Nutrient additions affecting matter turnover in forest and pasture ecosystems

Nutrient inputs into ecosystems of the tropical mountain rainforest region are projected to further increase in the next decades. To investigate whether important ecosystem services such as nutrient cycling and matter turnover in native forests and pasture ecosystems show different patterns of response, two nutrient addition experiments have been established: NUMEX in the forest and FERPAST at the pasture. Both ecosystems already responded 1.5 years after the start of nutrient application (N, P, NP, Ca). Interestingly, most nutrients remained in the respective systems. While the pasture grass was co-limited by N and P, most tree species responded to P addition. Soil microbial biomass in the forest litter layer increased after NP fertilization pointing to nutrient co-limitation. In pasture soils, microorganisms were neither limited by N nor P. The results support the hypothesis that multiple and temporally variable nutrient limitations can coexist in tropical ecosystems.

Seasonal controls on grassland microbial biogeography: Are they governed by plants, abiotic properties or both?

Temporal dynamics create unique and often ephemeral conditions that can influence soil microbial biogeography at different spatial scales. This study investigated the relation between decimeter to meter spatial variability of soil microbial community structure, plant diversity, and soil properties at six dates from April through November. We also explored the robustness of these interactions over time. An historically unfertilized, unplowed grassland in southwest Germany was selected to characterize how seasonal variability in the composition of plant communities and substrate quality changed the biogeography of soil microorganisms at the plot scale (10 m x 10 m). Microbial community spatial structure was positively correlated with the local environment, i.e. physical and chemical soil properties, in spring and autumn, while the density and diversity of plants had an additional effect in the summer period. Spatial relationships among plant and microbial communities were detected only in the early summer and autumn periods when aboveground biomass increase was most rapid and its influence on soil microbial communities was greatest due to increased demand by plants for nutrients. Individual properties exhibited varying degrees of spatial structure over the season. Differential responses of Gram positive and Gram negative bacterial communities to seasonal shifts in soil nutrients were detected. We concluded that spatial distribution patterns of soil microorganisms change over a season and that chemical soil properties are more important controlling factors than plant density and diversity. Finer spatial resolution, such as the mm to cm scale, as well as taxonomic resolution of microbial groups, could help determine the importance of plant species density, composition, and growth stage in shaping microbial community composition and spatial patterns. (C) 2014 The Authors. Published by Elsevier Ltd. All rights reserved.

Reprint of "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 cm(2) in each of six plots representing an area of c. 150 m(2). 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. (C) 2013 Elsevier B.V. All rights reserved.

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.

Determinants of Acidobacteria activity inferred from the relative abundances of 16S rRNA transcripts in German grassland and forest soils

16S rRNA genes and transcripts of Acidobacteria were investigated in 57 grassland and forest soils of three different geographic regions. Acidobacteria contributed 9-31% of bacterial 16S rRNA genes whereas the relative abundances of the respective transcripts were 4-16%. The specific cellular 16S rRNA content (determined as molar ratio of rRNA:rRNA genes) ranged between 3 and 80, indicating a low in situ growth rate. Correlations with flagellate numbers, vascular plant diversity and soil respiration suggest that biotic interactions are important determinants of Acidobacteria 16S rRNA transcript abundances in soils. While the phylogenetic composition of Acidobacteria differed significantly between grassland and forest soils, high throughput denaturing gradient gel electrophoresis and terminal restriction fragment length polymorphism fingerprinting detected 16S rRNA transcripts of most phylotypes in situ. Partial least squares regression suggested that chemical soil conditions such as pH, total nitrogen, C:N ratio, ammonia concentrations and total phosphorus affect the composition of this active fraction of Acidobacteria. Transcript abundance for individual Acidobacteria phylotypes was found to correlate with particular physicochemical (pH, temperature, nitrogen or phosphorus) and, most notably, biological parameters (respiration rates, abundances of ciliates or amoebae, vascular plant diversity), providing culture-independent evidence for a distinct niche specialization of different Acidobacteria even from the same subdivision.

Contrasting long-term records of biomass burning in wet and dry savannas of equatorial East Africa

Rainfall controls fire in tropical savanna ecosystems through impacting both the amount and flammability of plant biomass, and consequently, predicted changes in tropical precipitation over the next century are likely to have contrasting effects on the fire regimes of wet and dry savannas. We reconstructed the long-term dynamics of biomass burning in equatorial East Africa, using fossil charcoal particles from two well-dated lake-sediment records in western Uganda and central Kenya. We compared these high-resolution (5 years/sample) time series of biomass burning, spanning the last 3800 and 1200 years, with independent data on past hydroclimatic variability and vegetation dynamics. In western Uganda, a rapid (<100 years) and permanent increase in burning occurred around 2170 years ago, when climatic drying replaced semideciduous forest by wooded grassland. At the century time scale, biomass burning was inversely related to moisture balance for much of the next two millennia until ca. 1750 ad, when burning increased strongly despite regional climate becoming wetter. A sustained decrease in burning since the mid20th century reflects the intensified modern-day landscape conversion into cropland and plantations. In contrast, in semiarid central Kenya, biomass burning peaked at intermediate moisture-balance levels, whereas it was lower both during the wettest and driest multidecadal periods of the last 1200 years. Here, burning steadily increased since the mid20th century, presumably due to more frequent deliberate ignitions for bush clearing and cattle ranching. Both the observed historical trends and regional contrasts in biomass burning are consistent with spatial variability in fire regimes across the African savanna biome today. They demonstrate the strong dependence of East African fire regimes on both climatic moisture balance and vegetation, and the extent to which this dependence is now being overridden by anthropogenic activity.

Ecosystem functions are resistant to extreme changes to rainfall regimes in a mesotrophic grassland

Major changes to rainfall regimes are predicted for the future but the effect of such changes on terrestrial ecosystem function is largely unknown. We created a rainfall manipulation experiment to investigate the effects of extreme changes in rainfall regimes on ecosystem functioning in a grassland system. We applied two rainfall regimes; a prolonged drought treatment (30 % reduction over spring and summer) and drought/downpour treatment (long periods of no rainfall interspersed with downpours), with an ambient control. Both rainfall manipulations included increased winter rainfall. We measured plant community composition, CO2 fluxes and soil nutrient availability. Plant species richness and cover were lower in the drought/downpour treatment, and showed little recovery after the treatment ceased. Ecosystem processes were less affected, possibly due to winter rainfall additions buffering reduced summer rainfall, which saw relatively small soil moisture changes. However, soil extractable P and ecosystem respiration were significantly higher in rainfall change treatments than in the control. This grassland appears fairly resistant, in the short term, to even the more extreme rainfall changes that are predicted for the region, although prolonged study is needed to measure longer-term impacts. Differences in ecosystem responses between the two treatments emphasise the variety of ecosystem responses to changes in both the size and frequency of rainfall events. Given that model predictions are inconsistent there is therefore a need to assess ecosystem function under a range of potential climate change scenarios.

Factors controlling decomposition rates of fine root litter in temperate forests and grasslands

Background and aims Fine root decomposition contributes significantly to element cycling in terrestrial ecosystems. However, studies on root decomposition rates and on the factors that potentially influence them are fewer than those on leaf litter decomposition. To study the effects of region and land use intensity on fine root decomposition, we established a large scale study in three German regions with different climate regimes and soil properties. Methods In 150 forest and 150 grassland sites we deployed litterbags (100 μm mesh size) with standardized litter consisting of fine roots from European beech in forests and from a lowland mesophilous hay meadow in grasslands. In the central study region, we compared decomposition rates of this standardized litter with root litter collected on-site to separate the effect of litter quality from environmental factors. Results Standardized herbaceous roots in grassland soils decomposed on average significantly faster (24 ± 6 % mass loss after 12 months, mean ± SD) than beech roots in forest soils (12 ± 4 %; p < 0.001). Fine root decomposition varied among the three study regions. Land use intensity, in particular N addition, decreased fine root decomposition in grasslands. The initial lignin:N ratio explained 15 % of the variance in grasslands and 11 % in forests. Soil moisture, soil temperature, and C:N ratios of soils together explained 34 % of the variance of the fine root mass loss in grasslands, and 24 % in forests. Conclusions Grasslands, which have higher fine root biomass and root turnover compared to forests, also have higher rates of root decomposition. Our results further show that at the regional scale fine root decomposition is influenced by environmental variables such as soil moisture, soil temperature and soil nutrient content. Additional variation is explained by root litter quality.

Biotic resistance to plant invasion in grassland: Does seed predation increase with resident plant diversity?

Seed predation impacts heavily on plant populations and community composition in grasslands. In particular, generalist seed predators may contribute to biotic resistance, i.e. the ability of resident species in a community to reduce the success of non-indigenous plant invaders. However, little is known of predators' preferences for seeds of indigenous or non-indigenous plant species or how seed predation varies across communities. We hypothesize that seed predation does not differ between indigenous and non-indigenous plant species and that seed predation is positively related to plant species diversity in the resident community. The seed removal of 36 indigenous and non-indigenous grassland species in seven extensively or intensively managed hay meadows across Switzerland covering a species-richness gradient of 18-50 plant species per unit area (c. 2 m(2)) was studied. In mid-summer 2011, c. 24,000 seeds were exposed to predators in Petri dishes filled with sterilized soil, and the proportions of seeds removed were determined after three days' exposure. These proportions varied among species (9.2-62.5%) and hay meadows (17.8-48.6%). Seed removal was not related to seed size. Moreover, it did not differ between indigenous and non-indigenous species, suggesting that mainly generalist seed predators were active. However, seed predation was positively related to plant species richness across a gradient in the range of 18-38 species per unit area, representing common hay meadows in Switzerland. Our results suggest that generalist post-dispersal seed predation contributes to biotic resistance and may act as a filter to plant invasion by reducing the propagule pressure of non-local plant species.

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.

Vascular Plants and Biocrusts Modulate How Abiotic Factors Affect Wetting and Drying Events in Drylands

Understanding how organisms control soil water dynamics is a major research goal in dryland ecology. Although previous studies have mostly focused on the role of vascular plants on the hydrological cycle of drylands, recent studies highlight the importance of biological soil crusts formed by lichens, mosses, and cyanobacteria (biocrusts) as a major player in this cycle. We used data from a 6.5-year study to evaluate how multiple abiotic (rainfall characteristics, temperature, and initial soil moisture) and biotic (vascular plants and biocrusts) factors interact to determine wetting and drying processes in a semi-arid grassland from Central Spain. We found that the shrub Retama sphaerocarpa and biocrusts with medium cover (25–75%) enhanced water gain and slowed drying compared with bare ground areas (BSCl). Well-developed biocrusts (>75% cover) gained more water, but lost it faster than BSCl microsites. The grass Stipa tenacissima reduced water gain due to rainfall interception, but increased soil moisture retention compared to BSCl microsites. Biotic modulation of water dynamics was the result of different mechanisms acting in tandem and often in opposite directions. For instance, biocrusts promoted an exponential behavior during the first stage of the drying curve, but reduced the importance of soil characteristics that accentuate drying rates. Biocrust-dominated microsites gained a similar amount of water than vascular plants, although they lost it faster than vascular plants during dry periods. Our results emphasize the importance of biocrusts for water dynamics in drylands, and illustrate the potential mechanisms behind their effects. They will help to further advance theoretical and modeling efforts on the hydrology of drylands and their response to ongoing climate change.

Plant and arthropod community sensitivity to rainfall manipulation but not nitrogen enrichment in a successional grassland ecosystem

Grasslands provide many ecosystem services including carbon storage, biodiversity preservation and livestock forage production. These ecosystem services will change in the future in response to multiple global environmental changes, including climate change and increased nitrogen inputs. We conducted an experimental study over 3 years in a mesotrophic grassland ecosystem in southern England. We aimed to expose plots to rainfall manipulation that simulated IPCC 4th Assessment projections for 2100 (+15 % winter rainfall and −30 % summer rainfall) or ambient climate, achieving +15 % winter rainfall and −39 % summer rainfall in rainfall-manipulated plots. Nitrogen (40 kg ha−1 year−1) was also added to half of the experimental plots in factorial combination. Plant species composition and above ground biomass were not affected by rainfall in the first 2 years and the plant community did not respond to nitrogen enrichment throughout the experiment. In the third year, above-ground plant biomass declined in rainfall-manipulated plots, driven by a decline in the abundances of grass species characteristic of moist soils. Declining plant biomass was also associated with changes to arthropod communities, with lower abundances of plant-feeding Auchenorrhyncha and carnivorous Araneae indicating multi-trophic responses to rainfall manipulation. Plant and arthropod community composition and plant biomass responses to rainfall manipulation were not modified by nitrogen enrichment, which was not expected, but may have resulted from prior nitrogen saturation and/or phosphorus limitation. Overall, our study demonstrates that climate change may in future influence plant productivity and induce multi-trophic responses in grasslands.