ALBERT

Description: Conceptual models predict counteractive effects of herbivores and nutrient enrichment on plant diversity and reversed effects of grazers under different nutrient regimes. I tested these hypotheses in 11 field experiments with periphyton communities in three different aquatic habitats (a highly eutrophic lake, an meso-eutrophic lake, and an meso-eutrophic part of the Baltic Sea coast) and in different seasons. Grazer access and nutrient supply were manipulated in a factorial design. Species richness and evenness were chosen as response variables. Both manipulated factors had significant and contrasting effects on diversity, with variable effect strength between sites and seasons. From the two aspects of diversity, evenness well reflected the changes in community composition. Fertilization tended to increase the dominance of few species and thus to decrease evenness, whereas grazers counteracted these effects by removing dominant life forms. The response of species richness was not as expected, since grazers decreased richness throughout, whereas nutrients had weaker effects but tended to increase richness. Species richness rather reflected changes in periphyton architecture. Grazers reduced algal richness presumably by co-consumption of rare species in the tightly connected periphyton assemblages, whereas enrichment may increase richness by providing more structure via increased dominance of filamentous species. Although grazer and nutrient effects on richness and evenness were opposing, there was no change in the effect of one factor by manipulation of the other.

Description: Ecological stoichiometry describes the biochemical constraints of trophic interactions emerging from the different nutrient content and nutrient demand of producers and consumers, respectively. Most research on this topic originates from well-mixed pelagic food webs, whereas the idea has received far less attention in spatially structured habitats. Here, we test how light as well as grazing and nutrient regeneration by consumers affects growth and biomass of benthic primary producers. In the first laboratory experiment, we manipulated grazer presence (two different snail species plus ungrazed control), in the second experiment we factorially combined manipulation of grazer presence and light intensity. We monitored snail and periphyton biomass as well as dissolved and particulate nutrients (nitrogen and phosphorus) over time. Grazers significantly reduced algal biomass in both experiments. Grazers affected periphyton nutrient content depending on the prevailing nutrient limitation and their own body stoichiometry. In the nitrogen (N-) limited first experiment, grazers increased N both in the periphyton and in the water column. The effect was stronger for grazers with lower N-content. In the phosphorus (P-) limited second experiment, grazers increased the P-content of the periphyton, but the grazer with lower N-content had additionally positive effects on algal N. Light reduction did not affect periphyton biomass, but increased chlorophyll-, N- and P-content of the periphyton. These experiments revealed that the indirect effects of grazers on periphyton were bound by stoichiometric constraints of nutrient incorporation and excretion.

Description: Recent experiments, mainly in terrestrial environments, have provided evidence of the functional importance of biodiversity to ecosystem processes and properties. Compared to terrestrial systems, aquatic ecosystems are characterised by greater propagule and material exchange, often steeper physical and chemical gradients, more rapid biological processes and, in marine systems, higher metazoan phylogenetic diversity. These characteristics limit the potential to transfer conclusions derived from terrestrial experiments to aquatic ecosystems whilst at the same time provide opportunities for testing the general validity of hypotheses about effects of biodiversity on ecosystem functioning. Here, we focus on a number of unique features of aquatic experimental systems, propose an expansion to the scope of diversity facets to be considered when assessing the functional consequences of changes in biodiversity and outline a hierarchical classification scheme of ecosystem functions and their corresponding response variables. We then briefly highlight some recent controversial and newly emerging issues relating to biodiversity-ecosystem functioning relationships. Based on lessons learnt from previous experimental and theoretical work, we finally present four novel experimental designs to address largely unresolved questions about biodiversity-ecosystem functioning relationships. These include (1) investigating the effects of non-random species loss through the manipulation of the order and magnitude of such loss using dilution experiments; (2) combining factorial manipulation of diversity in interconnected habitat patches to test the additivity of ecosystem functioning between habitats; (3) disentangling the impact of local processes from the effect of ecosystem openness via factorial manipulation of the rate of recruitment and biodiversity within patches and within an available propagule pool; and (4) addressing how non-random species extinction following sequential exposure to different stressors may affect ecosystem functioning. Implementing these kinds of experimental designs in a variety of systems will, we believe, shift the focus of investigations from a species richness-centred approach to a broader consideration of the multifarious aspects of biodiversity that may well be critical to understanding effects of biodiversity changes on overall ecosystem functioning and to identifying some of the potential underlying mechanisms involved.

Description: Landscape connectivity can increase the capacity of communities to maintain their function when environments change by promoting the immigration of species or populations with adapted traits. However, high immigration may also restrict fine tuning of species compositions to local environmental conditions by homogenizing the community. Here we demonstrate that dispersal generates such a tradeoff between maximizing local biomass and the capacity of model periphyton metacommunities to recover after a simulated heat wave. In non-disturbed metacommunities, dispersal decreased the total biomass by preventing differentiation in species composition between the local patches making up the metacommunity. On the contrary, in metacommunities exposed to a realistic summer heat wave, dispersal promoted recovery by increasing the biomass of heat tolerant species in all local patches. Thus, the heat wave reorganized the species composition of the metacommunities and after an initial decrease in total biomass by 38.7%, dispersal fueled a full recovery of biomass in the restructured metacommunities. Although dispersal may decrease equilibrium biomass, our results highlight that connectivity is a key requirement for the response diversity that allows ecological communities to adapt to climate change through species sorting.

Description: Ecosystem functioning is affected by horizontal (within trophic groups) and vertical (across trophic levels) biodiversity. Theory predicts that the effects of vertical biodiversity depend on consumer specialization. In a microcosm experiment, we investigated ciliate consumer diversity and specialization effects on algal prey biovolume, evenness and composition, and on ciliate biovolume production. The experimental data was complemented by a process-based model further analyzing the ecological mechanisms behind the observed diversity effects. Overall, increasing consumer diversity had no significant effect on prey biovolume or evenness. However, consumer specialization affected the prey community. Specialist consumers showed a stronger negative impact on prey biovolume and evenness than generalists. The model confirmed that this pattern was mainly driven by a single specialist with a high per capita grazing rate, consuming the two most productive prey species. When these were suppressed, the prey assemblage became dominated by a less productive species, consequently decreasing prey biovolume and evenness. Consumer diversity increased consumer biovolume, which was stronger for generalists than for specialists and highest in mixed combinations, indicating that consumer functional diversity, i.e. more diverse feeding strategies, increased resource use efficiency. Overall, our results indicate that consumer diversity effects on prey and consumers strongly depend on species-specific growth and grazing rates, which may be at least equally important as consumer specialization in driving consumer diversity effects across trophic levels.

Description: Climatic warming is a primary driver of change in ecosystems worldwide. Here, we synthesize responses of species richness and evenness from 187 experimental warming studies in a quantitative meta-analysis. We asked 1) whether effects of warming on diversity were detectable and consistent across terrestrial, freshwater and marine ecosystems, 2) if effects on diversity correlated with intensity, duration, and experimental unit size of temperature change manipulations, and 3) whether these experimental effects on diversity interacted with ecosystem types. Using multilevel mixed linear models and model averaging, we also tested the relative importance of variables that described uncontrolled environmental variation and attributes of experimental units. Overall, experimental warming reduced richness across ecosystems (mean log-response ratio = –0.091, 95% bootstrapped CI: –0.13, –0.05) representing an 8.9% decline relative to ambient temperature treatments. Richness did not change in response to warming in freshwater systems, but was more strongly negative in terrestrial (–11.8%) and marine (–10.5%) experiments. In contrast, warming impacts on evenness were neutral overall and in aquatic systems, but weakly negative on land (7.6%). Intensity and duration of experimental warming did not explain variation in diversity responses, but negative effects on richness were stronger in smaller experimental units, particularly in marine systems. Model-averaged parameter estimation confirmed these main effects while accounting for variation in latitude, ambient temperature at the sites of manipulations, venue (field versus lab), community trophic type, and whether experiments were open or closed to colonization. These analyses synthesize extensive experimental evidence showing declines in local richness with increased temperature, particularly in terrestrial and marine communities. However, the more variable effects of warming on evenness were better explained by the random effect of site identity, suggesting that effects on species’ relative abundances were contingent on local species composition.

Description: 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).

Description: This data set contains plant species traits: 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). The traits were measured during the summer of 2012 on the plants grown in monoculture within a grassland trait diversity experiment (the Jena Trait Based Experiment). 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 plots were 3 x 3 m in size and established within the Jena Experiment, Germany, in 2011. Plots were maintained by manual weeding in March, July and September. Traits were measured during the summer of 2012. 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).

Description: This collection contains measurements of element concentrations in plants 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). Plots were maintained by bi-annual weeding and mowing. The following series of datasets are contained in this collection: 1. Carbon and nitrogen concentration in plants: C and N concentration in aboveground plant biomass was measured twice a year (once in 2002 and 2009) from 2002 to 2012. Plants were clipped at 3 cm above ground level in three or four rectangles of 20 x 50 cm size per plot. Target species were pooled per plot and harvest, dried at 70 °C for at least 48 h and cut up with an analysis mill (Kinematica, Littau, Schweiz). The cut material was milled in a ball-mill and carbon and nitrogen concentration was determined with an elemental analyzer. In 2010, phosphorous and potassium concentration was measured additionally. For this purpose, a subsample of the dried and cut material was milled and digested with HNO3 at 200 °C and at about 600-700 MPa using the microwave-assisted high pressure digestion unit (Ethos, Mikrowellen-Laborsysteme (MLS), Leutkirch, Germany). Phosphorus concentrations were determined in a Continuous Flow Analyzer, AA3-system (Bran and Lübbe, Hamburg-Norderstedt, Germany). For K measurement, atom absorption spectroscopy (AAS, Zeenit 700P, Analytik Jena, Jena, Germany) was used. 2. Carbon and nitrogen concentration in plants of the drought experiment: C and N concentration in aboveground plant biomass was measured once a year in 2008 and 2009 on the subplots of the drought experiment. Plants were harvested in rectangles of 20 x 50 cm size. Target species were dried at 70 °C for 48 h, grounded to powder and analyzed with an elemental analyzer. 3. Element analysis of phosphorous (P), calcium (Ca), potassium (K), sodium (Na) and magnesium (Mg) in plants: P, Ca, K, Na and Mg concentrations in aboveground plant biomass were measured twice a year (once in 2004) from 2003 to 2007. Plants were clipped at 3 cm above ground level in three or four rectangles of 20 x 50 cm size per plot. Target species were pooled per plot and harvest, dried at 70 °C for at least 48 h, shredded and milled. Each sample was digested with HNO3 at 200 °C and at about 600-700 MPa using the microwave-assisted high pressure digestion unit (Mars 5 Express, CEM, Lintfort, Germany). Phosphorus concentrations were determined in a Continuous Flow Analyzer, AA3-system (Bran and Lübbe, Hamburg-Norderstedt, Germany). For Ca, K, Na and Mg measurement, atom absorption spectroscopy (AAS, AS240FS Fast Sequential AAS, Varian, Palo Alto, USA) was used.

Description: This data set contains aboveground plant biomass (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 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 location of these rectangles was assigned prior to each harvest by random selection of coordinates within the core area of the plots (i.e. the central 10 x 15 m). The positions of the rectangles within plots were identical for all plots. 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.