Biodiversity Experiment Research Articles

Plant selection and soil legacy enhance long-term biodiversity effects.

Plant-plant and plant-soil interactions can help maintain plant diversity and ecosystem functions. Changes in these interactions may underlie experimentally observed increases in biodiversity effects over time via the selection of genotypes adapted to low or high plant diversity. Little is known, however, about such community-history effects and particularly the role of plant-soil interactions in this process. Soil-legacy effects may occur if co-evolved interactions with soil communities either positively or negatively modify plant biodiversity effects. We tested how plant selection and soil legacy influence biodiversity effects on productivity, and whether such effects increase the resistance of the communities to invasion by weeds. We used two plant selection treatments: parental plants growing in monoculture or in mixture over 8yr in a grassland biodiversity experiment in the field, which we term monoculture types and mixture types. The two soil-legacy treatments used in this study were neutral soil inoculated with live or sterilized soil inocula collected from the same plots in the biodiversity experiment. For each of the four factorial combinations, seedlings of eight species were grown in monocultures or four-species mixtures in pots in an experimental garden over 15 weeks. Soil legacy (live inoculum) strongly increased biodiversity complementarity effects for communities of mixture types, and to a significantly weaker extent for communities of monoculture types. This may be attributed to negative plant-soil feedbacks suffered by mixture types in monocultures, whereas monoculture types had positive plant-soil feedbacks, in both monocultures and mixtures. Monocultures of mixture types were most strongly invaded by weeds, presumably due to increased pathogen susceptibility, reduced biomass, and altered plant-soil interactions of mixture types. These results show that biodiversity effects in experimental grassland communities can be modified by the evolution of positive vs. negative plant-soil feedbacks of plant monoculture vs. mixture types.

Convergent high diversity in naturally colonized experimental grasslands is not related to increased productivity

Initial plant diversity might control subsequent community assembly processes and plant productivity. To study these effects, we used a biodiversity experiment (Jena Experiment) with subplots of different sown diversity that were never weeded and spontaneously colonized control plots of different size (3.5×3.5m, 20×20m) with and without mowing in an 8-year study. On non-sown bare plots without mowing, colonizer accumulation depended on plot size resulting in a loss of diversity in large, undisturbed (unmown) control plots after initial colonization. On sown plots that were mown, species richness converged to high levels due to the accumulation of internal colonists (species belonging to the experimental pool of sown species), while initially high species richness of external colonists (species not belonging to the experimental pool) and residents (species sown on the plot) declined over time. The convergence of total species richness at higher levels was paralleled by increased taxonomic (Simpson index, QSimp), phylogenetic (QPhylo) and trait (FDQ) diversity, whereby FDQ was greatest on plots with low resident species richness after several years. Rates of change in terms of species colonization and extinction decelerated over time irrespective of resident species richness, mowing or plot size. While the contribution of residents declined and that of colonists increased, community biomass production did not change over time. The biomasses of residents and colonists were greatest at higher levels of species richness, partial QPhylo and FDQ of the respective species group, but community biomass was consistently weakly related to total species richness and diversity indices. Our study shows that in contrast to the period of succession, “mature” plant communities resulting from natural assembly processes, favouring the coexistence of multiple species and thus high biodiversity, do no longer show significant relationships between species richness and variables related to ecosystem functioning such as primary productivity.

Functional composition rather than species richness determines root characteristics of experimental grasslands grown at different light and nutrient availability

Background and aims Results from many biodiversity experiments have established evidence for positive effects of diversity on aboveground plant productivity. However, less is known about the relationships between plant diversity and belowground plant community characteristics and their consistency at altered environmental conditions.

Complementarity among four highly productive grassland species depends on resource availability.

Positive species richness-productivity relationships are common in biodiversity experiments, but how resource availability modifies biodiversity effects in grass-legume mixtures composed of highly productive species is yet to be explicitly tested. We addressed this question by choosing two grasses (Arrhenatherum elatius and Dactylis glomerata) and two legumes (Medicago × varia and Onobrychis viciifolia) which are highly productive in monocultures and dominant in mixtures (the Jena Experiment). We established monocultures, all possible two- and three-species mixtures, and the four-species mixture under three different resource supply conditions (control, fertilization, and shading). Compared to the control, community biomass production decreased under shading (-56%) and increased under fertilization (+12%). Net diversity effects (i.e., mixture minus mean monoculture biomass) were positive in the control and under shading (on average +15 and +72%, respectively) and negative under fertilization (-10%). Positive complementarity effects in the control suggested resource partitioning and facilitation of growth through symbiotic N2 fixation by legumes. Positive complementarity effects under shading indicated that resource partitioning is also possible when growth is carbon-limited. Negative complementarity effects under fertilization suggested that external nutrient supply depressed facilitative grass-legume interactions due to increased competition for light. Selection effects, which quantify the dominance of species with particularly high monoculture biomasses in the mixture, were generally small compared to complementarity effects, and indicated that these species had comparable competitive strengths in the mixture. Our study shows that resource availability has a strong impact on the occurrence of positive diversity effects among tall and highly productive grass and legume species.

Trophic and Non-Trophic Interactions in a Biodiversity Experiment Assessed by Next-Generation Sequencing.

Plant diversity affects species richness and abundance of taxa at higher trophic levels. However, plant diversity effects on omnivores (feeding on multiple trophic levels) and their trophic and non-trophic interactions are not yet studied because appropriate methods were lacking. A promising approach is the DNA-based analysis of gut contents using next generation sequencing (NGS) technologies. Here, we integrate NGS-based analysis into the framework of a biodiversity experiment where plant taxonomic and functional diversity were manipulated to directly assess environmental interactions involving the omnivorous ground beetle Pterostichus melanarius. Beetle regurgitates were used for NGS-based analysis with universal 18S rDNA primers for eukaryotes. We detected a wide range of taxa with the NGS approach in regurgitates, including organisms representing trophic, phoretic, parasitic, and neutral interactions with P. melanarius. Our findings suggest that the frequency of (i) trophic interactions increased with plant diversity and vegetation cover; (ii) intraguild predation increased with vegetation cover, and (iii) neutral interactions with organisms such as fungi and protists increased with vegetation cover. Experimentally manipulated plant diversity likely affects multitrophic interactions involving omnivorous consumers. Our study therefore shows that trophic and non-trophic interactions can be assessed via NGS to address fundamental questions in biodiversity research.

Predicting the consequences of species loss using size-structured biodiversity approaches.

Understanding the consequences of species loss in complex ecological communities is one of the great challenges in current biodiversity research. For a long time, this topic has been addressed by traditional biodiversity experiments. Most of these approaches treat species as trait-free, taxonomic units characterizing communities only by species number without accounting for species traits. However, extinctions do not occur at random as there is a clear correlation between extinction risk and species traits. In this review, we assume that large species will be most threatened by extinction and use novel allometric and size-spectrum concepts that include body mass as a primary species trait at the levels of populations and individuals, respectively, to re-assess three classic debates on the relationships between biodiversity and (i) food-web structural complexity, (ii) community dynamic stability, and (iii) ecosystem functioning. Contrasting current expectations, size-structured approaches suggest that the loss of large species, that typically exploit most resource species, may lead to future food webs that are less interwoven and more structured by chains of interactions and compartments. The disruption of natural body-mass distributions maintaining food-web stability may trigger avalanches of secondary extinctions and strong trophic cascades with expected knock-on effects on the functionality of the ecosystems. Therefore, we argue that it is crucial to take into account body size as a species trait when analysing the consequences of biodiversity loss for natural ecosystems. Applying size-structured approaches provides an integrative ecological concept that enables a better understanding of each species' unique role across communities and the causes and consequences of biodiversity loss.

Further re‐analyses looking for effects of phylogenetic diversity on community biomass and stability

Species richness (SR) and phylogenetic diversity (PD) are highly correlated measures of plant diversity. Each, by itself, is significantly associated with plant community biomass in biodiversity experiments. As presented by Cadotte (2015) and as we present below, reasonable but alternative analyses that attempt to control for this correlation in different ways provide contradictory or inconclusive support for the hypothesis that PD is superior to SR as a predictor of community biomass. In Venail et al. (2015), we re-analysed data from 16 experimental manipulations of grassland SR to look at how SR and PD influence variation in plant community biomass through time. Using four types of analyses, we showed that, after statistically controlling for variation in SR, PD was not related to community biomass or to the temporal stability of biomass. We did, however, find that SR tends to increase the biomass production of plant communities after controlling for PD. In his comment, Cadotte expressed two concerns about our analyses. One is that we used non-random subsets of experiments, rather than the full data set, for some of our analyses (types 2, 3). We were clear in stating these analyses were based on non-random subsets that were specifically chosen to minimize the SR–PD correlation and avoid problems associated with multicollinearity. We acknowledge that our tests are conservative, a cost of which is that they sacrifice statistical power while, at the same time, minimizing the chance of drawing an incorrect conclusion. But we disagree with Cadotte’s suggestion that our use of non-random data subsets led to ‘biased’ conclusions, and demonstrate later in this response that his claim of bias is unsubstantiated. Cadotte’s second concern was that our analyses did not account for differences in biomass across studies. This is an important criticism to consider; we made a mistake by not controlling for variation in biomass. To address this issue, Cadotte used mixed models where study was included as a random effect, and ran analyses that standardized biomass among sites. Collectively, these led Cadotte to conclude ‘All analyses strongly support previous literature claims about the value of PD and I further show that: (i) PD provides a more powerful explanation of variation in biomass production than species richness; (ii) PD explains variation in biomass production after controlling for richness; and (iii) the use of data subsets inadvertently biased the conclusions’. We have two concerns with Cadotte’s re-analysis. First, Cadotte’s approach largely ignores the concerns we raised about multicollinearity. When two or more predictors

Can the Results of Biodiversity-Ecosystem Productivity Studies Be Translated to Bioenergy Production?

Biodiversity experiments show that increases in plant diversity can lead to greater biomass production, and some researchers suggest that high diversity plantings should be used for bioenergy production. However, many methods used in past biodiversity experiments are impractical for bioenergy plantings. For example, biodiversity experiments often use intensive management such as hand weeding to maintain low diversity plantings and exclude unplanted species, but this would not be done for bioenergy plantings. Also, biodiversity experiments generally use high seeding densities that would be too expensive for bioenergy plantings. Here we report the effects of biodiversity on biomass production from two studies of more realistic bioenergy crop plantings in southern Michigan, USA. One study involved comparing production between switchgrass (Panicum virgatum) monocultures and species-rich prairie plantings on private farm fields that were managed similarly to bioenergy plantings. The other study was an experiment where switchgrass was planted in monoculture and in combination with increasingly species-rich native prairie mixtures. Overall, we found that bioenergy plantings with higher species richness did not produce more biomass than switchgrass monocultures. The lack of a positive relationship between planted species richness and production in our studies may be due to several factors. Non-planted species (weeds) were not removed from our studies and these non-planted species may have competed with planted species and also prevented realized species richness from equaling planted species richness. Also, we found that low seeding density of individual species limited the biomass production of these individual species. Production in future bioenergy plantings with high species richness may be increased by using a high density of inexpensive seed from switchgrass and other highly productive species, and future efforts to translate the results of biodiversity experiments to bioenergy plantings should consider the role of seeding density.

Light and Nutrient Dependent Responses in Secondary Metabolites of Plantago lanceolata Offspring Are Due to Phenotypic Plasticity in Experimental Grasslands.

A few studies in the past have shown that plant diversity in terms of species richness and functional composition can modify plant defense chemistry. However, it is not yet clear to what extent genetic differentiation of plant chemotypes or phenotypic plasticity in response to diversity-induced variation in growth conditions or a combination of both is responsible for this pattern. We collected seed families of ribwort plantain (Plantago lanceolata) from six-year old experimental grasslands of varying plant diversity (Jena Experiment). The offspring of these seed families was grown under standardized conditions with two levels of light and nutrients. The iridoid glycosides, catalpol and aucubin, and verbascoside, a caffeoyl phenylethanoid glycoside, were measured in roots and shoots. Although offspring of different seed families differed in the tissue concentrations of defensive metabolites, plant diversity in the mothers' environment did not explain the variation in the measured defensive metabolites of P. lanceolata offspring. However secondary metabolite levels in roots and shoots were strongly affected by light and nutrient availability. Highest concentrations of iridoid glycosides and verbascoside were found under high light conditions, and nutrient availability had positive effects on iridoid glycoside concentrations in plants grown under high light conditions. However, verbascoside concentrations decreased under high levels of nutrients irrespective of light. The data from our greenhouse study show that phenotypic plasticity in response to environmental variation rather than genetic differentiation in response to plant community diversity is responsible for variation in secondary metabolite concentrations of P. lanceolata in the six-year old communities of the grassland biodiversity experiment. Due to its large phenotypic plasticity P. lanceolata has the potential for a fast and efficient adjustment to varying environmental conditions in plant communities of different species richness and functional composition.

Mechanisms driving diversity-productivity relationships differ between exotic and native communities and are affected by gastropod herbivory.

Biodiversity experiments have shown that productivity usually increases with plant species richness. However, most of those studies disregarded the importance of trophic interactions to the diversity–productivity relationship, and focused on the loss of native species while ignoring invasions by exotic species. Yet, as functional complementarity and the impact of plant antagonists are likely to differ between native and exotic communities, the diversity–productivity relationship may change when native communities are invaded by exotic species. We conducted a mesocosm experiment to test how diversity effects, evenness, and productivity differed between exotic and native assemblages of grassland plants, and how these communities were influenced by slug herbivory. In line with other experiments, we found higher productivity in exotic than in native communities. However, different mechanisms (complementarity vs. selection effect) contributed to the positive diversity–productivity relationships in exotic vs. native communities. Against expectations, native communities showed much lower evenness and a greater selection effect, suggesting that competitive dominance among native species may be even stronger than among exotic species. Slug herbivory decreased productivity independently of species origin and species diversity. However, exotic communities showed a threefold higher complementarity effect than native communities in the absence of slugs, which was mainly driven by differences in the responses of native and exotic legumes and nonleguminous herbs. Our results imply that underlying mechanisms for the positive diversity–productivity relationship differ between native and exotic communities in the early stages of community development, and that differential responses of plant functional groups to generalist herbivory can contribute to this pattern.Electronic supplementary materialThe online version of this article (doi:10.1007/s00442-015-3395-2) contains supplementary material, which is available to authorized users.

Plant species richness does not attenuate responses of soil microbial and nematode communities to a flood event

Human activities are causing climatic changes and alter the composition and biodiversity of ecosystems. Climate change has been and will be increasing the frequency and severity of extreme climate events and natural disasters like floods in many ecosystems. Therefore, it is important to investigate the effects of disturbances on ecosystems and identify potential stabilizing features of ecological communities. In this study, soil microbial and nematode communities were investigated in a grassland biodiversity experiment after a natural flood to investigate if plant diversity is able to attenuate or reinforce the magnitude of effects of the disturbance on soil food webs. In addition to community analyses of soil microorganisms and nematodes, the stability indices proportional resilience, proportional recovery, and proportional resistance were calculated. Generally, soil microbial biomass decreased significantly due to the flood with the strongest reduction in gram-negative bacteria, while gram-positive bacteria were less affected by flooding. Fungal biomass increased significantly three months after the flood compared to few days before the flood, reflecting elevated availability of dead plant biomass in response to the flood. Similar to the soil microbial community, nematode community structure changed considerably due to the flood by favoring colonizers (in the broadest sense r-strategists; c–p 1, 2 nematodes), particularly so at high plant diversity. None of the soil microbial community stability indices and few of the nematode stability indices were significantly affected by plant diversity, indicating limited potential of plant diversity to buffer soil food webs against flooding disturbance. However, plant diversity destabilized colonizer populations, while persister populations (in the broadest sense K-strategists; c–p 4 nematodes) were stabilized, suggesting that plant diversity can stabilize and destabilize populations depending on the ecology of the focal taxa. The present study shows that changes in plant diversity and subsequent alterations in resource availability may significantly modify the compositional shifts of soil food webs in response to disturbances.

Throughfall kinetic energy in young subtropical forests: Investigation on tree species richness effects and spatial variability

Soil erosion threatens ecosystem functioning by reducing soil organic carbon stocks or relocating nutrients. A common measure to protect soil against erosion is afforestation. There is growing evidence that mixed-species forest stands have beneficial effects on ecosystem functions (growth rates, nutrient cycling). In addition, species-rich forests tend to have higher and denser crown cover and thus might affect soil erosion. This study investigated the role of tree species richness on throughfall kinetic energy (TKE) as an important part of the soil erosion process and examined the spatial variability of TKE in mixed-species forest stands.The research was conducted within BEF-China, a large-scale forest biodiversity experiment in subtropical China. In summer 2013, 1800 TKE measurements were carried out during five rainfall events. TKE was measured using splash cups and related to tree height, crown base height, number of branches, leaf area index, stem ground diameter and crown area.Our experiment showed that TKE was not influenced by tree species richness at the plot level. This is likely due to the young age of the experimental forest where a dense and high tree canopy has not yet been developed. However, TKE was influenced by neighborhood tree species richness indicating that tree species richness only affected TKE on a small spatial scale within the direct neighborhood in young forests.TKE showed distinct spatial variability. Directly below the first branch of the tree individuals TKE was lowest (430J/m2) while it was highest in the middle of four tree individuals (556J/m2). Mean freefall kinetic energy (FKE) was 480J/m2. Lower TKE below the first branch than FKE can be attributed to low rain drop velocities due to short falling heights. Higher TKE in the middle of four tree individuals than FKE can be ascribed to a larger crown area on which drops can confluence resulting in an increase of drop mass. Furthermore, TKE was positively affected by the number of influencing tree individuals with a 13% increase of TKE from one to four influencing tree individuals. However, further investigation on TKE below mature trees and dense canopies is needed to confirm our findings for later successional stages.

Are Mixed Tropical Tree Plantations More Resistant to Drought than Monocultures?

Tropical tree plantations usually consist of a single exotic fast growing species, but recent research describes positive effects on ecosystem functions from mixed tropical tree plantations. In this review, we present the current knowledge of drought resistance of tropical mixed species plantations and summarize preliminary evidence from a tree biodiversity experiment in Panama. Converting mono-specific stands into mixed ones may improve stand stability and might reduce increasing abiotic and biotic disturbances due to climate change. However, little is known about the extent to which tropical tree species or tropical tree communities can resist increasing disturbances in the short term, e.g., water limitations due to increasing dry season intensity or length, or about their resilience after such disturbances and their capacity to adapt to changing conditions in the long term. Studies relating drought resistance and resilience to community diversity are missing. Further, we highlight the urgent need for a multifactorial manipulative throughfall reduction experiment in tropical environments. The outcome of such studies would greatly assist the forestry sector in tropical regions to maintain highly productive and ecologically sound forest plantations in a changing climate.

Diversity effects on root length production and loss in an experimental grassland community

Summary Advances in root ecology have revealed that root standing biomass is higher in species‐rich plant communities than in species‐poor communities. Currently, we do not know whether this below‐ground diversity effect is the result of enhanced root production or reduced root mortality or both, which is essential information to understand ecosystem functioning, as it determines C sequestration and N dynamics in soil. Minirhizotron observations were combined with root coring in five different plant communities (four monocultures and the respective mixture). Molecular markers were used to quantitatively determine species abundance in mixed root biomass samples in order to track shifts in below‐ground species composition. In addition, a litterbag experiment was performed to study root decomposition independent of root mortality. Root length production was greater and root length loss was lower in the mixture than expected from monocultures in all years. Simulations suggest that at least two species must have had reduced losses in mixture compared to monoculture. However, the diversity effects on root mortality may partially be explained by selection effects as the species with the longest root life span became dominant in the mixtures. Root length loss from minirhizotrons was very low; the combination of minirhizotron length measurements with root biomass estimates from coring suggested underestimation of root loss in minirhizotrons over time. Root decomposition was not affected by diversity. Diversity enhanced root length production and decreased root loss, resulting in below‐ground overyielding. With decomposition unaffected, our results suggest that root mortality is reduced with increasing diversity. Future studies have to reveal the generality of our observations in larger scale biodiversity experiments by using species having a wider variety of root traits.

Effects of Plant Diversity, Functional Group Composition, and Fertilization on Soil Microbial Properties in Experimental Grassland

BackgroundLoss of biodiversity and increased nutrient inputs are two of the most crucial anthropogenic factors driving ecosystem change. Although both received considerable attention in previous studies, information on their interactive effects on ecosystem functioning is scarce. In particular, little is known on how soil biota and their functions are affected by combined changes in plant diversity and fertilization.Methodology/Principal FindingsWe investigated the effects of plant diversity, functional community composition, and fertilization on the biomass and respiration of soil microbial communities in a long-term biodiversity experiment in semi-natural grassland (Jena Experiment). Plant species richness enhanced microbial basal respiration and microbial biomass, but did not significantly affect microbial specific respiration. In contrast, the presence of legumes and fertilization significantly decreased microbial specific respiration, without altering microbial biomass. The effect of legumes was superimposed by fertilization as indicated by a significant interaction between the presence of legumes and fertilization. Further, changes in microbial stoichiometry (C-to-N ratio) and specific respiration suggest the presence of legumes to reduce N limitation of soil microorganisms and to modify microbial C use efficiency.Conclusions/SignificanceOur study highlights the role of plant species and functional group diversity as well as interactions between plant community composition and fertilizer application for soil microbial functions. Our results suggest soil microbial stoichiometry to be a powerful indicator of microbial functioning under N limited conditions. Although our results support the notion that plant diversity and fertilizer application independently affect microbial functioning, legume effects on microbial N limitation were superimposed by fertilization, indicating significant interactions between the functional composition of plant communities and nutrient inputs for soil processes.

Contrasting effects of tree diversity on young tree growth and resistance to insect herbivores across three biodiversity experiments

Tree diversity is an important driver of forest ecosystem functioning, hypothesised to enhance tree growth and resistance to herbivores. To test this, we assessed the relative importance of tree species richness and functional diversity on tree height growth and insect herbivore damage across three tree diversity experiments in Finland, France and Germany, established within the last fifteen years. These experiments encompass species richness gradients from monocultures up to five species mixtures, with compositions drawn from a pool of eleven tree species. Tree height growth and total insect herbivory were evaluated at both the tree species and forest plot scales.Trees in mixtures tended to grow taller, but on average received more insect herbivory relative to monocultures. Gradients of tree species richness or functional diversity had only weak impact on the magnitude of these effects. Community weighted means of specific leaf area alone captured diversity effects on tree height growth, with stronger positive effects of diversity in mixtures with high community SLA. Tree species‐specific responses were highly variable. No species significantly benefited both in terms of increased growth and reduced herbivory when grown in mixtures. More species showed positive height growth responses in mixed assemblages, but only the two exotic conifers experienced associational resistance to herbivores.This large‐scale study shows that tree height growth in young forest plantations tends to be higher in species mixtures than in monocultures, but incremental increases in functional diversity have, at best, weak marginal growth benefits. Moreover, there appear to be contrasting effects at forest plot versus individual species scales. Thus, while some species show lower herbivore damage in mixtures, this is not a consistent trend and contrasts the higher overall damage in mixtures observed at the forest plot scale. To improve both tree growth and resistance to herbivores in tree species mixtures seems therefore challenging.

Plant diversity increases soil microbial activity and soil carbon storage

Plant diversity strongly influences ecosystem functions and services, such as soil carbon storage. However, the mechanisms underlying the positive plant diversity effects on soil carbon storage are poorly understood. We explored this relationship using long-term data from a grassland biodiversity experiment (The Jena Experiment) and radiocarbon ((14)C) modelling. Here we show that higher plant diversity increases rhizosphere carbon inputs into the microbial community resulting in both increased microbial activity and carbon storage. Increases in soil carbon were related to the enhanced accumulation of recently fixed carbon in high-diversity plots, while plant diversity had less pronounced effects on the decomposition rate of existing carbon. The present study shows that elevated carbon storage at high plant diversity is a direct function of the soil microbial community, indicating that the increase in carbon storage is mainly limited by the integration of new carbon into soil and less by the decomposition of existing soil carbon.

Early positive effects of tree species richness on herbivory in a large-scale forest biodiversity experiment influence tree growth.

Despite the importance of herbivory for the structure and functioning of species-rich forests, little is known about how herbivory is affected by tree species richness, and more specifically by random vs. non-random species loss. We assessed herbivore damage and its effects on tree growth in the early stage of a large-scale forest biodiversity experiment in subtropical China that features random and non-random extinction scenarios of tree mixtures numbering between one and 24 species. In contrast to random species loss, the non-random extinction scenarios were based on the tree species’ local rarity and specific leaf area – traits that may strongly influence the way herbivory is affected by plant species richness. Herbivory increased with tree species richness across all scenarios and was unaffected by the different species compositions in the random and non-random extinction scenarios. Whereas tree growth rates were positively related to herbivory on plots with smaller trees, growth rates significantly declined with increasing herbivory on plots with larger trees. Our results suggest that the effects of herbivory on growth rates increase from monocultures to the most species-rich plant communities and that negative effects with increasing tree species richness become more pronounced with time as trees grow larger. Synthesis. Our results indicate that key trophic interactions can be quick to become established in forest plantations (i.e. already 2.5 years after tree planting). Stronger herbivory effects on tree growth with increasing tree species richness suggest a potentially important role of herbivory in regulating ecosystem functions and the structural development of species-rich forests from the very start of secondary forest succession. The lack of significant differences between the extinction scenarios, however, contrasts with findings from natural forests of higher successional age, where rarity had negative effects on herbivory. This indicates that the effects of non-random species loss could change with forest succession.

Analysis of biodiversity experiments: A comparison of traditional and linear-model-based methods

The relationship between diversity and ecosystem functioning is often studied by biodiversity experiments. Traditionally, the mechanisms behind biodiversity effects observed in these experiments have been evaluated by relative yield, overyielding and Loreau and Hector's additive partitioning of net biodiversity effect, and more recently by linear-model-based methods such as Kirwan's diversity—interaction model and Bell's random partition model. We compared data required and results given by these traditional and linear-model-based methods using three data sets: a simulated data set and two pot biodiversity experiments. For the simulated data, we also compared expected outputs based on defined ecological species traits with actual outcomes of the methods. BEF experiments were designed to answer five fundamental questions: Q1) How does sown species richness change ecosystem functioning? Q2) What ecological mechanisms cause this change? Q3) How does the importance of ecological mechanisms change with sown species richness? Q4) Which species are responsible for given ecological mechanisms? Q5) How do other experimental treatments change answers to all questions above?We show that all methods were capable of answering Q1 and to some extent also Q5 although different methods use different procedures to reach the answer. Concerning Q2–Q4, we found that traditional methods provide more detailed insight into the ecological mechanisms than the linear-model-based methods which leave us just with brief information. A direct comparison between traditional biodiversity effects and effects from linear-model-based methods showed that species interactions from the diversity—interaction model were significantly positively correlated with the net effect, while species identity effects from diversity—interaction model were related to the species relative yield. The selection of an appropriate method for BEF experiment analysis thus depends on the questions we ask which in turn also affect the design of the BEF experiment.

Flooding disturbances increase resource availability and productivity but reduce stability in diverse plant communities.

The natural world is increasingly defined by change. Within the next 100 years, rising atmospheric CO₂ concentrations will continue to increase the frequency and magnitude of extreme weather events. Simultaneously, human activities are reducing global biodiversity, with current extinction rates at ~1,000 × what they were before human domination of Earth's ecosystems. The co-occurrence of these trends may be of particular concern, as greater biological diversity could help ecosystems resist change during large perturbations. We use data from a 200-year flood event to show that when a disturbance is associated with an increase in resource availability, the opposite may occur. Flooding was associated with increases in productivity and decreases in stability, particularly in the highest diversity communities. Our results undermine the utility of the biodiversity-stability hypothesis during a large number of disturbances where resource availability increases. We propose a conceptual framework that can be widely applied during natural disturbances.