and soil conditions. Although root biomass was on average greater in grasslands 5.1 ± 0.8 g (mean ± SE, n = 27) than in forests 3.1 ± 0.5 g (n = 27) (p < 0.05), the mean age of C in fine roots in forests averaged 11.3 ± 1.8 yr and was older and more variable compared to grasslands 1.7 ± 0.4 yr (p < 0.001). We further found that management affects the mean age of fine root C in temperate grasslands mediated by changes in plant species diversity and composition. Fine root mean C age is positively correlated with plant diversity (r = 0.65) and with the number of perennial species (r = 0.77). Fine root mean C age in grasslands was also affected by study region with averages of 0.7 ± 0.1 yr (n = 9) on mostly organic soils in northern Germany and of 1.8 ± 0.3 yr (n = 9) and 2.6 ± 0.3 (n = 9) in central and southern Germany (p < 0.05). This was probably due to differences in soil nutrient contents and soil moisture conditions between study regions, which affected plant species diversity and the presence of perennial species. Our results indicate more long-lived roots or internal redistribution of C in perennial species and suggest linkages between fine root C age and management in grasslands. These findings improve our ability to predict and model belowground C fluxes across broader spatial scales.

Direct and indirect associations between plant species richness and productivity in grasslands: regional differences preclude simple generalization of productivity-biodiversity relationships

Plant species richness of permanent grasslands has often been found to be significantly associated with productivity. Concentrations of nutrients in biomass can give further insight into these productivity- plant species richness relationships, e.g. by reflecting land use or soil characteristics. However, the consistency of such relationships across different regions has rarely been taken into account, which might significantly compromise our potential for generalization. We recorded plant species richness and measured above-ground biomass and concentrations of nutrients in biomass in 295 grasslands in three regions in Germany that differ in soil and climatic conditions. Structural equation modelling revealed that nutrient concentrations were mostly indirectly associated with plant species richness via biomass production. However, negative associations between the concentrations of different nutrients and biomass and plant species richness differed considerably among regions. While in two regions, more than 40% of the variation in plant species richness could be attributed to variation in biomass, K, P, and to some degree also N concentrations, in the third region only 15% of the variation could be explained in this way. Generally, highest plant species richness was recorded in grasslands where N and P were co-limiting plant growth, in contrast to N or K (co-) limitation. But again, this pattern was not recorded in the third region. While for two regions land-use intensity and especially the application of fertilizers are suggested to be the main drivers causing the observed negative associations with productivity, in the third region the little variance accounted for, low species richness and weak relationships implied that former intensive grassland management, ongoing mineralization of peat and fluctuating water levels in fen grasslands have overruled effects of current land-use intensity and productivity. Finally, we conclude that regional replication is of major importance for studies seeking general insights into productivity-diversity relationships.

Dry grasslands of NW Bulgarian mountains: first insights into diversity, ecology and syntaxonomy

We present the data of the 3rdresearch expedition of the European Dry Grasslands Group (EDGG), which was conducted in 2011 in two contrasting areas in NW Bulgarian mountains. The aim was to collect plot data for comparing Bulgarian dry grasslands with those of other parts of Europe in terms of syntaxonomy and biodiversity. We sampled 15 nested-plot series (0.0001–100 m²) and 68 normal plots(10 m²) covering the full variety of dry grassland types occurring in the Vratsa area (Balkan Mts.) and the Koprivshtitsa area (Sredna Gora Mt.). In the plots all vascular plants, terricolous non-vascular plants and a set of soil and other environmental parameters were determined. By applying modified TWIN-SPAN, we distinguished 10 floristically well characterised vegetation types at the association level. After comparison with the regional and European literature, we propose to place them within three classes and five orders: Festuco-Brometea with the orders Stipo pulcherrimae-Festucetalia pallentis (xerophilous dry grasslands of base-rich rocks; alliance Saturejion montanae), Brachypodietalia pinnate (meso-xeric, basiphilous grasslands; alliances Cirsio-Brachypodion pinnate and Chyrsopogono grylli-Danthonion calycinae),Calluno-Ulicetea with the order Nardetalia stricae (lowland to montane Nardus swards; alliance Violion caninae), and Koelerio-Corynephoretea with the orders Sedo-Scleranthetalia (open communities of skeleton-rich, acidic soils; alliance unclear) and Trifolioarvensis-Festucetalia ovinae(closed, meso-xeric, acidophilous grasslands; alliance Armerio rumelicae-Potentillion). The Violion caninae with the association Festuco rubrae-Genistelletum sagittalisis reported from Bulgaria for the first time, while the two occurring Koelerio-Corynephoretea communities are described as new associations (Cetrario aculeatae-Plantaginetum radicatae, Plantagini radicatae-Agrostietum capillaris). According to DCA the main floristic gradient was largely determined by soil conditions, differentiating the Festuco-Brometea communities on soils with high pH and high humus content from the Koelerio-Corynephoretea communities on acidic, humus-poor soils, while the Calluno-Ulicetea stands are the connecting link. At 10 m² Festuco-Brometea and Calluno-Ulicetea stands were richer in species across all investigated taxa and in vascular plants than Koelerio-Corynephoretea stands; the latter were richest in lichen species, while bryophyte richness did not differ significantly among syntaxa. Among the Bulgarian classes, the species-area relationships tended to be steepest in the Festuco-Brometea (i.e. highest beta diversity), but both alpha and beta diversity clearly fell behind the Festuco-Brometea communities in the Transylvanian Plateau, Romania, located less than 500 km north of the study region. Overall, our study contributes to a more adequate placement of the Bulgarian dry grasslands in the European syntaxonomic system and provides valuable data for large-scale analyses of biodiversity patterns

Designing forest biodiversity experiments: general considerations illustrated by a new large experiment in subtropical China

1. Biodiversity-ecosystem functioning (BEF) experiments address ecosystem-level consequences of species loss by comparing communities of high species richness with communities from which species have been gradually eliminated. BEF experiments originally started with microcosms in the laboratory and with grassland ecosystems. A new frontier in experimental BEF research is manipulating tree diversity in forest ecosystems, compelling researchers to think big and comprehensively. 2. We present and discuss some of the major issues to be considered in the design of BEF experiments with trees and illustrate these with a new forest biodiversity experiment established in subtropical China (Xingangshan, Jiangxi Province) in 2009/2010. Using a pool of 40 tree species, extinction scenarios were simulated with tree richness levels of 1, 2, 4, 8 and 16 species on a total of 566 plots of 25.8x25.8m each. 3. The goal of this experiment is to estimate effects of tree and shrub species richness on carbon storage and soil erosion; therefore, the experiment was established on sloped terrain. The following important design choices were made: (i) establishing many small rather than fewer larger plots, (ii) using high planting density and random mixing of species rather than lower planting density and patchwise mixing of species, (iii) establishing a map of the initial ecoscape' to characterize site heterogeneity before the onset of biodiversity effects and (iv) manipulating tree species richness not only in random but also in trait-oriented extinction scenarios. 4. Data management and analysis are particularly challenging in BEF experiments with their hierarchical designs nesting individuals within-species populations within plots within-species compositions. Statistical analysis best proceeds by partitioning these random terms into fixed-term contrasts, for example, species composition into contrasts for species richness and the presence of particular functional groups, which can then be tested against the remaining random variation among compositions. 5. We conclude that forest BEF experiments provide exciting and timely research options. They especially require careful thinking to allow multiple disciplines to measure and analyse data jointly and effectively. Achieving specific research goals and synergy with previous experiments involves trade-offs between different designs and requires manifold design decisions.

Comparison of inorganic nitrogen uptake dynamics following snowmelt and at peak biomass in subalpine grasslands

Subalpine grasslands are highly seasonal environments and likely subject to strong variability in nitrogen (N) dynamics. Plants and microbes typically compete for N acquisition during the growing season and particularly at plant peak biomass. During snowmelt, plants could potentially benefit from a decrease in competition by microbes, leading to greater plant N uptake associated with active growth and freeze-thaw cycles restricting microbial growth. In managed subalpine grasslands, we expect these interactions to be influenced by recent changes in agricultural land use, and associated modifications in plant and microbial communities. At several subalpine grasslands in the French Alps, we added pulses of 15N to the soil at the end of snowmelt, allowing us to compare the dynamics of inorganic N uptake in plants and microbes during this period with that previously reported at the peak biomass in July. In all grasslands, while specific shoot N translocation (per g of biomass) of dissolved inorganic nitrogen (DIN) was two to five times greater at snowmelt than at peak biomass, specific microbial DIN uptakes were similar between the two sampling dates. On an area basis, plant communities took more DIN than microbial communities at the end of snowmelt when aboveground plant biomasses were at least two times lower than at peak biomass. Consequently, inorganic N partitioning after snowmelt switches in favor of plant communities, allowing them to support their growing capacities at this period of the year. Seasonal differences in microbial and plant inorganic N-related dynamics were also affected by past (terraced vs. unterraced) rather than current (mown vs. unmown) land use. In terraced grasslands, microbial biomass N remained similar across seasons, whereas in unterraced grasslands, microbial biomass N was higher and microbial C : N lower at the end of snowmelt as compared to peak biomass. Further investigations on microbial community composition and their organic N uptake dynamics are required to better understand the decrease in microbial DIN uptake.

Stalagmite water content as a proxy for drip water supply in tropical and subtropical areas

In this pilot study water was extracted from samples of two Holocene stalagmites from Socotra Island, Yemen, and one Eemian stalagmite from southern continental Yemen. The amount of water extracted per unit mass of stalagmite rock, termed "water yield" hereafter, serves as a measure of its total water content. Based on direct correlation plots of water yields and δ18Ocalcite and on regime shift analyses, we demonstrate that for the studied stalagmites the water yield records vary systematically with the corresponding oxygen isotopic compositions of the calcite (δ18Ocalcite). Within each stalagmite lower δ18Ocalcite values are accompanied by lower water yields and vice versa. The δ18Ocalcite records of the studied stalagmites have previously been interpreted to predominantly reflect the amount of rainfall in the area; thus, water yields can be linked to drip water supply. Higher, and therefore more continuous drip water supply caused by higher rainfall rates, supports homogeneous deposition of calcite with low porosity and therefore a small fraction of water-filled inclusions, resulting in low water yields of the respective samples. A reduction of drip water supply fosters irregular growth of calcite with higher porosity, leading to an increase of the fraction of water-filled inclusions and thus higher water yields. The results are consistent with the literature on stalagmite growth and supported by optical inspection of thin sections of our samples. We propose that for a stalagmite from a dry tropical or subtropical area, its water yield record represents a novel paleo-climate proxy recording changes in drip water supply, which can in turn be interpreted in terms of associated rainfall rates.

Grazing response patterns indicate isolation of semi-natural European grasslands

Identifying drivers of species diversity is a major challenge in understanding and predicting the dynamics of species-rich semi-natural grasslands. In particular in temperate grasslands changes in land use and its consequences, i.e. increasing fragmentation, the on-going loss of habitat and the declining importance of regional processes such as seed dispersal by livestock, are considered key drivers of the diversity loss witnessed within the last decades. It is a largely unresolved question to what degree current temperate grassland communities already reflect a decline of regional processes such as longer distance seed dispersal. Answering this question is challenging since it requires both a mechanistic approach to community dynamics and a sufficient data basis that allows identifying general patterns. Here, we present results of a local individual- and trait-based community model that was initialized with plant functional types (PFTs) derived from an extensive empirical data set of species-rich grasslands within the `Biodiversity Exploratories' in Germany. Driving model processes included above- and belowground competition, dynamic resource allocation to shoots and roots, clonal growth, grazing, and local seed dispersal. To test for the impact of regional processes we also simulated seed input from a regional species pool. Model output, with and without regional seed input, was compared with empirical community response patterns along a grazing gradient. Simulated response patterns of changes in PFT richness, Shannon diversity, and biomass production matched observed grazing response patterns surprisingly well if only local processes were considered. Already low levels of additional regional seed input led to stronger deviations from empirical community pattern. While these findings cannot rule out that regional processes other than those considered in the modeling study potentially play a role in shaping the local grassland communities, our comparison indicates that European grasslands are largely isolated, i.e. local mechanisms explain observed community patterns to a large extent.