C and N mineralization and earthworm populations in a Norway spruce forest at Hasslöv (SW Sweden), 25 years after liming

During the last decades of the 20th century, acid rain affected many areas in Europe and Northern America. Soil acidification was considered a large problem for forest ecosystems, because it was expected that tree growth would be hampered by low pH, nutrient deficiencies and high concentrations of free Al. Furthermore low soil pH can change the soil fauna and soil microbial biomass. High acidity has also been shown to reduce C and N mineralization. Liming was expected to be a good measure against soil acidification. The present study assesses C and N mineralization and the earthworm community in the Hasslöv forest (SW Sweden; Norway spruce), 25 years after liming with a low dose of CaCO3 (1.75 t ha-1) and a low, medium and high dose (1.55, 3.45 and 8.75 t ha-1) of dolomitic lime. Soil pH correlated to the lime dose. Liming with medium and high doses of dolomitic lime increased C mineralization rates in the FH layer and the upper mineral soil. Liming did not increase N mineralization rates, though there was an increase in nitrification in the soil with the highest dose of lime. The C and N pool were lower in the heavily limed soil and to a lesser extent in the medium limed soil, compared to the control soil, mostly due to a reduction in the organic layer. Furthermore earthworm populations were larger with increasing doses of lime. The dominant species, Dendrobaena octaëdra, was present in all treatments whereas three other species were only found in the highest doses. When extrapolated to the field, both C and N mineralization expressed on an area basis were lower in soils with the highest lime treatment than in the control, and N mineralization was also lower in the medium limed soil. This can be explained by the fact that the C and N pools had been markedly reduced and that the increases in mineralization rate per gram soil could not make up for this reduction. In conclusion, addition of lime has long-lasting effects that can be seen as higher pH, higher C and N mineralization rates, lower soil pools of C and N and higher abundances of earthworms still after 25 years from the liming event. We recommend that as a measure against acidity liming should only be used with great caution

Plant diversity enhances production and downward transport of biodegradable dissolved organic matter

1. Plant diversity is an important driver of belowground ecosystem functions, such as root growth, soil organic matter (SOM) storage, and microbial metabolism, mainly by influencing the interactions between plant roots and soil. Dissolved organic matter (DOM), as the most mobile form of SOM, plays a crucial role for a multitude of soil processes that are central for ecosystem functioning. Thus, DOM is likely to be an important mediator of plant diversity effects on soil processes. However, the relationships between plant diversity and DOM have not been studied so far. 2. We investigated the mechanisms underlying plant diversity effects on concentrations of DOM using continuous soil water sampling across 6 years and 62 plant communities in a long‐term grassland biodiversity experiment in Jena, Germany. Furthermore, we investigated plant diversity effects on the molecular properties of DOM in a subset of the samples. 3. Although DOM concentrations were highly variable over the course of the year with highest concentrations in summer and autumn, we found that DOM concentrations consistently increased with plant diversity across seasons. The positive plant diversity effect on DOM concentrations was mainly mediated by increased microbial activity and newly sequestered carbon in topsoil. However, the effect of soil microbial activity on DOM concentrations differed between seasons, indicating DOM consumption in winter and spring, and DOM production in summer and autumn. Furthermore, we found increased contents of small and easily decomposable DOM molecules reaching deeper soil layers with high plant diversity. 4. Synthesis. Our findings suggest that plant diversity enhances the continuous downward transport of DOM in multiple ways. On the one hand, higher plant diversity results in higher DOM concentrations, on the other hand, this DOM is less degraded. The present study indicates, for the first time, that higher plant diversity enhances the downward transport of dissolved molecules that likely stimulate soil development in deeper layers and therefore increase soil fertility

Standing belowground plant biomass from the Jena Experiment (Main Experiment, year 2011)

This data set contains measurements of standing belowground plant biomass. Data presented here is from the Main Experiment plots of a large grassland biodiversity experiment (the Jena Experiment; see further details below). In the Main experiment, 82 grassland plots of 20 x 20 m were established from a pool of 60 species belonging to four functional groups (grasses, legumes, tall and small herbs). In May 2002, varying numbers of plant species from this species pool were sown into the plots to create a gradient of plant species richness (1, 2, 4, 8, 16 and 60 species) and functional richness (1, 2, 3, 4 functional groups). Plots were maintained in general by bi-annual weeding and mowing. Since 2010, plots were weeded three times per year. Plot size was reduced to 5 x 6 m since 2010. In 2011, standing root biomass was sampled in June. Three (two in few cases because of stones) soil cores with a 3.5 cm diameter per plot were taken to 40 cm depth and pooled plot-wise. The cores were immediately stored cool until further handling. The bulk material of the pooled cores was weighed and cut with scissors to 2 mm) and fine roots and only total root biomass is shown in this dataset

Effects of biodiversity strengthen over time as ecosystem functioning declines at low and increases at high biodiversity

Human-caused declines in biodiversity have stimulated intensive research on the consequences of biodiversity loss for ecosystem services and policy initiatives to preserve the functioning of ecosystems. Short-term biodiversity experiments have documented positive effects of plant species richness on many ecosystem functions, and longer-term studies indicate, for some ecosystem functions, that biodiversity effects can become stronger over time. Theoretically, a biodiversity effect can strengthen over time by an increasing performance of high-diversity communities, by a decreasing performance of low-diversity communities, or a combination of both processes. Which of these two mechanisms prevail, and whether the increase in the biodiversity effect over time is a general property of many functions remains currently unclear. These questions are an important knowledge gap as a continuing decline in the performance of low-diversity communities would indicate an ecosystem-service debt resulting from delayed effects of species loss on ecosystem functioning. Conversely, an increased performance of high-diversity communities over time would indicate that the benefits of biodiversity are generally underestimated in short-term studies. Analyzing 50 ecosystem variables over 11 years in the world's largest grassland biodiversity experiment, we show that overall plant diversity effects strengthened over time. Strengthening biodiversity effects were independent of the considered compartment (above- or belowground), organizational level (ecosystem variables associated with the abiotic habitat, primary producers, or higher trophic levels such as herbivores and pollinators), and variable type (measurements of pools or rates). We found evidence that biodiversity effects strengthened because of both a progressive decrease in functioning in species-poor and a progressive increase in functioning in species-rich communities. Our findings provide evidence that negative feedback effects at low biodiversity are as important for biodiversity effects as complementarity among species at high biodiversity. Finally, our results indicate that a current loss of species will result in a future impairment of ecosystem functioning, potentially decades beyond the moment of species extinction

Functional trait dissimilarity drives both species complementarity and competitive disparity

1. Niche complementarity and competitive disparity are driving mechanisms behind plant community assembly and productivity. Consequently, there is great interest in predicting species complementarity and their competitive differences from their functional traits as dissimilar species may compete less and result in more complete use of resources. 2. Here we assessed the role of trait dissimilarities for species complementarity and competitive disparities within an experimental gradient of plant species richness and functional trait dissimilarity. Communities were assembled using three pools of grass and forb species based on a priori knowledge of traits related to (1) above‐ and below‐ground spatial differences in resource acquisition, (2) phenological differences or (3) both. Complementarity and competitive disparities were assessed by partitioning the overyielding in mixed species communities into species complementarity and dominance effects. 3. Community overyielding and the underlying complementarity and competitive dominance varied strongly among the three plant species pools. Overyielding and complementarity were greatest among species that were assembled based on their variation in both spatial and phenological traits. Competitive dominance was greatest when species were assembled based on spatial resource acquisition traits alone. 4. In communities that were assembled based on species variation in only spatial or phenological traits, greater competitive dominance was predicted by greater differences in SLA and flowering initiation respectively, while greater complementarity was predicted by greater dissimilarity in leaf area and flowering senescence respectively. Greater differences in leaf area could also be linked to greater species complementarity in communities assembled based on variation in both phenological and spatial traits, but trait dissimilarity was unrelated to competitive dominance in these communities. 5. Our results indicate that complementarity and competitive disparity among species are both driven by trait dissimilarities. However, the identity of the traits that drives the complementarity and competitive disparity depends on the trait variation among species that comprise the community. Moreover, we demonstrate that communities assembled with the greater variation in both spatial and phenological traits show the greatest complementarity among species

Incorporation of mineral nitrogen into the soil food web as affected by plant community composition

Although nitrogen (N) deposition is increasing globally, N availability still limits many organisms, such as microorganisms and mesofauna. However, little is known to which extent soil organisms rely on mineral‐derived N and whether plant community composition modifies its incorporation into soil food webs. More diverse plant communities more effectively compete with microorganisms for mineral N likely reducing the incorporation of mineral‐derived N into soil food webs. We set up a field experiment in experimental grasslands with different levels of plant species and functional group richness. We labeled soil with 15NH415NO3 and analyzed the incorporation of mineral‐derived 15N into soil microorganisms and mesofauna over 3 months. Mineral‐derived N incorporation decreased over time in all investigated organisms. Plant species richness and presence of legumes reduced the uptake of mineral‐derived N into microorganisms. In parallel, the incorporation of mineral‐derived 15N into mesofauna species declined with time and decreased with increasing plant species richness in the secondary decomposer springtail Ceratophysella sp. Effects of both plant species richness and functional group richness on other mesofauna species varied with time. The presence of grasses increased the 15N incorporation into Ceratophysella sp., but decreased it in the primary decomposer oribatid mite Tectocepheus velatus sarekensis. The results highlight that mineral N is quickly channeled into soil animal food webs via microorganisms irrespective of plant diversity. The amount of mineral‐derived N incorporated into soil animals, and the plant community properties affecting this incorporation, differed markedly between soil animal taxa, reflecting species‐specific use of food resources. Our results highlight that plant diversity and community composition alter the competition for N in soil and change the transfer of N across trophic levels in soil food webs, potentially leading to changes in soil animal population dynamics and community composition. Sustaining high plant diversity may buffer detrimental effects of elevated N deposition on soil biota.ISSN:2045-775

Species-specific traits and aboveground species-specific plant biomass from the Jena Trait Based Experiment species monocultures (year 2012)

This data collection contains species-specific aboveground plant biomass that was collected from the Trait Based Experiment in 2012. (Sown plant species, Weed plant biomass, the biomass of dead plant material, and the biomass of unidentified plant material) per plots collected in 2012 from a grassland trait diversity experiment (the Jena Trait Based Experiment). The data collection also contains the traits of the species measured in their monoculture. The experiment consists of 20 plant species that were assigned to one of three species pools: 1. Species that vary along a gradient of spatial leaf and root trait similarity, 2. Species that vary along a gradient of phenological trait similarity and 3. Species that vary along a gradient of both spatial and phenological similarity (see Ebeling et al. 2014). The experiment consists of 138 grassland plots 3 x 3 m in size that was established within the Jena Experiment, Germany, in 2011. Plots vary in plant species richness (1, 2, 4, or 8 species) and functional diversity (1, 2, 3, 4 functional diversity levels, where 1 indicates species are most similar and 4 being most dissimilar in functional traits). Plots were maintained by manual weeding in March, July and September. Biomass was harvested twice in 2012 (during peak standing biomass in late May and in late August) on all experimental plots. Plots were mown to the same height directly following biomass harvest. Plant biomass was harvested by clipping the vegetation at 3 cm above ground in two 0.2 x 0.5 m quadrats per plot. The harvested biomass was sorted into categories: individual species of the sown plant species, 'Weed' plant species (species not sown in a plot), detached 'Dead' plant material, and remaining plant material that could not be assigned to any category ('Rest'). All biomass was dried to constant weight (70°C, >= 48 h) and weighed. The data from individual quadrats were averaged. The traits measured are: Flowering initiation, Flowering cessation, specific leaf area (SLA), leaf dry matter content (LDMC), leaf area, maximum canopy height, specific root length (SRL), mean rooting depth (MRD), root mass density (RMD) and root length density (RLD). Flowering initiation and cessation were measured respectively as the week in which flowering was first observed and flowering senesce had completed throughout the plot. Leaf area, leaf fresh mass were measured on approximately five fully expanded leaves from different individuals. These leaves were dried at 65°C for over 48 hours and massed to calculate the specific leaf area (SLA, area per dry mass), and the leaf dry matter content (LDMC, dry mass per fresh mass). Maximum canopy height was measured during peak biomass in May by taking the average of five measurements along a transect. Root traits were measured by taking soil cores, 4 cm in diameter and 40 cm deep and sectioned by depth: 0-5, 5-10, 10-20, 20-30 and 30-40 cm. Roots were washed and roots < 2 mm in diameter were stored in 70 % ethanol. Root length was determined by scanning stained roots with neutral red and scanning roots using WinRhizo software. Root traits were only measured in species pool 1 and 2. Roots were then dried at 65°C for over 48 hours and massed to determine the specific root length (SRL, root length per mass), mean rooting depth (MRD, the average depth weighed by root mass per depth), root mass density (RMD, the average root mass per cubic cm volume) and root length density (RLD, root mass per root length)

Collection of data on belowground plant biomass and morphological root parameters in the Jena Experiment

This collection contains measurements of standing below ground biomass, belowground biomass productivity and morphological root parameters measured on the Main Experiment plots of a large grassland biodiversity experiment (the Jena Experiment; see further details below). In the Main Experiment, 82 grassland plots of 20 x 20 m were established from a pool of 60 species belonging to four functional groups (grasses, legumes, tall and small herbs). In May 2002, varying numbers of plant species from this species pool were sown into the plots to create a gradient of plant species richness (1, 2, 4, 8, 16 and 60 species) and functional richness (1, 2, 3, 4 functional groups). Since 2010, plots were weeded three times per year. The following series of datasets are contained in this collection: 1. Standing below ground biomass: Coarse and fine root biomass was measured in 2003, 2004, 2006 and 2008 in 0 - 30 cm depth. In 2011 and 2014, total root biomass was sampled down to 40 cm depth. Some years report the data divided into sublayers. Every year, several soil cores were taken per plot and pooled before the whole bulk material or a subsample was washed for roots. Roots were dried at 60 - 70 °C and weighed. Standing root biomass was calculated as g m-2. 2. Below ground biomass productivity in 0 - 30 cm depth: Coarse and fine root biomass production from June to September 2003, September 2003 to July 2004 and July 2007 to June 2008 was measured by the ingrowth core method. In 2008, the data is reported divided into sublayers. Each time, five soil cores were taken per plot and replaced by root free soil from the field site. The initially root-free ingrowth cores were removed after a while and pooled plot-wise. To extract the newly formed roots, a subsample of the bulk material was washed for roots. Roots were dried at 70 °C and weighed. Root biomass productivity was calculated as g m-2. In addition, C- (only in 2003 and 2004) and N-concentration of the fine roots was determined. 3. Morphological root parameters of newly formed roots in 0 - 30 cm depth: Root length density and mean root diameter of newly formed roots from June to September 2003 and September 2003 to July 2004 were measured by the ingrowth core method. Each time, five soil cores were taken per plot and replaced by root free soil from the field site. The initially root-free ingrowth cores were removed after a while and pooled plot-wise. To extract the newly formed roots, a subsample of the bulk material was washed and scanned. Root length and mean diameter were determined by using WinRhizo (Regent Instruments, Quebec, Canada). 4. Morphological root parameters of standing roots in 0 - 30 cm depth: In 2004, mean diameter of standing roots was measured by sampling three soil cores per plot. To extract the standing roots, a subsample of the bulk material was washed and scanned. Mean diameter was determined by using WinRhizo (Regent Instruments, Quebec, Canada)

Functional trait dissimilarity drives both species complementarity and competitive disparity

Niche complementarity and competitive disparity are driving mechanisms behind plant community assembly and productivity. Consequently, there is great interest in predicting species complementarity and their competitive differences from their functional traits as dissimilar species may compete less and result in more complete use of resources. Here we assessed the role of trait dissimilarities for species complementarity and competitive disparities within an experimental gradient of plant species richness and functional trait dissimilarity. Communities were assembled using three pools of grass and forb species based on a priori knowledge of traits related to (1) above- and below-ground spatial differences in resource acquisition, (2) phenological differences or (3) both. Complementarity and competitive disparities were assessed by partitioning the overyielding in mixed species communities into species complementarity and dominance effects. Community overyielding and the underlying complementarity and competitive dominance varied strongly among the three plant species pools. Overyielding and complementarity were greatest among species that were assembled based on their variation in both spatial and phenological traits. Competitive dominance was greatest when species were assembled based on spatial resource acquisition traits alone. In communities that were assembled based on species variation in only spatial or phenological traits, greater competitive dominance was predicted by greater differences in SLA and flowering initiation respectively, while greater complementarity was predicted by greater dissimilarity in leaf area and flowering senescence respectively. Greater differences in leaf area could also be linked to greater species complementarity in communities assembled based on variation in both phenological and spatial traits, but trait dissimilarity was unrelated to competitive dominance in these communities. Our results indicate that complementarity and competitive disparity among species are both driven by trait dissimilarities. However, the identity of the traits that drives the complementarity and competitive disparity depends on the trait variation among species that comprise the community. Moreover, we demonstrate that communities assembled with the greater variation in both spatial and phenological traits show the greatest complementarity among species. A plain language summary is available for this article