Janneke Ravenek

Plant species diversity affects infiltration capacity in an experimental grassland through changes in soil properties

Background and aimsSoil hydraulic properties drive water distribution and availability in soil. There exists limited knowledge of how plant species diversity might influence soil hydraulic properties.MethodsWe quantified the change in infiltration capacity affected by soil structural variables (soil bulk density, porosity and organic carbon content) along a gradient of soil texture, plant species richness (1, 2, 4, 8, 16 and 60) and functional group composition (grasses, legumes, small herbs, tall herbs). We conducted two infiltration measurement campaigns (May and October 2012) using a hood infiltrometer.ResultsPlant species richness significantly increased infiltration capacity in the studied grasslands. Both soil porosity (or inversely bulk density) and organic carbon played an important role in mediating the plant species richness effect. Soil texture did not correlate with infiltration capacity. In spring 2012, earthworm biomass increased infiltration capacity, but this effect could not be attributed to changes in soil structural variables.ConclusionsWe experimentally identified important ecological drivers of infiltration capacity, suggesting complex interactions between plant species richness, earthworms, and soil structural variables, while showing little impact of soil texture. Changes in plant species richness may thus have significant effects on soil hydraulic properties with potential consequences for surface run-off and soil erosion.

No Evidence of Complementary Water Use along a Plant Species Richness Gradient in Temperate Experimental Grasslands

Niche complementarity in resource use has been proposed as a key mechanism to explain the positive effects of increasing plant species richness on ecosystem processes, in particular on primary productivity. Since hardly any information is available for niche complementarity in water use, we tested the effects of plant diversity on spatial and temporal complementarity in water uptake in experimental grasslands by using stable water isotopes. We hypothesized that water uptake from deeper soil depths increases in more diverse compared to low diverse plant species mixtures. We labeled soil water in 8 cm (with 18O) and 28 cm depth (with ²H) three times during the 2011 growing season in 40 temperate grassland communities of varying species richness (2, 4, 8 and 16 species) and functional group number and composition (legumes, grasses, tall herbs, small herbs). Stable isotope analyses of xylem and soil water allowed identifying the preferential depth of water uptake. Higher enrichment in 18O of xylem water than in ²H suggested that the main water uptake was in the upper soil layer. Furthermore, our results revealed no differences in root water uptake among communities with different species richness, different number of functional groups or with time. Thus, our results do not support the hypothesis of increased complementarity in water use in more diverse than in less diverse communities of temperate grassland species.

Linking root traits and competitive success in grassland species

Background and aimsCompetition is an important force shaping plant communities. Here we test the hypothesis that high overall root length density and selective root placement in nutrient patches, as two alternative strategies, confer competitive advantage in species mixtures.MethodsWe performed a full-factorial pairwise competition experiment with eight grassland species in soil with homogeneously distributed nutrients, or with nutrients concentrated in a single patch. We measured species-specific relative growth rate, root length density, selective root placement, and ion uptake rates of all species in monocultures and in mixtures.ResultsGrasses showed higher specific root length overall and forbs a higher selective root placement in the nutrient patch. However, relative growth rate and root length density were more strongly related to competitive ability (measured as relative yield per plant), with little distinction between grasses and forbs.ConclusionsOur results suggest that short-term competitive success was related to fast growth and high root densities, irrespective of nutrient heterogeneity. Developing a large root mass quickly may overwhelm the importance of other traits in the establishment phase of plants, although these other traits may prove to be important in the long run.

Plants are less negatively affected by flooding when growing in species‐rich plant communities

Flooding is expected to increase in frequency and severity in the future. The ecological consequences of flooding are the combined result of species-specific plant traits and ecological context. However, the majority of past flooding research has focused on individual model species under highly controlled conditions. An early summer flooding event in a grassland biodiversity experiment in Jena, Germany, provided the opportunity to assess flooding responses of 60 grassland species in monocultures and 16-species mixtures. We examined plant biomass, species-specific traits (plant height, specific leaf area (SLA), root aerenchyma, starch content) and soil porosity. We found that, on average, plant species were less negatively affected by the flood when grown in higher-diversity plots in July 2013. By September 2013, grasses were unaffected by the flood regardless of plant diversity, and legumes were severely negatively affected regardless of plant diversity. Plants with greater SLA and more root aerenchyma performed better in September. Soil porosity was higher in higher-diversity plots and had a positive effect on plant performance. As floods become more frequent and severe in the future, growing flood-sensitive plants in higher-diversity communities and in soil with greater soil aeration may attenuate the most negative effects of flooding.

Plant diversity shapes microbe-rhizosphere effects on P mobilisation from organic matter in soil

Plant species richness (PSR) increases nutrient uptake which depletes bioavailable nutrient pools in soil. No such relationship between plant uptake and availability in soil was found for phosphorus (P). We explored PSR effects on P mobilisation [phosphatase activity (PA)] in soil. PA increased with PSR. The positive PSR effect was not solely due to an increase in Corg concentrations because PSR remained significant if related to PA:Corg . An increase in PA per unit Corg increases the probability of the temporal and spatial match between substrate, enzyme and microorganism potentially serving as an adaption to competition. Carbon use efficiency of microorganisms (Cmic :Corg ) increased with increasing PSR while enzyme exudation efficiency (PA:Cmic ) remained constant. These findings suggest the need for efficient C rather than P cycling underlying the relationship between PSR and PA. Our results indicate that the coupling between C and P cycling in soil becomes tighter with increasing PSR.

Functional trait dissimilarity drives both species complementarity and competitive disparity

Summary 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 on 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 belowground 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 was 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 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. This article is protected by copyright. All rights reserved.

Below-ground complementarity effects in a grassland biodiversity experiment are related to deep-rooting species

Summary 1.Belowground resource partitioning is often proposed as the underlying mechanism for the positive relationship between plant species richness and productivity. For example, if species have different root distributions, a mixture of plant species may be able to use the available resources more completely than the individual species in a monoculture. However, there is little experimental evidence for differentiation in vertical root distributions among species and its contribution to biodiversity effects. 2.We determined species-specific root standing biomass over depth using molecular techniques (real time-qPCR) in a large grassland biodiversity experiment (1-8 plant species mixtures), in two years. Species-specific root biomass data were used to disentangle the effects of positive interactions between species (complementarity effects) and effects due to dominance of productive species (selection effects) on root biomass in mixtures. In a next step, these biodiversity effects were linked to the diversity of rooting depths and the averaged rooting depth of the community. 3.Root biomass increased with species richness. This was mainly due to positive interactions (the complementarity effect), which increased with species richness belowground. In contrast, the selection effect decreased with species richness. Although there was considerable variation in vertical root distribution between species in monocultures, the diversity of rooting strategies did not explain the complementarity effect. Rather, the abundance of deep-rooting species in mixtures (i.e. high community weighted mean) was significantly related to the complementarity effect. Comparing the ‘predicted’ root distribution (based on monocultures) to the actual distribution in mixtures, we found that mixtures rooted deeper than expected, but this did not better explain the complementarity effect. 4.Synthesis: This study demonstrates that vertical root distributions of species provide only subtle evidence for resource partitioning. We found no evidence that functional diversity in vertical rooting patterns was important for the complementarity effect, in contrast to our expectation that the enhancement of productivity was due to resource partitioning. Alternatively, we found significant but weak relationships between the complementarity effect and deep-rooting communities, based on the community weighted mean root distribution. This suggests that factors other than belowground resource partitioning alone may drive the biodiversity-productivity relationship. This article is protected by copyright. All rights reserved.

Below-ground resource partitioning alone cannot explain the biodiversity–ecosystem function relationship: a field test using multiple tracers

Below-ground resource partitioning is among the most prominent hypotheses for driving the positive biodiversity-ecosystem function relationship. However, experimental tests of this hypothesis in biodiversity experiments are scarce, and the available evidence is not consistent. We tested the hypothesis that resource partitioning in space, in time or in both space and time combined drives the positive effect of diversity on both plant productivity and total community resource uptake. At the community level, we predicted that total community resource uptake and biomass production above- and below-ground will increase with increased species richness or functional group richness. We predicted that, at the species level, resource partition breadth will become narrower, and that overlap between the resource partitions of different species will become smaller with increasing species richness or functional group richness. We applied multiple resource tracers (Li and Rb as potassium analogues, the water isotopologues-H2 18O and 2H2O, and 15N) in three seasons at two depths across a species and functional group richness gradient at a grassland biodiversity experiment. We used this multidimensional resource tracer study to test if plant species partition resources with increasing plant diversity across space, time or both simultaneously. At the community level, total community resource uptake of nitrogen and potassium and above- and below-ground biomass increased significantly with increasing species richness but not with increasing functional group richness. However, we found no evidence that resource partition breadth or resource partition overlap decreased with increasing species richness for any resource in space, time or both space and time combined. Synthesis. These findings indicate that below-ground resource partitioning may not drive the enhanced resource uptake or biomass production found here. Instead, other mechanisms such as facilitation, species-specific biotic feedback or above-ground resource partitioning are likely necessary for enhanced overall ecosystem function.

Plant species richness and functional groups have different effects on soil water content in a decade‐long grassland experiment

The temporal and spatial dynamics of soil water are closely interlinked with terrestrial ecosystems functioning. The interaction between plant community properties such as species composition and richness and soil water mirrors fundamental ecological processes determining above-ground–below-ground feedbacks. Plant–water relations and water stress have attracted considerable attention in biodiversity experiments. Yet, although soil scientific research suggests an influence of ecosystem productivity on soil hydraulic properties, temporal changes of the soil water content and soil hydraulic properties remain largely understudied in biodiversity experiments. Thus, insights on how plant diversity—productivity relationships affect soil water are lacking. Here, we determine which factors related to plant community composition (species and functional group richness, presence of plant functional groups) and soil (organic carbon concentration) affect soil water in a long-term grassland biodiversity experiment (The Jena Experiment). Both plant species richness and the presence of particular functional groups affected soil water content, while functional group richness played no role. The effect of species richness changed from positive to negative and expanded to deeper soil with time. Shortly after establishment, increased topsoil water content was related to higher leaf area index in species-rich plots, which enhanced shading. In later years, higher species richness increased topsoil organic carbon, likely improving soil aggregation. Improved aggregation, in turn, dried topsoils in species-rich plots due to faster drainage of rainwater. Functional groups affected soil water distribution, likely due to plant traits affecting root water uptake depths, shading, or water-use efficiency. For instance, topsoils in plots containing grasses were generally drier, while plots with legumes were moister. Synthesis. Our decade-long experiment reveals that the maturation of grasslands changes the effects of plant richness from influencing soil water content through shading effects to altering soil physical characteristics in addition to modification of water uptake depth. Functional groups affected the soil water distribution by characteristic shifts of root water uptake depth, but did not enhance exploitation of the overall soil water storage. Our results reconcile previous seemingly contradictory results on the relation between grassland species diversity and soil moisture and highlight the role of vegetation composition for soil processes.