Species richness, environmental fluctuations, and temporal change in total community biomass

Theory and empirical results suggest that high biodiversity should often cause lower temporal variability in aggregate community properties such as total community biomass. We assembled microbial communities containing 2 to 8 species of competitors in aquatic microcosms and found that the temporal change in total community biomass was positively but insignificantly associated with diversity in a constant temperature environment. There was no evidence of any trend in variable temperature environments. Three non-exclusive mechanisms might explain the lack of a net stabilising effect of species richness on temporal change. (1) A direct destabilising effect of diversity on population level variances caused some populations to vary more when embedded in more diverse communities. (2) Similar responses of the different species to environmental variability might have limited any insurance effect of increased species richness. (3) Large differences in the population level variability of different species (i.e., unevenness) could weaken the relation between species richness and community level stability. These three mechanisms may outweigh the stabilising effects of increases in total community biomass with diversity, statistical averaging, and slightly more negative covariance in more diverse communities. Our experiment and analyses advocate for further experimental investigations of diversity-variability relations.     

Predicting ecosystem stability from community composition and biodiversity

As biodiversity is declining at an unprecedented rate, an important current scientific challenge is to understand and predict the consequences of biodiversity loss. Here, we develop a theory that predicts the temporal variability of community biomass from the properties of individual component species in monoculture. Our theory shows that biodiversity stabilises ecosystems through three main mechanisms: (1) asynchrony in species’ responses to environmental fluctuations, (2) reduced demographic stochasticity due to overyielding in species mixtures and (3) reduced observation error (including spatial and sampling variability). Parameterised with empirical data from four long‐term grassland biodiversity experiments, our prediction explained 22–75% of the observed variability, and captured much of the effect of species richness. Richness stabilised communities mainly by increasing community biomass and reducing the strength of demographic stochasticity. Our approach calls for a re‐evaluation of the mechanisms explaining the effects of biodiversity on ecosystem stability.     

Population and community variability in randomly fluctuating environments

The prediction that environmental fluctuations may destabilise populations and yet stabilise aggregate community properties has remained largely untested. We examined population and community stability under constant and fluctuating temperatures in simple planktonic assemblages of differing algal richness. Temperature dependent resource competition produced a highly asymmetric community structure where algal community biomass was dominated by one species. For a given level of species richness, temperature fluctuations induced lower community covariance and thus stabilised community biomass. However, increasing algal species richness increased the variability of population abundance and growth rates, as well as population and community variability. Consumer dynamics were directly destabilised by environmental fluctuations. These results confirm recent theoretical studies suggesting a stabilising effect of environmental fluctuations at the community level. However, they also support the theoretical prediction that increasing species richness may be of limited value for community stability, most especially in asymmetric communities, when competition directly affects population variability.     

Does Tropical Forest Fragmentation Increase Long-Term Variability of Butterfly Communities?

Habitat fragmentation is a major driver of biodiversity loss. Yet, the overall effects of fragmentation on biodiversity may be obscured by differences in responses among species. These opposing responses to fragmentation may be manifest in higher variability in species richness and abundance (termed hyperdynamism), and in predictable changes in community composition. We tested whether forest fragmentation causes long-term hyperdynamism in butterfly communities, a taxon that naturally displays large variations in species richness and community composition. Using a dataset from an experimentally fragmented landscape in the central Amazon that spanned 11 years, we evaluated the effect of fragmentation on changes in species richness and community composition through time. Overall, adjusted species richness (adjusted for survey duration) did not differ between fragmented forest and intact forest. However, spatial and temporal variation of adjusted species richness was significantly higher in fragmented forests relative to intact forest. This variation was associated with changes in butterfly community composition, specifically lower proportions of understory shade species and higher proportions of edge species in fragmented forest. Analysis of rarefied species richness, estimated using indices of butterfly abundance, showed no differences between fragmented and intact forest plots in spatial or temporal variation. These results do not contradict the results from adjusted species richness, but rather suggest that higher variability in butterfly adjusted species richness may be explained by changes in butterfly abundance. Combined, these results indicate that butterfly communities in fragmented tropical forests are more variable than in intact forest, and that the natural variability of butterflies was not a buffer against the effects of fragmentation on community dynamics.     

物种多样性与生态系统功能时间变异性的关系研究进展

近年来物种多样性的急剧丧失使得物种多样性与生态系统功能的时间变异性的关系及其机制问题的研究成为了生态学研究的一个热点。综述了物种多样性与群落集合性质变异性以及种群性质变异性的关系及其机制的最新研究成果：1、理论上探讨造成物种多样性与群落集合性质变异性负相关关系的机制包括：抽样效应、资源利用分化假说、统计平均效应、保险假说、种群变异性的均匀度效应等;但实验研究对理论预期的支持并不是普遍的;2．多样性与种群变异性之间的关系主要依赖于均值-方差尺度系数Z;理论上大部分自然群落是种群变异性应该随着多样性的增加而增加;但有研究表明：在变动环境中多样性对单个组分物种的种群水平有稳定性作用;而经验研究并不能得出多样性对种群变异性效应的清晰模式。讨论了目前的理论和实验研究中存在的和今后研究中需要认真思考的问题。     

Linking biodiversity to ecosystem function: implications for conservation ecology

We evaluate the empirical and theoretical support for the hypothesis that a large proportion of native species richness is required to maximize ecosystem stability and sustain function. This assessment is important for conservation strategies because sustenance of ecosystem functions has been used as an argument for the conservation of species. If ecosystem functions are sustained at relatively low species richness, then arguing for the conservation of ecosystem function, no matter how important in its own right, does not strongly argue for the conservation of species. Additionally, for this to be a strong conservation argument the link between species diversity and ecosystem functions of value to the human community must be clear. We review the empirical literature to quantify the support for two hypotheses: (1) species richness is positively correlated with ecosystem function, and (2) ecosystem functions do not saturate at low species richness relative to the observed or experimental diversity. Few empirical studies demonstrate improved function at high levels of species richness. Second, we analyze recent theoretical models in order to estimate the level of species richness required to maintain ecosystem function. Again we find that, within a single trophic level, most mathematical models predict saturation of ecosystem function at a low proportion of local species richness. We also analyze a theoretical model linking species number to ecosystem stability. This model predicts that species richness beyond the first few species does not typically increase ecosystem stability. One reason that high species richness may not contribute significantly to function or stability is that most communities are characterized by strong dominance such that a few species provide the vast majority of the community biomass. Rapid turnover of species may rescue the concept that diversity leads to maximum function and stability. The role of turnover in ecosystem function and stability has not been investigated. Despite the recent rush to embrace the linkage between biodiversity and ecosystem function, we find little support for the hypothesis that there is a strong dependence of ecosystem function on the full complement of diversity within sites. Given this observation, the conservation community should take a cautious view of endorsing this linkage as a model to promote conservation goals. Received: 2 September 1999 / Accepted: 26 October 1999     

Nutrient enrichment weakens the stabilizing effect of species richness

With global freshwater biodiversity declining at an even faster rate than in the most disturbed terrestrial ecosystems, understanding the effects of changing environmental conditions on relationships between biodiversity and the variability of community and population processes in aquatic ecosystems is of significant interest. Evidence is accumulating that biodiversity loss results in more variable communities; however, the mechanisms underlying this effect have been the subject of considerable debate. We manipulated species richness and nutrients in outdoor aquatic microcosms composed of naturally occurring assemblages of zooplankton and benthic invertebrates to determine how the relationship between species richness and variability might change under different nutrient conditions. Temporal variability of populations and communities decreased with increasing species richness in low nutrient microcosms. In contrast, we found no relationship between species richness and either population or community variability in nutrient enriched microcosms. Of the different mechanisms we investigated (e.g. overyielding, statistical averaging, insurance effects, and the stabilizing effect of species richness on populations) the only one that was consistent with our results was that increases in species richness led to more stable community abundances through the stabilizing effect of species richness on the component populations. While we cannot conclusively determine the mechanism(s) by which species richness stabilized populations, our results suggest that more complete resource-use in the more species-rich low nutrient communities may have dampened population fluctuations.     

Spatio-temporal dynamics of species richness in coastal fish communities

Determining patterns of change in species richness and the processes underlying the dynamics of biodiversity are of key interest within the field of ecology, but few studies have investigated the dynamics of vertebrate communities at a decadal temporal scale. Here, we report findings on the spatio-temporal variability in the richness and composition of fish communities along the Norwegian Skagerrak coast having been surveyed for more than half a century. Using statistical models incorporating non-detection and associated sampling variance, we estimate local species richness and changes in species composition allowing us to compute temporal variability in species richness. We tested whether temporal variation could be related to distance to the open sea and to local levels of pollution. Clear differences in mean species richness and temporal variability are observed between fjords that were and were not exposed to the effects of pollution. Altogether this indicates that the fjord is an appropriate scale for studying changes in coastal fish communities in space and time. The year-to-year rates of local extinction and turnover were found to be smaller than spatial differences in community composition. At the regional level, exposure to the open sea plays a homogenizing role, possibly due to coastal currents and advection.     

Opposite effects of environmental variability and species richness on temporal turnover of species in a complex habitat mosaic

Species are continuously lost and added to a local community. Dynamics of this process in a complex habitat mosaic (multiple habitats in a landscape), particularly of its rates (species turnover) are of primary concern for biodiversity conservation. Various studies suggest that species traits such as habitat specialization should affect species turnover. In communities where habitat specialization is a function of abiotic constraints, habitat specialists should respond faster to changing environment than generalists. We thus predicted a higher temporal turnover for specialists than for generalists in the presence of environmental variability (EV). In addition, we predicted that temporal turnover should decrease with increasing species richness of the communities they live in. We tested these predictions in a model system of 49 natural rock pools inhabited by 70 invertebrate species for which long-term (9 years) environmental and population dynamics data are available. We computed standard deviation of salinity measurements to represent EV for each pool. We further obtained the number of combined colonization and extinction events weighted by the number of years a species was recorded as a temporal turnover for each species in individual pools. We found that EV induced greater temporal turnover, however, the turnover depended on the species habitat traits (habitat specialization)—it has been higher in specialists but that relationship between EV and temporal turnover dissolved with increasing niche breadth (generalists). We further found that for some species, temporal turnover decreased with higher species richness and for other species, temporal turnover increased with higher species richness. The effect of species richness on temporal turnover was unrelated to species traits. This study suggests that whenever habitat is complex and heterogeneous and species pool diversified, local community dynamics becomes a composite of differential responses.     

Diversity promotes temporal stability across levels of ecosystem organization in experimental grasslands

The diversity-stability hypothesis states that current losses of biodiversity can impair the ability of an ecosystem to dampen the effect of environmental perturbations on its functioning. Using data from a long-term and comprehensive biodiversity experiment, we quantified the temporal stability of 42 variables characterizing twelve ecological functions in managed grassland plots varying in plant species richness. We demonstrate that diversity increases stability i) across trophic levels (producer, consumer), ii) at both the system (community, ecosystem) and the component levels (population, functional group, phylogenetic clade), and iii) primarily for aboveground rather than belowground processes. Temporal synchronization across studied variables was mostly unaffected with increasing species richness. This study provides the strongest empirical support so far that diversity promotes stability across different ecological functions and levels of ecosystem organization in grasslands.     

Biodiversity, productivity and the temporal stability of productivity: patterns and processes

Theory predicts that the temporal stability of productivity, measured as the ratio of the mean to the standard deviation of community biomass, increases with species richness and evenness. We used experimental species mixtures of grassland plants to test this hypothesis and identified the mechanisms involved. Additionally, we tested whether biodiversity, productivity and temporal stability were similarly influenced by particular types of species interactions. We found that productivity was less variable among years in plots planted with more species. Temporal stability did not depend on whether the species were planted equally abundant (high evenness) or not (realistically low evenness). Greater richness increased temporal stability by increasing overyielding, asynchrony of species fluctuations and statistical averaging. Species interactions that favoured unproductive species increased both biodiversity and temporal stability. Species interactions that resulted in niche partitioning or facilitation increased both productivity and temporal stability. Thus, species interactions can promote biodiversity and ecosystem services.     

Effect of (a)synchronous light fluctuation on diversity,functional and structural stability of a marine phytoplankton metacommunity

Disentangling the mechanisms that maintain the stability of communities and ecosystem properties has become a major research focus in ecology in the face of anthropogenic environmental change. Dispersal plays a pivotal role in maintaining diversity in spatially subdivided communities, but only a few experiments have simultaneously investigated how dispersal and environmental fluctuation affect community dynamics and ecosystem stability. We performed an experimental study using marine phytoplankton species as model organisms to test these mechanisms in a metacommunity context. We established three levels of dispersal and exposed the phytoplankton to fluctuating light levels, where fluctuations were either spatially asynchronous or synchronous across patches of the metacommunity. Dispersal had no effect on diversity and ecosystem function (biomass), while light fluctuations affected both evenness and community biomass. The temporal variability of community biomass was reduced by fluctuating light and temporal beta diversity was influenced interactively by dispersal and fluctuation, whereas spatial variability in community biomass and beta diversity were barely affected by treatments. Along the establishing gradient of species richness and dominance, community biomass increased but temporal variability of biomass decreased, thus highest stability was associated with species-rich but highly uneven communities and less influenced by compensatory dynamics. In conclusion, both specific traits (dominance) and diversity (richness) affected the stability of metacommunities under fluctuating conditions.     

Impact of exotic invasion on the temporal stability of natural annual plant communities

Much work in ecology has focused on understanding how changes in community diversity and composition will affect the temporal stability of communities (the degree of fluctuations in community abundance or biomass over time). While theory suggests diversity and dominant species can enhance temporal stability, empirical work has tended to focus on testing the effect of diversity, often using synthetic communities created with high species evenness. We use a complementary approach by studying the temporal stability of natural plant communities invaded by a dominant exotic, Erodium cicutarium. Invasion was associated with a significant decline in community diversity and change in the identity of the dominant species allowing us to evaluate predictions about how these changes might affect temporal stability. Community temporal stability was not correlated with community richness or diversity prior to invasion. Following invasion, community stability was again not correlated with community richness but was negatively correlated with community diversity. Before and after invasion, community stability was positively correlated with the stability of the most dominant species in the community, even though the identity of the dominant species changed from a native (prior to invasion) to an exotic species. Our results demonstrate that invasion by a dominant exotic species may reduce diversity without negatively affecting the temporal stability of natural communities. These findings add support to the idea that dominant species can strongly affect temporal stability, independent of community diversity.     

Species richness–variability relationships in multi-trophic aquatic microcosms

While species loss may affect the temporal variability of populations and communities differently in multi- versus single-trophic level communities, the nature of these differences are poorly understood. Here, we report on an experiment where we manipulated species richness of multi-trophic rock pool invertebrate communities to determine the effects of species richness, S, on the temporal variability of communities, populations, and individual species. As in single-trophic level studies, temporal variability in community abundance decreased with increasing species richness. However, in contrast to most studies in single-trophic level systems, temporal variability of populations also decreased as species richness increased. Furthermore, the variability of the constituent populations strongly correlated with variability of community abundance suggesting that, in rock pools, S affects community variability through its stabilizing effect on component populations. Our results suggest that species loss may affect population and community variability differently in multi-trophic versus single trophic level communities. If this is so, then the mechanisms proposed to underlie the effects of S on community variability in single-trophic communities may have to be supplemented by those that describe contributions to population stability in order to fully describe the patterns observed in multi-trophic communities.     

Using root traits to understand temporal changes in biodiversity effects in grassland mixtures

Biodiversity–ecosystem functioning (BEF) studies typically show that species richness enhances community biomass, but the underlying mechanisms remain debated. Here, we combine metrics from BEF research that distinguish the contribution of dominant species (selection effects, SE) from those due to positive interactions such as resource partitioning (complementarity effects, CE) with a functional trait approach in an attempt to reveal the functional characteristics of species that drive community biomass in species mixtures. In a biodiversity experiment with 16 plant species in monocultures, 4‐species and 16‐species mixtures, we used aboveground biomass to determine the relative contributions of CE and SE to biomass production in mixtures in the second, dry year of the experiment. We also measured root traits (specific root length, root length density, root tissue density and the deep root fraction) of each species in monocultures and linked the calculated community weighted mean (CWM) trait values and trait diversity of mixtures to CE and SE. In the second year of the experiment, community biomass, CE and SE increased compared to the first year. The contribution of SE to this positive effect was greater than that of CE. The increased contribution of SE was associated with root traits: SE increased most in communities with high abundance of species with deep, thick and dense roots. In contrast, changes in CE were not related to trait diversity or CWM trait values. Together, these results suggest that increased positive effects of species richness on community biomass in a dry year were mainly driven by increased dominance of deep‐rooting species, supporting the insurance hypothesis of biodiversity. Positive CE indicates that other positive interactions did occur, but we could not find evidence that belowground resource partitioning or facilitation via root trait diversity was important for community productivity in our biodiversity experiment.     

Herbivores control effects of algal species richness on community biomass and stability in a laboratory microcosm experiment

Hundreds of studies that have explored how biodiversity affects the productivity and stability of ecosystems have produced a consensus that communities composed of more species tend to have higher biomass that is more stable through time. However, the majority of this work stems from studies performed using highly simplified food webs, often composed of just primary producers competing for inorganic resources in the absence of trophic interactions. When studies have incorporated trophic interactions, diversity‐function relationships have been more variable, leaving open the question of how biodiversity affects the functioning of ecosystems with more trophic levels. Here we report the results of a laboratory experiment that used freshwater microcosms to test for effects of algal diversity (one or four species) on community biomass and temporal variability in the presence and absence of two different herbivore species (cladocerans Ceriodaphnia dubia and Daphnia pulex). When no herbivores were present, we found the classic pattern observed in hundreds of other studies – as species richness of algae increased, algal biomass increased, and the temporal variation in biomass decreased. This pattern was retained when one of the herbivores (C. dubia) was present. Ceriodaphnia dubia exhibited weak and non‐selective grazing on the focal algae, leaving the effect of diversity on biomass and variability essentially intact. In contrast, D. pulex exhibited strong and selective grazing in algal polycultures that qualitatively altered both diversity–function relationships. As algal richness increased, total algal biomass decreased and variation through time increased. These changes were coupled with larger and less variable populations of D. pulex. Our results show that herbivory leads to a richer array of diversity–function relationships than often observed in studies focused on just one trophic level, and suggests trophic interactions should be given more attention in work that seeks to determine how biodiversity impacts the functioning of ecosystems.     

Nutrient concentrations and fibre contents of plant community biomass reflect species richness patterns along a broad range of land-use intensities among agricultural grasslands

Understanding changes in biodiversity in agricultural landscapes in relation to land-use type and intensity is a major issue in current ecological research. In this context nutrient enrichment has been identified as a key mechanism inducing species loss in Central European grassland ecosystems. At the same time, insights into the linkage between agricultural land use and plant nutrient status are largely missing. So far, studies on the relationship between chemical composition of plant community biomass and biodiversity have mainly been restricted to wetlands and all these studies neglected the effects of land use. Therefore, we analyzed aboveground biomass of 145 grassland plots covering a gradient of land-use intensities in three regions across Germany. In particular, we explored relationships between vascular plant species richness and nutrient concentrations as well as fibre contents (neutral and acid detergent fibre and lignin) in the aboveground community biomass.We found the concentrations of several nutrients in the biomass to be closely linked to plant species richness and land use. Whereas phosphorus concentrations increased with land-use intensity and decreased with plant species richness, nitrogen and potassium concentrations showed less clear patterns. Fibre fractions were negatively related to nutrient concentrations in biomass, but hardly to land-use measures and species richness. Only high lignin contents were positively associated with species richness of grasslands. The N:P ratio was strongly positively related to species richness and even more so to the number of endangered plant species, indicating a higher persistence of endangered species under P (co-)limited conditions. Therefore, we stress the importance of low P supply for species-rich grasslands and suggest the N:P ratio in community biomass to be a useful proxy of the conservation value of agriculturally used grasslands.     

Temperature mean and variance alter phytoplankton biomass and biodiversity in a long‐term microcosm experiment

Global predictions of increasing mean temperature and its variance result in concerns about the fate of biodiversity under future climatic conditions. On the local scale of interacting species, the response of diversity to warming will depend much on the change in interspecific interactions such as competition or predation. Using a phytoplankton model community, we conducted a 2 × 2 × 2 factorial long‐term microcosm experiment (16 months) with an underlying seasonal temperature profile. We manipulated the mean temperature (T_mean), the temperature variance (T_var), and the presence of consumers (ciliates) and monitored treatment effects on algal biomass, species composition and species richness. Temperature effects on algal biomass depended on the seasonal development and consumer presence. Algal biomass decreased with increasing T_mean at consumer absence, but increased at consumer presence. Overall, algal biomass decreased with increasing T_var. Consumer presence reduced algal biomass from summer to winter, but then ciliates and the consumer effect disappeared. Almost all treatment combinations collapsed to monocultures after 16 months, but extinction occurred faster at higher T_mean (especially at consumer absence) and slower at consumer presence (especially at higher T_mean). Contrasting our predictions, increasing T_var reduced richness and increased extinction rate. The treatment effects on biomass and richness were not independent, as algal species richness and biomass were positively correlated. Moreover, accelerated loss of species was consistently correlated to higher temporal variability in biomass. In conclusion, altered temperature regimes strongly affected algal biomass and diversity by interdependently altering competitive and consumer interactions.     

Adaptive survival mechanisms and growth limitations of small-stature herb species across a plant diversity gradient

Several biodiversity experiments have shown positive effects of species richness on aboveground biomass production, but highly variable responses of individual species. The well-known fact that the competitive ability of plant species depends on size differences among species, raises the question of effects of community species richness on small-stature subordinate species. We used experimental grasslands differing in species richness (1-60 species) and functional group richness (one to four functional groups) to study biodiversity effects on biomass production and ecophysiological traits of five small-stature herbs (Bellis perennis, Plantago media, Glechoma hederacea, Ranunculus repens and Veronica chamaedrys). We found that ecophysiological adaptations, known as typical shade-tolerance strategies, played an important role with increasing species richness and in relation to a decrease in transmitted light. Specific leaf area and leaf area ratio increased, while area-based leaf nitrogen decreased with increasing community species richness. Community species richness did not affect daily leaf carbohydrate turnover of V. chamaedrys and P. media indicating that these species maintained efficiency of photosynthesis even in low-light environments. This suggests an important possible mechanism of complementarity in such grasslands, whereby smaller species contribute to a better overall efficiency of light use. Nevertheless, these species rarely contributed a large proportion to community biomass production or achieved higher yields in mixtures than expected from monocultures. It seems likely that the allocation to aboveground plant organs to optimise carbon assimilation limited the investment in belowground organs to acquire nutrients and thus hindered these species from increasing their performance in multi-species mixtures.     

Long‐term study of root biomass in a biodiversity experiment reveals shifts in diversity effects over time

Biodiversity experiments generally report a positive effect of plant biodiversity on aboveground biomass (overyielding), which typically increases with time. Various studies also found overyielding for belowground plant biomass, but this has never been measured over time. Also, potential underlying mechanisms have remained unclear. Differentiation in rooting patterns among plant species and plant functional groups has been proposed as a main driver of the observed biodiversity effect on belowground biomass, leading to more efficient belowground resource use with increasing diversity, but so far there is little evidence to support this. We analyzed standing root biomass and its distribution over the soil profile, along a 1–16 species richness gradient over eight years in the Jena Experiment in Germany, and compared belowground to aboveground overyielding. In our long‐term dataset, total root biomass increased with increasing species richness but this effect was only apparent after four years. The increasingly positive relationship between species richness and root biomass, explaining 12% of overall variation and up to 28% in the last year of our study, was mainly due to decreasing root biomass at low diversity over time. Functional group composition strongly affected total standing root biomass, explaining 44% of variation, with grasses and legumes having strong overall positive and negative effects, respectively. Functional group richness or interactions between functional group presences did not strongly contribute to overyielding. We found no support for the hypothesis that vertical root differentiation increases with species richness, with functional group richness or composition. Other explanations, such as stronger negative plant–soil feedbacks in low‐diverse plant communities on standing root biomass and vertical distribution should be considered.