Plant community controls small-scale variation in nutrient stoichiometry in a Patagonian peatland

Elemental stoichiometry of plant litter is typically interpreted to reflect nutrient availability and limitation, e.g. the N:P ratio indicates whether plant growth is N- or P-limited and might point towards the presence of N-fixation. However, in the case of plant litter and peat organic matter, resorption of nutrients during senescence, and preferential loss of nutrients during decomposition have to be taken into account. Here we study how small scale variability in species composition within peatlands affects the stoichiometry and long term apparent uptake rates of nutrients (C, N, P, K, S, Ca, Mg) in an ombrotrophic peatland in southern Patagonia. Assuming that nutrient availability is similar within one site, observed variation should be driven by vegetation and decomposition processes linked to microtopography. We studied a transect spanning 800 m. where the vegetation changed from cushion plant (Astelia pumila) dominated, to graminoid dominated, ending in Sphagnum magellanicum dominated. From six peat cores along this transect we analysed nutrient concentrations by X-ray fluorescence. The peat decomposition state, expressed as FTIR-humification index, was the best predictor of stoichiometric variation (particularly ratios C:N, C:S, and N:P), followed by current plant species composition. Comparison of average peat core stoichiometry across the transect showed that C:N and C:S ratios were larger in Sphagnum cores than cushion plant and graminoid cores (C:N 56±14 vs. 38±6; C:S 312±61 vs. 268±57; respectively), controlled by lower decomposition state in Sphagnum cores and larger C:N ratios in living biomass of Sphagnum vs. A. pumila. Larger N:P ratios in cushion plant and graminoid vs. Sphagnum cores (N:P 50±12 vs. 38±11; respectively) could furthermore indicate the presence of N-fixation in the former. Comparison with two additional Patagonian bogs showed similar distinction in C:N and C:S ratios (both: Sphagnum > cushion plant), but variation between cores within sites was more pronounced than between different peatlands. Taking the variable peat accumulation rate (0.09 ? 0.52 mm yr-1) into account, there was notable variation in apparent long term nutrient uptake rates along the transect. N and S uptake rates were larger in cushion plant and graminoid versus Sphagnum cores, while Mg uptake rates were largest in Sphagnum cores. Overall, the stoichiometry of these Patagonian peatlands suggests lower availability of N, P, and Ca compared to peatlands in Ontario, Canada, resulting in lower apparent N, P, and Ca uptake rates. In contrast, apparent Mg uptake rates were larger in Patagonia than Ontario. These results indicate that small scale variability in long term accumulation of nutrients in these ecosystems might be more pronounced than variability in long term C accumulation, and highlights the variability in nutrient availability between peatlands of different regions.Fil: Mathijssen, Paul. University of Münster; AlemaniaFil: Münchberger, Wiebke. University of Münster; AlemaniaFil: Borken, Werner. University of Bayreuth; AlemaniaFil: Pancotto, Veronica Andrea. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Austral de Investigaciones Científicas; ArgentinaFil: Kleinebecker, Till. Justus Liebig Universitat Giessen; AlemaniaFil: Knorr, Klaus Holger. University of Bayreuth; Alemania21th European Geosciences Union General AssemblyVienaAustriaAsamblea General de European Geosciences Unio

Decomposition disentangled: A test of the multiple mechanisms by which nitrogen enrichment alters litter decomposition

1. Nitrogen (N) enrichment has direct effects on ecosystem functioning by altering soil abiotic conditions and indirect effects by reducing plant diversity and shifting plant functional composition from dominance by slow to fast growing species. Litter decomposition is a key ecosystem function and is affected by N enrichment either by a change in litter quality (the recalcitrance of the plant material) or through a change in soil quality (the abiotic and biotic components of the soil that affect decomposition). How the direct and indirect effects of N alter soil and litter quality remains poorly known. 2. We designed a large grassland field experiment manipulating N enrichment, plant species richness and functional composition in a full factorial design. We used three complementary litterbag experiments, combined in a structural equation model (SEM), to quantify the effects of the treatments and various measures of functional composition and diversity on litter and soil quality and overall decomposition. 3. Our results revealed multiple drivers of litter quality and showed that nutrient concentrations (N and calcium) were about twice as important as structural components (leaf dry matter content, fibres) in determining litter quality. Overall the experimental results suggest that N enrichment increases litter decomposition mostly indirectly through a shift in functional composition toward faster growing plant species, producing higher quality litter. N enrichment also altered soil quality and thereby litter decomposition, through its effects on vegetation cover. 4. Our approach provides a mechanistic tool to test the drivers of litter decomposition across different ecosystems. Our results show that litter quality is determined by several nutrient and structure traits and highlight the importance of considering shifts in plant species composition when assessing the effects of N enrichment on decomposition.This study was supported by funding of the Swiss National Science Foundation. S.S. was supported by the Spanish Government under a Ramón y Cajal contract (RYC-2016-20604)

High-resolution classification of south patagonian peat bog microforms reveals potential gaps in up-scaled CH4 fluxes by use of Unmanned Aerial System (UAS) and CIR imagery

South Patagonian peat bogs are little studied sources of methane (CH4). Since CH4 fluxes can vary greatly on a small scale of meters, high-quality maps are needed to accurately quantify CH4 fluxes from bogs. We used high-resolution color infrared (CIR) images captured by an Unmanned Aerial System (UAS) to investigate potential uncertainties in total ecosystem CH4 fluxes introduced by the classification of the surface area. An object-based approach was used to classify vegetation both on species and microform level. We achieved an overall Kappa Index of Agreement (KIA) of 0.90 for the species- and 0.83 for the microform-level classification, respectively. CH4 fluxes were determined by closed chamber measurements on four predominant microforms of the studied bog. Both classification approaches were employed to up-scale CH4 closed chamber measurements in a total area of around 1.8 hectares. Including proportions of the surface area where no chamber measurements were conducted, we estimated a potential uncertainty in ecosystem CH4 fluxes introduced by the classification of the surface area. This potential uncertainty ranged from 14.2 mg·m-2· day-1 to 26.8 mg·m-2· day-1. Our results show that a simple classification with only few classes potentially leads to pronounced bias in total ecosystem CH4 fluxes when plot-scale fluxes are up-scaled.Fil: Lehmann, Jan R. K.. Westfalische Wilhelms Universitat; AlemaniaFil: Münchberger, Wiebke. Westfalische Wilhelms Universitat; AlemaniaFil: Knoth, Christian. Westfalische Wilhelms Universitat; AlemaniaFil: Blodau, Christian. Westfalische Wilhelms Universitat; AlemaniaFil: Nieberding, Felix. Westfalische Wilhelms Universitat; AlemaniaFil: Prinz, Torsten. Westfalische Wilhelms Universitat; AlemaniaFil: Pancotto, Veronica Andrea. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Austral de Investigaciones Científicas; ArgentinaFil: Kleinebecker, Till. Westfalische Wilhelms Universitat; Alemani

Long-term success of floodplain meadow restoration on species-poor grassland

Restoration of floodplain meadows remains a challenge, as many degraded sites suffer from seed limitation. The transfer of seed-containing plant material from species-rich donor sites is a widely used method to restore semi-natural grasslands. However, most studies on the success of such restoration projects comprise limited time frames. As factors determining restoration success may only become evident after many years, long-term observations are crucial. We re-investigated 20 restored grassland sites in the floodplain of the Northern Upper Rhine 13–16 years after plant material transfer with different soil preparation treatments. To this end, we carried out vegetation surveys on 254 permanent plots and studied the potential influence of soil preparation, soil nutrients, and hydrology on plant species composition, diversity, and transfer of target species. Since sustainable agricultural use is important to ensure the long-term stability of restored semi-natural grasslands, we further investigated biomass productivity and feeding value. While most target species increased in frequency or remained stable over time, we found no positive long-term effect of soil preparation on vegetation development and target species establishment. Instead, increased biomass yield and flooding frequency led to reduced restoration success, while higher soil C/N ratios had a positive effect. Overall, restoration measures did not affect the agricultural value of the restored grasslands, which had higher dry matter biomass yields compared with the donor sites. Our results indicate that the positive effect of soil preparation on the number and cover of target species, which is regularly reported in short-term studies, diminishes over time, and other factors such as site conditions become increasingly important. Furthermore, additional plant material transfer or manual seeding may be necessary to support target species establishment. Concerning agricultural usability, the integration of restored floodplain meadows in farming systems is possible and can ensure long-term management and thus stability of these ecosystems. Our study shows that long-term monitoring of restoration projects is necessary, as factors determining restoration success may only become evident in the long-term

Organic vs. Conventional Grassland Management: Do 15N and 13C 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 δ15N and δ13C isotopic composition of soil and hay samples of 21 organic and 34 conventional grasslands in two German regions. We also used Δδ15N (δ15N plant - δ15N 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 δ13C in hay and δ15N in both soil and hay between management types, but showed that δ13C abundances were significantly lower in soil of organic compared to conventional grasslands. Δδ15N values implied that management types did not substantially differ in nitrogen cycling. Only δ13C 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

Temporal and small-scale spatial variation in grassland productivity, biomass quality, and nutrient limitation

Characterization of spatial and temporal variation in grassland productivity and nutrition is crucial for a comprehensive understanding of ecosystem function. Although within-site heterogeneity in soil and plant properties has been shown to be relevant for plant community stability, spatiotemporal variability in these factors is still understudied in temperate grasslands. Our study aimed to detect if soil characteristics and plant diversity could explain observed small-scale spatial and temporal variability in grassland productivity, biomass nutrient concentrations, and nutrient limitation. Therefore, we sampled 360 plots of 20 cm × 20 cm each at six consecutive dates in an unfertilized grassland in Southern Germany. Nutrient limitation was estimated using nutrient ratios in plant biomass. Absolute values of, and spatial variability in, productivity, biomass nutrient concentrations, and nutrient limitation were strongly associated with sampling date. In April, spatial heterogeneity was high and most plots showed phosphorous deficiency, while later in the season nitrogen was the major limiting nutrient. Additionally, a small significant positive association between plant diversity and biomass phosphorus concentrations was observed, but should be tested in more detail. We discuss how low biological activity e.g., of soil microbial organisms might have influenced observed heterogeneity of plant nutrition in early spring in combination with reduced active acquisition of soil resources by plants. These early-season conditions are particularly relevant for future studies as they differ substantially from more thoroughly studied later season conditions. Our study underlines the importance of considering small spatial scales and temporal variability to better elucidate mechanisms of ecosystem functioning and plant community assembly

Direct and plant community mediated effects of management intensity on annual nutrient leaching risk in temperate grasslands

Grassland management intensity influences nutrient cycling both directly, by changing nutrient inputs and outputs from the ecosystem, and indirectly, by altering the nutrient content, and the diversity and functional composition of plant and microbial communities. However, the relative importance of these direct and indirect processes for the leaching of multiple nutrients is poorly studied. We measured the annual leaching of nitrate, ammonium, phosphate and sulphate at a depth of 10 cm in 150 temperate managed grasslands using a resin method. Using Structural Equation Modeling, we distinguished between various direct and indirect effects of management intensity (i.e. grazing and fertilization) on nutrient leaching. We found that management intensity was positively associated with nitrate, ammonium and phosphate leaching risk both directly (i.e. via increased nutrient inputs) and indirectly, by changing the stoichiometry of soils, plants and microbes. In contrast, sulphate leaching risk was negatively associated with management intensity, presumably due to increased outputs with mowing and grazing. In addition, management intensification shifted plant communities towards an exploitative functional composition (characterized by high tissue turnover rates) and, thus, further promoted the leaching risk of inorganic nitrogen. Plant species richness was associated with lower inorganic nitrogen leaching risk, but most of its effects were mediated by stoichiometry and plant community functional traits. Maintaining and restoring diverse plant communities may therefore mitigate the increased leaching risk that management intensity imposes upon grasslands

Nutrient stoichiometry and land use rather than species richness determine plant functional diversity

Ajuts: Deutsche Forschungsgemeinschaft. Grant Numbers: FI 1246/6-1, HO 3830/2-1, KL 2265/5-1 - TRY initiative on plant traits DIVERSITAS/Future Earth and the German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig - Open Access Publication Fund of University of Muenster.Plant functional traits reflect individual and community ecological strategies. They allow the detection of directional changes in community dynamics and ecosystemic processes, being an additional tool to assess biodiversity than species richness. Analysis of functional patterns in plant communities provides mechanistic insight into biodiversity alterations due to anthropogenic activity. Although studies have consi-dered of either anthropogenic management or nutrient availability on functional traits in temperate grasslands, studies combining effects of both drivers are scarce. Here, we assessed the impacts of management intensity (fertilization, mowing, grazing), nutrient stoichiometry (C, N, P, K), and vegetation composition on community-weighted means (CWMs) and functional diversity (Rao's Q) from seven plant traits in 150 grasslands in three regions in Germany, using data of 6 years. Land use and nutrient stoichiometry accounted for larger proportions of model variance of CWM and Rao's Q than species richness and productivity. Grazing affected all analyzed trait groups; fertilization and mowing only impacted generative traits. Grazing was clearly associated with nutrient retention strategies, that is, investing in durable structures and production of fewer, less variable seed. Phenological variability was increased. Fertilization and mowing decreased seed number/mass variability, indicating competition-related effects. Impacts of nutrient stoichiometry on trait syndromes varied. Nutrient limitation (large N:P, C:N ratios) promoted species with conservative strategies, that is, investment in durable plant structures rather than fast growth, fewer seed, and delayed flowering onset. In contrast to seed mass, leaf-economics variability was reduced under P shortage. Species diversity was positively associated with the variability of generative traits. Synthesis. Here, land use, nutrient availability, species richness, and plant functional strategies have been shown to interact complexly, driving community composition, and vegetation responses to management intensity. We suggest that deeper understanding of underlying mechanisms shaping community assembly and biodiversity will require analyzing all these parameters

The Evolution of Ecological Diversity in Acidobacteria

Acidobacteria occur in a large variety of ecosystems worldwide and are particularly abundant and highly diverse in soils. In spite of their diversity, only few species have been characterized to date which makes Acidobacteria one of the most poorly understood phyla among the domain Bacteria. We used a culture-independent niche modeling approach to elucidate ecological adaptations and their evolution for 4,154 operational taxonomic units (OTUs) of Acidobacteria across 150 different, comprehensively characterized grassland soils in Germany. Using the relative abundances of their 16S rRNA gene transcripts, the responses of active OTUs along gradients of 41 environmental variables were modeled using hierarchical logistic regression (HOF), which allowed to determine values for optimum activity for each variable (niche optima). By linking 16S rRNA transcripts to the phylogeny of full 16S rRNA gene sequences, we could trace the evolution of the different ecological adaptations during the diversification of Acidobacteria. This approach revealed a pronounced ecological diversification even among acidobacterial sister clades. Although the evolution of habitat adaptation was mainly cladogenic, it was disrupted by recurrent events of convergent evolution that resulted in frequent habitat switching within individual clades. Our findings indicate that the high diversity of soil acidobacterial communities is largely sustained by differential habitat adaptation even at the level of closely related species. A comparison of niche optima of individual OTUs with the phenotypic properties of their cultivated representatives showed that our niche modeling approach (1) correctly predicts those physiological properties that have been determined for cultivated species of Acidobacteria but (2) also provides ample information on ecological adaptations that cannot be inferred from standard taxonomic descriptions of bacterial isolates. These novel information on specific adaptations of not-yet-cultivated Acidobacteria can therefore guide future cultivation trials and likely will increase their cultivation success

Above- and belowground biodiversity jointly tighten the P cycle in agricultural grasslands

Experiments showed that biodiversity increases grassland productivity and nutrient exploitation, potentially reducing fertiliser needs. Enhancing biodiversity could improve P-use efficiency of grasslands, which is beneficial given that rock-derived P fertilisers are expected to become scarce in the future. Here, we show in a biodiversity experiment that more diverse plant communities were able to exploit P resources more completely than less diverse ones. In the agricultural grasslands that we studied, management effects either overruled or modified the driving role of plant diversity observed in the biodiversity experiment. Nevertheless, we show that greater above- (plants) and belowground (mycorrhizal fungi) biodiversity contributed to tightening the P cycle in agricultural grasslands, as reduced management intensity and the associated increased biodiversity fostered the exploitation of P resources. Our results demonstrate that promoting a high above- and belowground biodiversity has ecological (biodiversity protection) and economical (fertiliser savings) benefits. Such win-win situations for farmers and biodiversity are crucial to convince farmers of the benefits of biodiversity and thus counteract global biodiversity loss