Does species diversity limit productivity in natural grassland communities?

Theoretical analyses and experimental studies of synthesized assemblages indicate that under particular circumstances species diversity can enhance community productivity through niche complementarity. It remains unclear whether this process has important effects in mature natural ecosystems where competitive feedbacks and complex environmental influences affect diversity–productivity relationships. In this study, we evaluated diversity–productivity relationships while statistically controlling for environmental influences in 12 natural grassland ecosystems. Because diversity–productivity relationships are conspicuously nonlinear, we developed a nonlinear structural equation modeling (SEM) methodology to separate the effects of diversity on productivity from the effects of productivity on diversity. Meta-analysis was used to summarize the SEM findings across studies. While competitive effects were readily detected, enhancement of production by diversity was not. These results suggest that the influence of small-scale diversity on productivity in mature natural systems is a weak force, both in absolute terms and relative to the effects of other controls on productivity.     

Defoliation and competition effects in a productivity gradient for a semiarid Mediterranean annual grassland community

Grazing and competition are two main factors shaping range plant communities; however, few studies have investigated their interaction. The current study aimed to investigate the effects of defoliation, competition and their interaction on production of annual grasses in semiarid Mediterranean areas. Competition treatments (absence/presence of neighbors) were combined with three defoliation intensities (0%, 30% and 60%) in a complete factorial design. Competition significantly reduced grass biomass. However, the role of competition was eliminated under heavy defoliation or under dry growth conditions. Defoliation showed variable results on final biomass (FB) and cumulative biomass (CB). While heavy defoliation (60% clipping intensity) reduced grass FB down to 80% during the two growing seasons, light defoliation (30%) significantly increased CB. Results showed that competition may limit the direct effect of defoliation on dominant grass species. Further, the relationship between site productivity and competition effect was best explained by a negative linear model. This hypothesized model may suggest that facilitation and competition alternatively affect grassland communities along a productivity gradient. The results suggest that light grazing may sustain or even enhance grassland productivity. The results also indicated the suitability of annual grass species to re-vegetate degraded rangeland in semi-arid climate. Further, optimum grazing practices to conserve biodiversity of Avena grassland may involve moderate stocking rate.     

Early-successional plants regulate grassland productivity and species composition: a removal experiment

Grime's (1998) "mass-ratio" hypothesis holds that ecosystem processes depend in the short term on functional properties of dominant plants and in the longer term on how resident species influence the recruitment of dominants. The latter of these effects may be especially important among early-successional species in disturbed ecosystems, but experimental tests are few. We removed two groups of early-successional species, an annual forb Gutierrezia dracunculoides (DC.) S. F. Blake and annual species (mostly grasses) that complete growth early in the growing season [early-season (ES) species], from a heavily-grazed grassland in central Texas, USA dominated by a C₄ perennial grass. We sought to determine effects of annuals on grassland functioning [productivity, water balance, soil and plant nitrogen (N)] and composition. Removals did not impact N retention in the soil/plant system during the two years of this study, but removing ES annuals increased the amount of water between 30 and 120 cm in the soil profile early in each growing season. Production and N accumulation by vegetation declined following the removal of ES annuals in approximate proportion to the contribution of annuals to aboveground biomass and N, consistent with the mass-ratio hypothesis. By the second year, production and N uptake by initially sub-dominant species increased to fully compensate for the loss of annuals. These results are consistent with the view that ecosystem functions are more strongly linked to species attributes than to diversity per se. Longer-term effects of annuals on grassland composition were evident in a dramatic increase in biomass of perennial forbs after annuals were removed. Because perennial forbs differ from the dominant grass in this grassland in traits that influence ecosystem functioning, ES annuals may affect grassland functioning more by regulating the composition of vegetation than by directly affecting process rates.     

Experimental Manipulation of Grassland Plant Diversity Induces Complex Shifts in Aboveground Arthropod Diversity

Changes in producer diversity cause multiple changes in consumer communities through various mechanisms. However, past analyses investigating the relationship between plant diversity and arthropod consumers focused only on few aspects of arthropod diversity, e.g. species richness and abundance. Yet, shifts in understudied facets of arthropod diversity like relative abundances or species dominance may have strong effects on arthropod-mediated ecosystem functions. Here we analyze the relationship between plant species richness and arthropod diversity using four complementary diversity indices, namely: abundance, species richness, evenness (equitability of the abundance distribution) and dominance (relative abundance of the dominant species). Along an experimental gradient of plant species richness (1, 2, 4, 8, 16 and 60 plant species), we sampled herbivorous and carnivorous arthropods using pitfall traps and suction sampling during a whole vegetation period. We tested whether plant species richness affects consumer diversity directly (i), or indirectly through increased productivity (ii). Further, we tested the impact of plant community composition on arthropod diversity by testing for the effects of plant functional groups (iii). Abundance and species richness of both herbivores and carnivores increased with increasing plant species richness, but the underlying mechanisms differed between the two trophic groups. While higher species richness in herbivores was caused by an increase in resource diversity, carnivore richness was driven by plant productivity. Evenness of herbivore communities did not change along the gradient in plant species richness, whereas evenness of carnivores declined. The abundance of dominant herbivore species showed no response to changes in plant species richness, but the dominant carnivores were more abundant in species-rich plant communities. The functional composition of plant communities had small impacts on herbivore communities, whereas carnivore communities were affected by forbs of small stature, grasses and legumes. Contrasting patterns in the abundance of dominant species imply different levels of resource specialization for dominant herbivores (narrow food spectrum) and carnivores (broad food spectrum). That in turn could heavily affect ecosystem functions mediated by herbivorous and carnivorous arthropods, such as herbivory or biological pest control.     

Species richness peaks for intermediate levels of biomass in a fractal succession with quasi-neutral interactions

The mechanisms that promote species richness, including net community interactions, are considered central to the investigation of the consequences of biodiversity loss for ecosystem functioning. Recently, some empirical studies at large spatiotemporal scales suggest that increasing species richness within natural communities results in a finer division of biomass among species rather than an increase in total biomass. In parallel, the most common pattern observed in nature is the peaked relationship between diversity and productivity estimated as total biomass. Thus, the aim of our study is to provide model predictions for the diversity–biomass relationship with various levels of net species interactions within communities: negative, neutral, quasi-neutral and positive. Using a scaling relationship between the number of species and total community biomass, we propose a new self-similar process of biomass partitioning during a community assembly process. At each step of the succession, K more species appear that are A times less abundant on average giving K=A^d; the parameter d being a fractal dimension related to the nature of interactions among coexisting species. Our results, compared to those from meta-analyses about empirical diversity–productivity relationships, illustrate that quasi-neutral interactions among coexisting species lead to the most commonly observed pattern: an 'envelope' where diversity peaks at intermediate values of total biomass, i.e. that the area below the hump-backed line (considered as the upper boundary) is filled with data points.     

Community maturity, species saturation and the variant diversity-productivity relationships in grasslands

Detailed knowledge of the relationship between plant diversity and productivity is critical for advancing our understanding of ecosystem functioning and for achieving success in habitat restoration efforts. However, effects and interactions of diversity, succession and biotic invasions on productivity remain elusive. We studied newly established communities in relation to preexisting homogeneous vegetation invaded by exotic plants in the northern Great Plains, USA, at four study sites for 3 years. We observed variant diversity–productivity relationships for the seeded communities (generally positive monotonic at three sites and non-monotonic at the other site) but no relationships for the resident community or the seeded and resident communities combined at all sites and all years. Community richness was enhanced by seeding additional species but productivity was not. The optimal diversity (as indicated by maximum productivity) changed among sites and as the community developed. The findings shed new light on ecosystem functioning of biodiversity under different conditions and have important implications for restoration.     

Genotypic diversity of a dominant C4 grass across a long-term fire frequency gradient

Aims and Methods Diversity-disturbance research has focused on community diversity, but disturbance frequency could impact diversity within species as well, with important consequences for community diversity and ecosystem function. We examined patterns of genetic diversity of a dominant grass species, Andropogon gerardii, in native North American tallgrass prairie sites located in eastern Kansas that have been subjected to a gradient of fire frequency treatments (burned every 1, 2, 4 or 20 years) since the 1970s. In addition, we were able to assess the relationships between genetic diversity of A. gerardii, species diversity and productivity across this range of fire frequencies.Important findings We found no significant relationships between genetic diversity of A. gerardii at the local scale (1 m 2 plot level) and disturbance frequency (burned 2 to 32 times over a 38-year period). However, at the site level (i.e. across all plots sampled within a site, ~100 m 2) there were differences in genotype richness and composition, as well as genomic dissimilarity among individuals of A. gerardii. Genotype richness was greatest for the site burned at an intermediate (4-year) frequency and lowest for the infrequently (20-year) burned site. In addition, genotypes found in the frequently burned sites were more similar from each other than expected by random chance than those found in the infrequently burned sites. Genotype composition of A. gerardii was not significantly different between the frequently burned sites (annual vs. 2 year) but did differ between frequently burned and infrequently burned sites (1 and 2 year vs. 4 and 20 year, etc.). Together, these results suggest site-level ecological sorting of genotypes in intact prairie across a broad gradient of disturbance frequencies, likely driven by alterations in environmental conditions. Frequent fire promotes the abundance of dominant grass species, reduces plant community diversity and impacts ecosystem processes such as productivity. Our study suggests that genetic diversity within dominant grass species also may be affected by disturbance frequency, which could have important implications for how species are able to respond to disturbance.     

Species richness, temporal variability and resistance of biomass production in a Mediterranean grassland

We studied the temporal variability and resistance to perturbation of the biomass production of grassland communities from an experimental diversity gradient (the Portuguese BIODEPTH project site). With increasing species richness relative temporal variability (CV) of plant populations increased but that of communities decreased, supporting the insurance hypothesis and related theory. Species‐rich communities were more productive than species‐poor communities in all three years although a natural climatic perturbation in the third year (frequent frost and low precipitation) caused an overall decrease in biomass production. Resistance to this perturbation was constant across the experimental species richness gradient in relative terms, supporting a similar response from the Swiss BIODEPTH experiment. The positive biomass response was generated by different combinations of the complementarity and selection effects in different years. Complementarity effects were positive across mixtures on average in all three years and positively related to diversity in one season. The complementarity effect declined following perturbation in line with total biomass but, counter to predictions, in relative terms overyielding was maintained in all years. Selection effects were positively related to diversity in one year and negative overall in the other two years. The response to perturbation varied among species and for the same species growing in monoculture and mixture, but following the frost communities were more strongly dominated by species with lower monoculture biomass and the selection effect was more negative. In total, our results support previous findings of a positive relationship between diversity and productivity and between diversity and the temporal stability of production, but of no effect of diversity on the resistance to perturbation. We demonstrate for the first time that the relative strength of overyielding remained constant during an exceptional natural environmental perturbation.     

Floristic diversity of an eastern Mediterranean dwarf shrubland: the importance of soil pH

Questions: Does plant species richness and composition of eastern Mediterranean dwarf shrubland (phrygana) correlate with soil pH? How important is the effect of pH on species diversity in relation to other environmental factors in this ecosystem? What is the evolutionary background of the diversity–pH relationship? Location: Western Crete, Greece. Methods: Species composition of vascular plants, soil and other environmental variables were sampled in 100‐m² plots on acidic and basic bedrock in phrygana vegetation. The relationships between species composition and environmental variables (including climate) were tested using canonical correspondence analysis, and relationships between species richness and environment using correlation and regression analyses. Data were analysed separately for different plant functional types based on life form and life span. Results: Although soil pH varied across a narrow range (5.9‐8.1), species composition changed significantly along the pH gradient within all plant functional types. For most functional types, the effect of soil pH on species composition was stronger than that of other environmental variables. Species richness of annuals, geophytes and suffruticose chamaephytes increased with soil pH, while richness of hemicryptophytes and shrubs was not correlated with pH. Conclusions: The results are consistent with the evolutionary species pool hypothesis. High numbers of calcicole annuals, geophytes and suffruticose chamaephytes may be a result of the evolution of these groups on base‐rich dry soils in the Mediterranean climate. In contrast, hemicryptophytes, a life form typical of the temperate zone, evolved on both acidic and basic soils and therefore their species numbers do not respond to soil pH across the narrow range studied. The lack of a relationship between shrub species richness and pH is difficult to explain: it may reflect the more diverse or older origin of Mediterranean woody species and their conservative niches.     

Species–area relationships in Mediterranean-climate plant communities

Aim To determine the best‐fit model of species–area relationships for Mediterranean‐type plant communities and evaluate how community structure affects these species–area models. Location Data were collected from California shrublands and woodlands and compared with literature reports for other Mediterranean‐climate regions. Methods The number of species was recorded from 1, 100 and 1000 m² nested plots. Best fit to the power model or exponential model was determined by comparing adjusted r² values from the least squares regression, pattern of residuals, homoscedasticity across scales, and semi‐log slopes at 1–100 m² and 100–1000 m². Dominance–diversity curves were tested for fit to the lognormal model, MacArthur's broken stick model, and the geometric and harmonic series. Results Early successional Western Australia and California shrublands represented the extremes and provide an interesting contrast as the exponential model was the best fit for the former, and the power model for the latter, despite similar total species richness. We hypothesize that structural differences in these communities account for the different species–area curves and are tied to patterns of dominance, equitability and life form distribution. Dominance–diversity relationships for Western Australian heathlands exhibited a close fit to MacArthur's broken stick model, indicating more equitable distribution of species. In contrast, Californian shrublands, both postfire and mature stands, were best fit by the geometric model indicating strong dominance and many minor subordinate species. These regions differ in life form distribution, with annuals being a major component of diversity in early successional Californian shrublands although they are largely lacking in mature stands. Both young and old Australian heathlands are dominated by perennials, and annuals are largely absent. Inherent in all of these ecosystems is cyclical disequilibrium caused by periodic fires. The potential for community reassembly is greater in Californian shrublands where only a quarter of the flora resprout, whereas three quarters resprout in Australian heathlands. Other Californian vegetation types sampled include coniferous forests, oak savannas and desert scrub, and demonstrate that different community structures may lead to a similar species–area relationship. Dominance–diversity relationships for coniferous forests closely follow a geometric model whereas associated oak savannas show a close fit to the lognormal model. However, for both communities, species–area curves fit a power model. The primary driver appears to be the presence of annuals. Desert scrub communities illustrate dramatic changes in both species diversity and dominance–diversity relationships in high and low rainfall years, because of the disappearance of annuals in drought years. Main conclusions Species–area curves for immature shrublands in California and the majority of Mediterranean plant communities fit a power function model. Exceptions that fit the exponential model are not because of sampling error or scaling effects, rather structural differences in these communities provide plausible explanations. The exponential species–area model may arise in more than one way. In the highly diverse Australian heathlands it results from a rapid increase in species richness at small scales. In mature California shrublands it results from very depauperate richness at the community scale. In both instances the exponential model is tied to a preponderance of perennials and paucity of annuals. For communities fit by a power model, coefficients z and log c exhibit a number of significant correlations with other diversity parameters, suggesting that they have some predictive value in ecological communities.     

Effects of diversity on diversity: consequences of competition and facilitation

Diversity in one group of species or genotypes is often correlated with diversity in a second group – prominent examples including native vs exotic species, and genetic diversity in a focal species vs species diversity in the rest of the community. I used simulation models to investigate the roles of competition and facilitation among species or genotypes in creating diversity–diversity relationships, with a focus on facilitation, which has received little theoretical attention. When competitive interactions dominate, increasing diversity in one group reduces diversity in the second group via filling of available niche space. Facilitation can create positive diversity–diversity relationships via a sampling effect, whereby a strong facilitator of the second group is more likely to be present as diversity increases in the first group, and also via one group acting as a source of biotic heterogeneity (i.e. diversifying selection) on the second group. However, the biotic heterogeneity effect is expected only under restricted conditions – with asymmetric facilitation, only during a transient period, or only over a small range of species diversity levels – and therefore seems unlikely to operate within trophic levels in natural communities. More generally, the simultaneous operation of competition and facilitation results in several different diversity–diversity relationships and underlying mechanisms. The results clarify the potential roles of positive and negative interactions in creating diversity–diversity relationships, and in determining the outcome of community dynamics in general. This study also highlights some important difficulties in incorporating facilitation into ecological theory for communities with many species.     

How do subordinate and dominant species in semi-natural mountain grasslands relate to productivity and land-use change?

Changes in agricultural practices of semi-natural mountain grasslands are expected to modify plant community structure and shift dominance patterns. Using vegetation surveys of 11 sites in semi-natural grasslands of the Swiss Jura and Swiss and French Alps, we determined the relative contribution of dominant, subordinate and transient plant species in grazed and abandoned communities and observed their changes along a gradient of productivity and in response to abandonment of pasturing.The results confirm the humpbacked diversity–productivity relationship in semi-natural grassland, which is due to the increase of subordinate species number at intermediate productivity levels. Grazed communities, at the lower or higher end of the species diversity gradient, suffered higher species loss after grazing abandonment. Species loss after abandonment of pasturing was mainly due to a higher reduction in the number of subordinate species, as a consequence of the increasing proportion of dominant species.When plant biodiversity maintenance is the aim, our results have direct implications for the way grasslands should be managed. Indeed, while intensification and abandonment have been accelerated since few decades, our findings in this multi-site analysis confirm the importance of maintaining intermediate levels of pasturing to preserve biodiversity.     

Plant responsiveness to variation in precipitation and nitrogen is consistent across the compositional diversity of a California annual grassland

Question: How does responsiveness to water and Nitrogen (N) availability vary across the compositional and functional diversity that exists in a mesic California annual grassland plant community? Location: Northern California annual grassland. Methods: A mesocosm system was used to simulate average annual precipitation totals and dry and wet year extremes observed in northern California mesic grasslands. The effects of precipitation and N availability on biomass and fecundity were measured on three different vegetation types, a mixed grass forb community, and a forb and a grass monoculture. The treatment effects on plant community composition were examined in the mixed species community. Results: While growth and seed production of the three vegetation types was inherently different, their responses to variation in precipitation and N were statistically similar. Plant density, shoot biomass, and seed production tended to increase with greater water availability in all vegetation types, with the exception of a consistent growth reduction in high precipitation (1245 mm) plots in the first year of the study. Shoot biomass responded positively to N addition, an effect that increased with greater water availability. Nitrogen addition had little effect on plant density or seed production. In the mixed grass‐forb community, biomass responsiveness to water and N treatments were consistently driven by the shoot growth of Avena barbata, the dominant grass species. Conclusions: Vegetation responses to changes in precipitation and N availability were consistent across a range of composition and structural diversity in this study. Plant growth and seed production were sensitive to both increased and decreased precipitation totals, and the magnitude of these responses to N availability varied depending on soil moisture conditions. Our results suggest the impacts of changing precipitation regimes and N deposition on annual productivity of California grasslands may be predictable under different climate scenarios across a range of plant communities.     

On the relative importance of the effects of selection and complementarity as drivers of diversity–productivity relationships in Mediterranean shrublands

Extensive research has been devoted to understanding the role of biodiversity as a driver of ecosystem functioning. However, no previous study has evaluated the relative contribution of complementarity and selection to productivity in shrublands. We have attempted to do this for a Mediterranean shrubland dominated by Quercus coccifera , Cistus albidus , Ulex parviflorus and Rosmarinus officinalis . We found a highly significant and linear positive relationship between productivity and species richness. No selection effect was apparent, but both the complementarity and net effects were highly significant. The magnitude of these effects increased from two to three species, but became non-significant in the four-species mixtures. Analysis of pairwise interactions revealed that legumes did not promote overyielding. Complementarity was mostly driven by Cistus , which always performed better when growing with other species than when growing with conspecifics. Our results are an addition to the still scarce literature dealing with diversity–productivity relationships in communities dominated by woody species, and show that methodologies commonly used to assess complementarity may not provide a precise estimation when a given species has negative effects on its conspecifics.     

The effects of species pool,dispersal and competition on the diversity–productivity relationship

Aim The diversity–productivity relationship is a controversial issue in ecology. Diversity is sometimes seen to increase with productivity but a unimodal relationship has often been reported. Competitive exclusion was cited initially to account for the decrease of diversity at high productivity. Subsequently, the roles of evolutionary history (species pool size) and dispersal rate have been acknowledged. We explore how the effects of species pool, dispersal and competition combine to produce different diversity–productivity relationships. Methods We use a series of simulations with a spatially explicit, individual‐based model. Following empirical expectations, we used four scenarios to characterize species pool size along the productivity gradient (uniformly low and high, linear increase and unimodal). Similarly, the dispersal rate varied along the productivity gradient (uniformly low and high, and unimodal). We considered both neutral communities and communities with competitive exclusion. Results and main conclusions Our model predicts that competitive interactions will result in unimodal diversity–productivity relationships. The model often predicts unimodal patterns in neutral communities as well, although the decline in richness at high productivity is less than in competing communities. A positive diversity–productivity relationship is simulated for neutral communities when the species pool size increases with productivity and the dispersal rate is high. This scenario is probably more widespread in nature than the others since positive diversity–productivity relationships have been observed more frequently than previously expected, especially in the tropics and for woody species. Our simulated effects of species pool, dispersal and competition on diversity patterns can be linked to empirical observations to uncover mechanisms behind the diversity–productivity relationship.     

Relationships among Bacterial Cell Size, Productivity, and Genetic Diversity in Aquatic Environments using Cell Sorting and Flow Cytometry

Abstract The study of relationships between cell size and productivity is of key importance in microbial ecology to understand which members of natural aquatic communities are responsible for the overall activity and/or productivity. Flow sorting of microorganisms from different environmental samples was used to analyze the activity of bacterial cells depending on their biovolume. Bacterial cells from five different natural samples taken along the Mediterranean coast including fresh- and seawaters were incubated with tritiated leucine, then stained with SYTO 13 and sorted by flow cytometry according to their average side-angle-scattered (SSC) light. In all samples, a bell-shaped relationship was found between cell biovolume and activity, whereas activity of a given cell-size class varied between samples. In contrast, an inverse relationship was found between biovolumes and abundances. These results suggest that medium-sized cells with highest growth rates are probably submitted to intense grazing. For one sample, bacteria within five different size classes were sorted and the genetic diversity of cells within each sorted size class and that of the whole community were analyzed by the denaturing gradient gel electrophoresis (DGGE) method. The genetic diversity, as determined at the community level was highly represented into the pool of small cells, whereas only few species were present into larger cell subpopulations. The results suggest that only a few genotypes may be dominant within the largest and most productive cells. Furthermore, cell size polymorphism as well as heterogeneous cellular activities were found within some species. Received: January 2000; Accepted: April 2000; Online Publication: 28 August 2000     

Invertebrate herbivory increases along an experimental gradient of grassland plant diversity

Plant diversity is a key driver of ecosystem functioning best documented for its influence on plant productivity. The strength and direction of plant diversity effects on species interactions across trophic levels are less clear. For example, with respect to the interactions between herbivorous invertebrates and plants, a number of competing hypotheses have been proposed that predict either increasing or decreasing community herbivory with increasing plant species richness. We investigated foliar herbivory rates and consumed leaf biomass along an experimental grassland plant diversity gradient in year eight after establishment. The gradient ranged from one to 60 plant species and manipulated also functional group richness (from one to four functional groups—legumes, grasses, small herbs, and tall herbs) and plant community composition. Measurements in monocultures of each plant species showed that functional groups differed in the quantity and quality of herbivory damage they experienced, with legumes being more damaged than grasses or non-legume herbs. In mixed plant communities, herbivory increased with plant diversity and the presence of two key plant functional groups in mixtures had a positive (legumes) and a negative (grasses) effect on levels of herbivory. Further, plant community biomass had a strong positive impact on consumed leaf biomass, but little effect on herbivory rates. Our results contribute detailed data from a well-established biodiversity experiment to a growing body of evidence suggesting that an increase of herbivory with increasing plant diversity is the rule rather than an exception. Considering documented effects of herbivory on other ecosystem functions and the increase of herbivory with plant diversity, levels of herbivory damage might not only be a result, but also a trigger within the diversity–productivity relationship.     

Diversity patterns of leaf-associated aquatic hyphomycetes along a broad latitudinal gradient

Information about the global distribution of aquatic hyphomycetes is scarce, despite the primary importance of these fungi in stream ecosystem functioning. In particular, the relationship between their diversity and latitude remains unclear, due to a lack of coordinated surveys across broad latitudinal ranges. This study is a first report on latitudinal patterns of aquatic hyphomycete diversity associated with native leaf-litter species in five streams located along a gradient extending from the subarctic to the tropics. Exposure of leaf litter in mesh bags of three different mesh sizes facilitated assessing the effects of including or excluding different size-classes of litter-consuming invertebrates. Aquatic hyphomycete evenness was notably constant across all sites, whereas species richness and diversity, expressed as the Hill number, reached a maximum at mid-latitudes (Mediterranean and temperate streams). These latitudinal patterns were consistent across litter species, despite a notable influence of litter identity on fungal communities at the local scale. As a result, the bell-shaped distribution of species richness and Hill diversity deviated markedly from the latitudinal patterns of most other groups of organisms. Differences in the body-size distribution of invertebrate communities colonizing the leaves had no effect on aquatic hyphomycete species richness, Hill diversity or evenness, but invertebrates could still influence fungal communities by depleting litter, an effect that was not captured by the design of our experiment.     

Species richness destabilizes ecosystem functioning in experimental aquatic microcosms

The relationship between species diversity and ecosystem stability has long interested ecologists, yet no consensus has been reached and the underlying mechanisms remain unclear. We used five unicellular algal species, cultured in all possible combinations, to assemble microcosms containing 1 to 5 algal species, on which a cold perturbation was imposed. Our aim was to find whether and how species richness begets ecosystem resistance and resilience. In the experiment, the species-rich communities produced more biomass than the species-poor ones, either in pre-, under- or post-perturbation conditions. The positive diversity–biomass relationship was weakened by the perturbation, and fully restored one week after the perturbation. The diverse communities showed greater absolute biomass reduction during the perturbation than did species-poor systems. Resistance of community, measured by the relative change in biomass from pre- to under-perturbation, decreased with species richness. All the species showed significant reduction in biomass when stressed, without any density compensation among species in diverse communities; and the ratio of biomass change in each species was independent of diversity. The overyielding effect, measured as relative yield total, remained constant from pre- to under-perturbation; and the selection and complementarity effects played equal roles for the biodiversity effect on biomass production, and their relative importance was not altered by the perturbation. These results suggest that similar responses of different species to environmental perturbations may limit the insurance effect of biodiversity, and lead to an inverse diversity–resistance relationship.     

Mechanisms responsible for the positive diversity–productivity relationship in Minnesota grasslands

Species’ extinctions have spurred debate on whether interactions among few or among many species cause a positive diversity–productivity relationship in experimentally assembled grasslands. We addressed this question by quantifying the productivity of 14 species across an experimental diversity gradient in Minnesota. We found that interspecific interactions leading to coexistence and competitive displacement both determine which species overyield; i.e. are more productive at high diversity. Overyielding species were either superior N competitors (C4 grasses) or N fixers (legumes). Surprisingly, these species were not most productive in monoculture, thus, the ‘selection’ of productive species in diverse plots did not cause the positive diversity–productivity relationship. Both positive (with legumes) and negative interspecific interactions (with C4 grasses) determined whether individual species overyielded. Foliar pathogens did not cause overyielding, although other natural enemies may be responsible. Overyielding species are not displacing underyielding species over time, implying that other diversity‐promoting interactions also operate in this experiment.