Ecological realism and mechanisms by which diversity begets stability

We investigated how ecological realism might impact the outcome of three experimental manipulations of species richness to determine whether the patterns and the mechanisms underlying richness–variability relationships differ as ecological communities are increasingly exposed to external forces that may drive richness–variability patterns in nature. To test for such an effect, we conducted experiments using rock pool meio‐invertebrate communities housed in three experimental venues: controlled laboratory microcosms, artificially constructed rock pools in the field, and naturally occurring rock pools in the field. Our results showed that experimental venue can have a strong effect on the outcome of richness manipulation experiments. As ecological realism increased, the strength of the relationship between species richness and community variability declined from 32.9% in the laboratory microcosms to 16.8% in the artificial pools to no effect of species richness on community variability in the natural rock pools. The determinants of community variability also differed as ecological realism increased. In laboratory microcosms, community variability was driven solely by mechanisms related to increasing species richness. In artificial rock pools, community variability was driven by a combination of direct and indirect environmental factors as well as mechanisms related to increasing species richness. In the natural rock pools community variability was independent of species richness and was only related to environmental factors. In summary, we found that stabilizing mechanisms associated with species interactions were influential in establishing species richness–variability relations only in the less realistic experimental venues (the laboratory microcosms and the artificial rock pools in the field), and that these mechanisms diminished in importance as ecological realism and complexity of the experimental venue increased. Our results suggest that the effects of diversity might be more difficult to detect in natural systems due to the combined effects of biotic and abiotic forcing, which can mask our ability to detect richness effects.     

Dominant species constrain effects of species diversity on temporal variability in biomass production of tallgrass prairie

Species diversity is thought to stabilize functioning of plant communities. An alternative view is that stability depends more on dynamics of dominant species than on diversity. We compared inter-annual variability (inverse of stability) of aboveground biomass in paired restored and remnant tallgrass prairies at two locations in central Texas, USA. Data from these two locations were used to test the hypothesis that greater richness and evenness in remnant than restored prairies would reduce variability in aboveground biomass in response to natural variation in rainfall. Restored prairies were chosen to be similar to paired remnant prairies in characteristics other than species diversity that affect temporal variability in biomass. Variability was measured as the coefficient of variation among years (square root of variance/mean; CV), where variance in community biomass equals the sum of variances of individual plant species plus the summed covariances between species pairs. Species diversity over five years was greater by a factor of 2 or more in remnant than restored prairies because richness and evenness were greater in remnant than restored prairies. Still, the CV of community biomass during spring and CV of annual biomass production did not differ consistently between prairie types. Neither the sum of species covariances nor total community biomass differed between prairies. Biomass varied relatively little in restored compared to remnant prairies because biomass of the dominant species in restored prairies (the grass Schizachyrium scoparium ) varied less than did biomass of other dominant and sub-dominant species. In these grasslands, biomass response to natural variation in precipitation depended as much on characteristics of a dominant grass as on differences in diversity.     

Spatial heterogeneity reduces temporal variability in stream insect communities

Although all natural systems are heterogeneous, the direct influence of spatial heterogeneity on most ecological variables is unknown. In many systems, spatial heterogeneity is positively correlated with both microhabitat refugia and species richness. Both an increased number of microhabitat refugia and the effects of statistical averaging via increased species richness should lead to an inverse relationship between spatial heterogeneity and variability in community composition. To test this prediction, I measured diversity and temporal variability of invertebrate communities in a northern New Hampshire stream along a natural gradient of spatial heterogeneity formed by variation in stream substrates. On average, there was a 42% decrease in community variability along a gradient of increasing heterogeneity. This pattern was robust to changes in metrics of both heterogeneity and community variability. There was also a significant positive relationship between taxon richness and spatial heterogeneity with predicted taxon richness increasing c. 1.5× along the heterogeneity gradient. By resampling community abundance data, I estimated that statistical averaging accounted for only 4% of the observed decrease in community variability in this study. I concluded that the remaining decrease was very likely explained by a greater number of refugia from predation and/or flooding in high‐heterogeneity habitats. The results of this study suggest that maximizing heterogeneity in ecological restoration programmes may promote temporally stable and diverse communities and may aid in responsible management of aquatic resources.     

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.     

Soil moisture's underestimated role in climate change impact modelling in low‐energy systems

Shifts in precipitation regimes are an inherent component of climate change, but in low‐energy systems are often assumed to be less important than changes in temperature. Because soil moisture is the hydrological variable most proximally linked to plant performance during the growing season in arctic‐alpine habitats, it may offer the most useful perspective on the influence of changes in precipitation on vegetation. Here we quantify the influence of soil moisture for multiple vegetation properties at fine spatial scales, to determine the potential importance of soil moisture under changing climatic conditions. A fine‐scale data set, comprising vascular species cover and field‐quantified ecologically relevant environmental parameters, was analysed to determine the influence of soil moisture relative to other key abiotic predictors. Soil moisture was strongly related to community composition, species richness and the occurrence patterns of individual species, having a similar or greater influence than soil temperature, pH and solar radiation. Soil moisture varied considerably over short distances, and this fine‐scale heterogeneity may contribute to offsetting the ecological impacts of changes in precipitation for species not limited to extreme soil moisture conditions. In conclusion, soil moisture is a key driver of vegetation properties, both at the species and community level, even in this low‐energy system. Soil moisture conditions represent an important mechanism through which changing climatic conditions impact vegetation, and advancing our predictive capability will therefore require a better understanding of how soil moisture mediates the effects of climate change on biota.     

The inherent multidimensionality of temporal variability: how common and rare species shape stability patterns

Empirical knowledge of diversity–stability relationships is mostly based on the analysis of temporal variability. Variability, however, often depends on external factors that act as disturbances, which makes comparisons across systems difficult to interpret. Here, we show how variability can reveal inherent stability properties of ecological communities. This requires that we abandon one‐dimensional representations, in which a single variability measurement is taken as a proxy for how stable a system is, and instead consider the whole set of variability values generated by all possible stochastic perturbations. Despite this complexity, in species‐rich systems, a generic pattern emerges from community assembly, relating variability to the abundance of perturbed species. Strikingly, the contrasting contributions of different species abundance classes to variability, driven by different types of perturbations, can lead to opposite diversity–stability patterns. We conclude that a multidimensional perspective on variability helps reveal the dynamical richness of ecological systems and the underlying meaning of their stability patterns.     

Stability and species richness in complex communities

Using both numerical simulations and analytical methods, we investigate how the stability of ecological communities depends on the number of species they contain. To investigate complex communities, we construct communities from modular "subcommunities" that can have arbitrary community structure; e.g. subcommunities could consist of pairs of predator and prey species, trios of prey, specialist predator and generalist predator, or any collection of interacting species. By building entire communities from subcommunities, we can change the number of species in the community without changing community structure. We further suppose that species sharing the same ecological role in different subcommunities act additively on the per capita population growth rates of other species. Under these assumptions, the inter-actions between species from different subcommunities have no effect on community-level stability, measured by the variability in the combined densities of species sharing the same ecological role in different subcommunities. Furthermore, increasing species richness (i.e. the number of subcommunities comprising the community) increases community-level stability only when it introduces species that respond differently to environmental fluctuations. Therefore, our results support the "insurance hypothesis" that species richness increases community-level stability by insuring that some species in a community are tolerant of different environmental fluctuations.     

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.     

Bottom–up and top–down forces structuring consumer communities in an experimental grassland

Synthesis The interplay between bottom‐up and top‐down effects is certainly a general manifestation of any changes in both species abundances and diversity. Summary variables, such as species numbers, diversity indices or lumped species abundances provide too limited information about highly complex ecosystems. In contrast, species by species analyses of ecological communities comprising hundreds of species are inevitably only snapshot‐like and lack generality in explaining processes within communities. Our synthesis, based on species matrices of functional groups of all trophic levels, simplifies community complexity to a manageable degree while retaining full species‐specific information. Taking into account plant species richness, plant biomass, soil properties and relevant spatial scales, we decompose variance of abundance in consumer functional groups to determine the direction and the magnitude of community controlling processes. After decades of intensive research, the relative importance of top–down and bottom–up control for structuring ecological communities is still a particularly disputed issue among ecologists. In our study, we determine the relative role of bottom–up and top–down forces in structuring the composition of 13 arthropod functional groups (FG) comprising different trophic consumer levels. Based on species‐specific plant biomass and arthropod abundance data from 50 plots of a grassland biodiversity experiment, we quantified the proportions of bottom–up and top–down forces on consumer FG composition while taking into account direct and indirect effects of plant diversity, functional diversity, community biomass, soil properties and spatial arrangement of these plots. Variance partitioning using partial redundancy analysis explained 21–44% of total variation in arthropod functional group composition. Plant‐mediated bottom–up forces accounted for the major part of the explainable variation within the composition of all FGs. Predator‐mediated top–down forces, however, were much weaker, yet influenced the majority of consumer FGs. Plant functional group composition, notably legume composition, had the most important impact on virtually all consumer FGs. Compared to plant species richness and plant functional group richness, plant community biomass explained a much higher proportion of variation in consumer community composition.     

Biodiversity may regulate the temporal variability of ecological systems

The effect of biodiversity on natural communities has recently emerged as a topic of considerable ecological interest. We review studies that explicitly test whether the number of species in a community (species richness) regulates the temporal variability of aggregate community (total biomass, productivity, nutrient cycling) and population (density, biomass) properties. Theoretical studies predict that community variability should decline with increasing species richness, while population variability should increase. Many, but not all, empirical studies support these expectations. However, a closer look reveals that several empirical studies have either imperfect experimental designs or biased methods of calculating variability. Furthermore, most theoretical studies rely on highly unrealistic assumptions. We conclude that evidence to support the claim that biodiversity regulates temporal variability is accumulating, but not unequivocal. More research, in a broader array of ecosystem types and with careful attention to methodological considerations, is needed before we can make definitive statements regarding richness‐variability relationships.     

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.     

Trait response in communities to environmental change: effect of interspecific competition and trait covariance structure

The response of ecological communities to environmental disturbances depends not just on the number of species they contain but also on the functional diversity of the constituent species; greater variation in the tolerance of species to different environmental disturbances is generally thought to confer greater resistance to the community. Here, I investigate how the functional diversity of communities changes with environmental disturbances. Specifically, I assume that there is variation in traits among species that confer tolerance or sensitivity to environmental disturbances. When a disturbance occurs, variation in species tolerances causes changes in the relative abundances of species, which in turn changes the average tolerance of the community. For example, if tolerance to an environmental disturbance is conferred by large body size, then the environmental disturbance should be expected to increase the average body size of individuals in the community. Despite this expectation, ecological interactions among species can affect the average community response. For example, if larger species are also strong competitors with each other, then this might reduce the increase in average body size in the community, because interspecific competition limits the grow in population density of large bodied species. Similarly, when disturbances affect multiple traits, the covariance in the distribution of trait values among species may restrict the response of any one trait; if two traits provide tolerance to the same disturbance but negatively covary among species, then the response of one trait will limit the response of the other trait at the community level. Using a Lotka–Volterra model for competitive communities, I derive general formulae that generate explicit predictions about the changes in average trait values in a community subject to environmental disturbances. These formulae demonstrate that competition can impede the change in average community trait values. However, the impediment is not considerable in comparison to the predominant factors of trait variances and species selection effects when species with the most similar trait values also experience the greatest interspecific competition. Similarly, negative covariances among different traits that confer resistance to the same environmental disturbance will impede their responses. I illustrate these results using phytoplankton data from a whole-lake experiment in which manipulation to the zooplankton community created a disturbance to the phytoplankton that changed the selective consumption of large vs. small phytoplankton.     

Process-based models of species distributions and the mid-domain effect

Null models that place species ranges at random within a bounded geographical domain produce hump-shaped species richness gradients (the "mid-domain effect," or MDE). However, there is debate about the extent to which these models are a suitable null expectation for effects of environmental gradients on species richness. Here, I present a process-based framework for modeling species distributions within a bounded geographical domain. Analysis of null models consistent with the mid-domain hypothesis shows that MDEs are indeed likely to be ubiquitous consequences of geographical domain boundaries. Comparing the probability distributions of range locations for the process-based and randomization-based models reveals that randomization models probably overestimate the contribution of MDEs to empirical patterns of species richness, but it also indicates that other testable predictions from randomization models are likely to be robust. I also show how this process-based framework can be extended beyond null models to incorporate effects of environmental gradients within the domain. This study provides a first step toward an ecological theory of species distributions in geographical space that can incorporate both "geometric constraints" and effects of environmental gradients, and it shows how such a theory can inform our understanding of species richness gradients in nature.     

Phylogenetic distance among beneficiary species in a cushion plant species explains interaction outcome

Determining which drivers lead to a specific species assemblage is a central issue in community ecology. Although many processes are involved, plant–plant interactions are among the most important. The phylogenetic limiting similarity hypothesis states that closely related species tend to compete stronger than distantly related species, although evidence is inconclusive. We used ecological and phylogenetic data on alpine plant communities along an environmental severity gradient to assess the importance of phylogenetic relatedness in affecting the interaction between cushion plants and the whole community, and how these interactions may affect community assemblage and diversity. We first measured species richness and individual biomass of species growing within and outside the nurse cushion species, Arenaria tetraquetra. We then assembled the phylogenetic tree of species present in both communities and calculated the phylogenetic distance between the cushion species and its beneficiary species, as well as the phylogenetic community structure. We also estimated changes in species richness at the local level due to the presence of cushions. The effects of cushions on closely related species changed from negative to positive as environmental conditions became more severe, while the interaction with distantly related species did not change along the environmental gradient. Overall, we found an environmental context‐dependence in patterns of phylogenetic similarity, as the interaction outcome between nurses and their close and distantly‐related species showed an opposite pattern with environmental severity.     

SESAM – a new framework integrating macroecological and species distribution models for predicting spatio‐temporal patterns of species assemblages

Two different approaches currently prevail for predicting spatial patterns of species assemblages. The first approach (macroecological modelling, MEM) focuses directly on realized properties of species assemblages, whereas the second approach (stacked species distribution modelling, S‐SDM) starts with constituent species to approximate the properties of assemblages. Here, we propose to unify the two approaches in a single ‘spatially explicit species assemblage modelling’ (SESAM) framework. This framework uses relevant designations of initial species source pools for modelling, macroecological variables, and ecological assembly rules to constrain predictions of the richness and composition of species assemblages obtained by stacking predictions of individual species distributions. We believe that such a framework could prove useful in many theoretical and applied disciplines of ecology and evolution, both for improving our basic understanding of species assembly across spatio‐temporal scales and for anticipating expected consequences of local, regional or global environmental changes. In this paper, we propose such a framework and call for further developments and testing across a broad range of community types in a variety of environments.     

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.     

Two-phase species–time relationships in North American land birds

The species–time relationship (STR) is a macroecological pattern describing the increase in the observed species richness with the length of time censused. Understanding STRs is important for understanding the ecological processes underlying temporal turnover and species richness. However, accurate characterization of the STR has been hampered by the influence of sampling. I analysed STRs for 521 breeding bird survey communities. I used a model of sampling effects to demonstrate that the increase in richness was not due exclusively to sampling. I estimated the time scale at which ecological processes became dominant over sampling effects using a two‐phase model combining a sampling phase and either a power function or logarithmic ecological phase. These two‐phase models performed significantly better than sampling alone and better than simple power and logarithmic functions. Most community dynamics were dominated by ecological processes over scales <5 years. This technique provides an example of a rigorous, quantitative approach to separating sampling from ecological processes.     

Weaving animal temperament into food webs: implications for biodiversity

Recent studies into community level dynamics are revealing processes and patterns that underpin the biodiversity and complexity of natural ecosystems. Theoretical food webs have suggested that species‐rich and highly complex communities are inherently unstable, but incorporating certain characteristics of empirical communities, such as allometric body size scaling and non‐random interaction distributions, have been shown to enhance stability and facilitate species coexistence. Incorporating individual level traits and variability into food web theory is seen as a future pathway for this research and our growing knowledge of individual behaviours, in the form of temperament (or personality) traits, can inform the direction of this research. Temperament traits are consistent differences in behaviour between individuals, which are repeatable across time and/or across ecological contexts, such as aggressive or boldness behaviours that commonly differ between individuals of the same species. These traits, under the framework of behavioural reaction norms, show both individual consistency as well as contextual and phenotypic plasticity. This is likely to contribute significantly to the effects of individual trait variability and adaptive trophic behaviour on the structure and dynamics of food webs, which are apparently stabilizing. Exploring the role of temperament in the context of community ecology is a unique opportunity for cross‐pollination between ecological fields, and can provide new insights into community stability and biodiversity.     

On the importance of intraspecific variability for the quantification of functional diversity

Functional diversity (FD) is a key facet of biodiversity used to address central questions in ecology. Despite recent methodological advances, FD remains a complex concept and no consensus has been reached either on how to quantify it, or on how it influences ecological processes. Here we define FD as the distribution of trait values within a community. When and how to account for intraspecific trait variability (ITV) when measuring FD remains one of the main current debates. It remains however unclear to what extent accounting for population‐level ITV would modify FD quantification and associated conclusions. In this paper, we address two critical questions: (1) How sensitive are different components of FD to the inclusion of population‐level ITV? (2) Does the omission of ITV obscure the understanding of ecological patterns? Using a mixture of empirical data and simulation experiments, we conducted a sensitivity analysis of four commonly used FD indices (community weighted mean traits, functional richness, Rao's quadratic entropy, Petchey and Gaston's FD index) and their relationships with environmental gradients and species richness, by varying both the extent (plasticity or not) and the structure (contingency to environmental gradient due to local adaptation) of population‐level ITV. Our results suggest that ITV may strongly alter the quantification of FD and the detection of ecological patterns. Our analysis highlights that 1) species trait values distributions within communities are crucial to the sensitivity to ITV, 2) ITV structure plays a major role in this sensitivity and 3) different indices are not evenly sensitive to ITV, the single‐trait FD from Petchey and Gaston being the most sensitive among the four metrics tested. We conclude that the effects of intraspecific variability in trait values should be more systematically tested before drawing central conclusions on FD, and suggest the use of simulation studies for such sensitivity analyses.     

A general multi-trait-based framework for studying the effects of biodiversity on ecosystem functioning

Environmental change is as multifaceted as are the species and communities that respond to these changes. Current theoretical approaches to modeling ecosystem response to environmental change often deal only with single environmental drivers or single species traits, simple ecological interactions, and/or steady states, leading to concern about how accurately these approaches will capture future responses to environmental change in real biological systems. To begin addressing this issue, we generalize a previous trait-based framework to incorporate aspects of frequency dependence, functional complementarity, and the dynamics of systems composed of species that are defined by multiple traits that are tied to multiple environmental drivers. The framework is particularly well suited for analyzing the role of temporal environmental fluctuations in maintaining trait variability and the resultant effects on community response to environmental change. Using this framework, we construct simple models to investigate two ecological problems. First, we show how complementary resource use can significantly enhance the nutrient uptake of plant communities through two different mechanisms related to increased productivity (over-yielding) and larger trait variability. Over-yielding is a hallmark of complementarity and increases the total biomass of the community and, thus, the total rate at which nutrients are consumed. Trait variability also increases due to the lower levels of competition associated with complementarity, thus speeding up the rate at which more efficient species emerge as conditions change. Second, we study systems in which multiple environmental drivers act on species defined by multiple, correlated traits. We show that correlations in these systems can increase trait variability within the community and again lead to faster responses to environmental change. The methodological advances provided here will apply to almost any function that relates species traits and environmental drivers to growth, and should prove useful for studying the effects of climate change on the dynamics of biota.