Does species richness drive community production or vice versa? Reconciling historical and contemporary paradigms in competitive communities

Studies examining the relationship between species richness and the productivity of ecological communities have taken one of two opposite viewpoints, viewing either productivity as a primary driver of richness or richness as a driver of productivity. Recently, verbal and graphical hypotheses have been proposed that attempt to merge these perspectives by clarifying the causal pathways that link resource supply, species richness, resource use, and biomass production. Here we present mathematical models that formalize how these pathways can operate simultaneously in a single ecological system. Using a metacommunity framework in which classic consumer-resource competition theory governs species interactions within patches, we show that the mechanisms by which resource supply influences species richness are inherently linked to the mechanisms by which species richness controls resource use and biomass production. Unlike prior hypotheses, our models show that resource supply can affect species richness and that richness can affect productivity simultaneously at a single spatial scale. Our models also reproduce scale-dependent associations between species richness and community biomass that have been reported elsewhere. By detailing the pathways by which resource supply, species richness, biomass production, and resource use are connected, our models move closer to resolving the nature of causality in diversity-productivity relationships.     

Coexistence and community structure in a consumer resource model with implicit stoichiometry

We combine stoichiometry theory and optimal foraging theory into the MacArthur consumer-resource model. This generates predictions for diet choice, coexistence, and community structure of heterotroph communities. Tradeoffs in consumer resource-garnering traits influence community outcomes. With scarce resources, consumers forage opportunistically for complementary resources and may coexist via tradeoffs in resource encounter rates. In contrast to single currency models, stoichiometry permits multiple equilibria. These alternative stable states occur when tradeoffs in resource encounter rates are stronger than tradeoffs in elemental conversion efficiencies. With abundant resources consumers exhibit partially selective diets for essential resources and may coexist via tradeoffs in elemental conversion efficiencies. These results differ from single currency models, where adaptive diet selection is either opportunistic or selective. Interestingly, communities composed of efficient consumers share many of the same properties as communities based on substitutable resources. However, communities composed of relatively inefficient consumers behave similarly to plant communities as characterized by Tilman’s consumer resource theory. The results of our model indicate that the effects of stoichiometry theory on community ecology are dependent upon both consumer foraging behavior and the nature of resource garnering tradeoffs.     

Constrained proteome allocation affects coexistence in models of competitive microbial communities

Microbial communities are ubiquitous and play crucial roles in many natural processes. Despite their importance for the environment, industry and human health, there are still many aspects of microbial community dynamics that we do not understand quantitatively. Recent experiments have shown that the structure and composition of microbial communities are intertwined with the metabolism of the species that inhabit them, suggesting that properties at the intracellular level such as the allocation of cellular proteomic resources must be taken into account when describing microbial communities with a population dynamics approach. In this work, we reconsider one of the theoretical frameworks most commonly used to model population dynamics in competitive ecosystems, MacArthur’s consumer-resource model, in light of experimental evidence showing how proteome allocation affects microbial growth. This new framework allows us to describe community dynamics at an intermediate level of complexity between classical consumer-resource models and biochemical models of microbial metabolism, accounting for temporally-varying proteome allocation subject to constraints on growth and protein synthesis in the presence of multiple resources, while preserving analytical insight into the dynamics of the system. We first show with a simple experiment that proteome allocation needs to be accounted for to properly understand the dynamics of even the simplest microbial community, i.e. two bacterial strains competing for one common resource. Then, we study our consumer-proteome-resource model analytically and numerically to determine the conditions that allow multiple species to coexist in systems with arbitrary numbers of species and resources.Subject terms: Biodiversity, Microbial ecology, Microbial ecology, Bacterial physiology     

Biological invasions and the neutral theory

Aim  The invasion of natural communities by alien species represents a serious threat, but creates opportunities to learn about community functions. Neutral theory proposes that the niche concept may not be needed to explain the assemblage and diversity of natural communities, challenging the classical view of community ecology and generating a lasting debate. Biological invasions, when considered as natural experiments, can be used to contrast some of the predictions of neutral and classic niche theories.
Location  Global.
Methods  We use data from biological invasions as natural experiments to contrast some of the fundamental predictions of neutral theory.
Results  Some emerging patterns did not differ from neutral model expectations (e.g. the relationship between native and exotic species richness, invasibility of resource-rich habitats, and the relationship between propagule release and invasion success). Nevertheless, other patterns (e.g. experimental evidence of the relationship between diversity and susceptibility to invasion, the invasion of communities with a low resource availability, invasiveness related to species traits) contrasted with the predictions that can be inferred from neutral theory.
Main conclusions  Neutral theory correctly highlights the need to include randomness in models of community structure. Biological invasion patterns show that neutral forces are important in structuring natural communities, but the patterns differ from those inferred from a complete neutral model. For biodiversity-conservation purposes, the implications of accepting or not accepting neutral theory as a process-based theory are very important.     

Prediction of species composition of plant communities in a rural landscape based on species traits

The species composition of a community is a subset of the regional species pool, and predicting the species composition of a community from ecological traits of organisms is an important objective in ecology. If such a prediction can be made feasible, we could assess the risk of invasion of locally new species (alien species and genetically modified species) into natural communities. We developed and tested statistical models to predict a community’s species composition from ecological traits of the species pool. Various types of communities (forest, meadow, and weed communities) exist in a small area of traditional rural landscape in Japan, and have been maintained by human activities. These communities and the tracheophytes species pool in the 1-km² research area were considered. We used logistic regression and decision-tree analysis to construct predictive models of community species composition based on plant traits, using the presence or absence of species in a community as the dependent variable and ecological traits as independent variables. Plant traits were grouped by cluster analysis, and the average in each trait group was used for model building to avoid multiple collinearity. Statistical prediction models were significant in all communities. About 60–75% of species composition could be predicted from the measured plant traits in forest communities, but 33–56% in the meadow and weed communities. Our results showed the possibility of predicting the species composition of plant communities from the ecological traits of the plant species together with the information on local species pool.     

Adaptive change in the resource-exploitation traits of a generalist consumer: the evolution and coexistence of generalists and specialists

Mathematical models of consumer-resource systems are used to explore the evolution of traits related to resource acquisition in a generalist consumer species that is capable of exploiting two resources. The analysis focuses on whether evolution of traits determining the capture rates of two resources by a consumer species produce one generalist, two specialists, or all three types, when all types are characterized by a common fitness function. In systems with a stable equilibrium, evolution produces one generalist or two specialists, depending on the second derivative of the trade-off relationship. When there are sustained population fluctuations, the nature of the trade-off between the consumer's capture rates of the two resources still plays a key role in determining the evolutionary outcome. If the trade-off is described by a choice variable between zero and one that is raised to a power n, polymorphic states are possible when n > 1, which implies a positive second derivative of the curve. These states are either dimorphism, with two relatively specialized consumer types, or trimorphism, with a single generalist type and two specialists. Both endogenously driven consumer-resource cycles, and fluctuations driven by an environmental variable affecting resource growth are considered. Trimorphic evolutionary outcomes are relatively common in the case of endogenous cycles. In contrast to a previous study, these trimorphisms can often evolve even when new lineages are constrained to have phenotypes very similar to existing lineages. Exogenous cycles driven by environmental variation in resource growth rates appear to be much less likely to produce a mixture of generalists and specialists than are endogenous consumer-resource cycles.     

Integrating phylogenetic community structure with species distribution models: an example with plants of rock barrens

Approaches using phylogenetic pattern in ecological communities to deduce processes of community assembly have been criticised as disconnected from foundations in ecological mechanism, especially with respect to lack of data about abiotic and biotic niches. These criticisms can be addressed with analyses of organismal traits that underlie environmental filtering, competitive exclusion, and other candidate processes; however, the difficulty of assembling large trait databases means that such studies remain uncommon. We suggest a synthesis of phylogenetic community structure analysis and species distribution modeling that we believe can allow inference about community processes without prohibitive data requirements. We illustrate this method for angiosperm communities of rock barrens in eastern Canada. First, we analyzed phylogenetic community structure of four rock‐barren sites at three nested spatial scales (quadrat to region). For the nine most common species in our barrens, we used regional occurrence records to build species distribution models identifying environmental drivers of the nine species’ distributions. Coefficients of these models represent implicit trait data that summarize each species’ response to the environmental drivers in the model. We then tested for phylogenetic signal in these traits, to ask whether ecological forces acting on them could be generating phylogenetic community structure. We found strong phylogenetic clustering at the quadrat level, while patterns at larger scales were complex. Our distribution model suggested drought stress as the dominant driver for distributions of all the species, consistent with local correlations with soil depth, and the species’ responses to drought showed strong phylogenetic signal. The convergence of results from phylogenetic community structure and species distribution modeling suggests that barren communities are structured at the quadrat level by environmental filtering effects of moisture stress, to which species have phylogenetically patterned responses.     

Phylogenetic analysis of trophic associations

Ecologists frequently collect data on the patterns of association between adjacent trophic levels in the form of binary or quantitative food webs. Here, we develop statistical methods to estimate the roles of consumer and resource phylogenies in explaining patterns of consumer-resource association. We use these methods to ask whether closely related consumer species are more likely to attack the same resource species and whether closely related resource species are more likely to be attacked by the same consumer species. We then show how to use estimates of phylogenetic signals to predict novel consumer-resource associations solely from the phylogenetic position of species for which no other (or only partial) data are available. Finally, we show how to combine phylogenetic information with information about species' ecological characteristics and life-history traits to estimate the effects of species traits on consumer-resource associations while accounting for phylogenies. We illustrate these techniques using a food web comprising species of parasitoids, leaf-mining moths, and their host plants.     

Discrete consumers,small scale resource heterogeneity,and population stability

We present a consumer-resource model in which individual consumers subsist on a continuum of resource distributed over a very large number of small “bite-sized” patches, each patch being sufficiently small that all its resource is eaten whenever a consumer visits. This form of consumer–resource interaction forces a heterogeneous distribution of resource among the patches, and may dampen out the large amplitude, consumer-resource cycles that are predicted by traditional models of well-mixed, spatially homogeneous systems. The resource equilibrium does not increase with enrichment, a prediction that distinguishes this model from models that invoke direct or indirect consumer density dependence as a stabilizing mechanism.     

Subsidies mediate interactions between communities across space

Most spatial ecology focuses on how species dispersal affects community dynamics and coexistence. Ecosystems, however, are also commonly connected by flows of resources. We experimentally tested how neighbouring communities indirectly influence each other in absence of dispersal, via resource exchanges. Using two‐patch microcosm meta‐ecosystems, we manipulated community composition and dynamics, by varying separately species key functional traits (autotroph versus heterotroph species and size of consumer species) and trophic structure of aquatic communities (species growing alone or in presence of competitors or predators). We then analysed the effects of species functional traits and trophic structure on communities connected through spatial subsidies in the absence of actual dispersal. Both functional traits and trophic structure strongly affected dynamics across neighbouring communities. Heterotroph communities connected to autotroph neighbours developed better than with heterotroph neighbours, such that coexistence of competitors was determined by the functional traits of the neighbouring community. Densities in autotroph communities were also strikingly higher when receiving subsidies from heterotroph communities compared to their own subsidies when grown in isolated ecosystems. In contrast, communities connected to predator‐dominated ecosystems collapsed, without any direct contact with the predators. Our results demonstrate that because community composition and structure modify the distribution of biomass within a community, they may also affect communities connected through subsidies through quantitative and qualitative changes of detritus flows. This stresses that ecosystem management should account for such interdependencies mediated by spatial subsidies, given that local community alterations cascade across space onto other ecosystems even if species dispersal is completely absent.     

Thinking inside the box: community-level consequences of stage-structured populations

Ecologists have historically represented consumer-resource interactions with boxes and arrows. A key assumption of this conceptualization is that all individuals inside a box are functionally equivalent. Demographic stage structure, however, is a widespread source of heterogeneity inside the boxes. Synthesizing recent studies, we show that stage structure can modify the dynamics of consumer-resource communities owing to stage-related shifts in the nature and strength of interactions that occur within and between populations. As a consequence, stage structure can stabilize consumer-resource dynamics, create possibilities for alternative community states, modify conditions for coexistence of competitors, and alter the strength and direction of trophic cascades. Consideration of stage structure can thus lead to outcomes that are not expected based on unstructured approaches.     

A mechanistic approach for modeling temperature-dependent consumer-resource dynamics

Paramount to our ability to manage and protect biological communities from impending changes in the environment is an understanding of how communities will respond. General mathematical models of community dynamics are often too simplistic to accurately describe this response, partly to retain mathematical tractability and partly for the lack of biologically pleasing functions representing the model/environment interface. We address these problems of tractability and plausibility in community/environment models by incorporating the Boltzmann factor (temperature dependence) in a bioenergetic consumer-resource framework. Our analysis leads to three predictions for the response of consumer-resource systems to increasing mean temperature (warming). First, mathematical extinctions do not occur with warming; however, stable systems may transition into an unstable (cycling) state. Second, there is a decrease in the biomass density of resources with warming. The biomass density of consumers may increase or decrease depending on their proximity to the feasibility (extinction) boundary. Third, consumer biomass density is more sensitive to warming than resource biomass density (with some exceptions). These predictions are in line with many current observations and experiments. The model presented and analyzed here provides an advancement in the testing framework for global change scenarios and hypotheses of latitudinal and elevational species distributions.     

Low-dimensional trade-offs fail to explain richness and structure in species-rich plant communities

Mathematical models and ecological theory suggest that low-dimensional life history trade-offs (i.e. negative correlation between two life history traits such as competition vs. colonisation) may potentially explain the maintenance of species diversity and community structure. In the absence of trade-offs, we would expect communities to be dominated by ‘super-types’ characterised by mainly positive trait expressions. However, it has proven difficult to find strong empirical evidence for such trade-offs in species-rich communities. We developed a spatially explicit, rule-based and individual-based stochastic model to explore the importance of low-dimensional trade-offs. This model simulates the community dynamics of 288 virtual plant functional types (PFTs), each of which is described by seven life history traits. We consider trait combinations that fit into the trade-off concept, as well as super-types with little or no energy constraints or resource limitations, and weak PFTs, which do not exploit resources efficiently. The model is parameterised using data from a fire-prone, species-rich Mediterranean-type shrubland in southwestern Australia. We performed an exclusion experiment, where we sequentially removed the strongest PFT in the simulation and studied the remaining communities. We analysed the impact of traits on performance of PFTs in the exclusion experiment with standard and boosted regression trees. Regression tree analysis of the simulation results showed that the trade-off concept is necessary for PFT viability in the case of weak trait expression combinations such as low seed production or small seeds. However, species richness and diversity can be high despite the presence of super-types. Furthermore, the exclusion of super-types does not necessarily lead to a large increase in PFT richness and diversity. We conclude that low-dimensional trade-offs do not provide explanations for multi-species co-existence contrary to the prediction of many conceptual models.     

Zero sum, the niche, and metacommunities: long-term dynamics of community assembly

Recent models of community assembly, structure, and dynamics have incorporated, to varying degrees, three mechanistic processes: resource limitation and interspecific competition, niche requirements of species, and exchanges between a local community and a regional species pool. Synthesizing 30 years of data from an intensively studied desert rodent community, we show that all of these processes, separately and in combination, have influenced the structural organization of this community and affected its dynamical response to both natural environmental changes and experimental perturbations. In addition, our analyses suggest that zero-sum constraints, niche differences, and metacommunity processes are inextricably linked in the ways that they affect the structure and dynamics of this system. Explicit consideration of the interaction of these processes should yield a deeper understanding of the assembly and dynamics of other ecological communities. This synthesis highlights the role that long-term data, especially when coupled with experimental manipulations, can play in assessing the fundamental processes that govern the structure and function of ecological communities.     

Linking traits to species diversity and community structure in phytoplankton

In addition to answering Hutchinson’s question “Why are there so many species?”, we need to understand why certain species are found only under certain environmental conditions and not others. Trait-based approaches are being increasingly used in ecology to do just that: explain and predict species distributions along environmental gradients. These approaches can be successful in understanding the diversity and community structure of phytoplankton. Among major traits shaping phytoplankton distributions are resource utilization traits, morphological traits (with size being probably the most influential), grazer resistance traits, and temperature responses. We review these trait-based approaches and give examples of how trait data can explain species distributions in both freshwater and marine systems. We also outline new directions in trait-based approaches applied to phytoplankton such as looking simultaneously at trait and phylogenetic structure of phytoplankton communities and using adaptive dynamics models to predict trait evolution.     

Evolutionarily stable consumer home range size in relation to resource demography and consumer spatial organization

There is a large variation in home range size within species, yet few models relate that variation to demographic and life-history traits. We derive an approximate deterministic population dynamics model keeping track of spatial structure, via spatial moment equations, from an individual-based spatial consumer-resource model; where space-use of consumers resembles that of central place foragers. Using invasion analyses, we investigate how the evolutionarily stable home range size of the consumer depends on a number of ecological and behavioral traits of both the resource and the consumer. We show that any trait variation leading to a decreased overall resource production or an increased spatial segregation between consumer and resource acts to increase consumer home range size. In this way, we extend theoretical predictions on optimal territory size to a larger range of ecological scenarios where home ranges overlap and population dynamics feedbacks are possible. Consideration of spatial traits such as dispersal distances also generates new results: (1) consumer home range size decreases with increased resource dispersal distance, and (2) when consumer agonistic behavior is weak, more philopatric consumers have larger home ranges. Finally, our results emphasize the role of the spatial correlation between consumer and resource distributions in determining home range size, and suggest resource dispersion is less important.     

Trait plasticity in species interactions: a driving force of community dynamics

Evolutionary community ecology is an emerging field of study that includes evolutionary principles such as individual trait variation and plasticity of traits to provide a more mechanistic insight as to how species diversity is maintained and community processes are shaped across time and space. In this review we explore phenotypic plasticity in functional traits and its consequences at the community level. We argue that resource requirement and resource uptake are plastic traits that can alter fundamental and realised niches of species in the community if environmental conditions change. We conceptually add to niche models by including phenotypic plasticity in traits involved in resource allocation under stress. Two qualitative predictions that we derive are: (1) plasticity in resource requirement induced by availability of resources enlarges the fundamental niche of species and causes a reduction of vacant niches for other species and (2) plasticity in the proportional resource uptake results in expansion of the realized niche, causing a reduction in the possibility for coexistence with other species. We illustrate these predictions with data on the competitive impact of invasive species. Furthermore, we review the quickly increasing number of empirical studies on evolutionary community ecology and demonstrate the impact of phenotypic plasticity on community composition. Among others, we give examples that show that differences in the level of phenotypic plasticity can disrupt species interactions when environmental conditions change, due to effects on realized niches. Finally, we indicate several promising directions for future phenotypic plasticity research in a community context. We need an integrative, trait-based approach that has its roots in community and evolutionary ecology in order to face fast changing environmental conditions such as global warming and urbanization that pose ecological as well as evolutionary challenges.     

Effects of multiple timescales of resource supply on the maintenance of species and functional diversity

It is well known that variable resource supply can allow competitors to coexist on a single limiting resource, and this is one mechanism that may explain the maintenance of diversity in paradoxically speciose communities. Ecosystems experience fluctuations in resource supply on a range of timescales, but we have a poor understanding of how multiple frequencies of resource supply affect the maintenance of diversity and community structure. Here we explore this question using a model of phytoplankton competition for a limiting nutrient, parameterized using empirical tradeoffs between rapid growth, nutrient storage capacity and nutrient uptake affinity. Compared to a single frequency of nutrient supply, we find that multiple frequencies of nutrient supply increase functional diversity, by permitting the coexistence of strategies adapted to different frequencies of supply. Species richness is also promoted by multiple modes of nutrient supply, but not as consistently as functional diversity. Although this model is parameterized for phytoplankton, the fundamental dynamics and tradeoffs likely occur in a variety of ecosystems. Our results suggest that the spectrum of temporal variation driving communities should be further investigated in the context of the maintenance of diversity and the functional composition of communities under different environmental regimes.     

Adaptive plasticity in ontogenetic niche shifts stabilizes consumer-resource dynamics

Ontogenetic niche shifts, changes in the diet or habitats of organisms during their ontogeny, are widespread among various animal taxa. Ontogenetic niche shifts introduce stage structure in a population with different stages interacting with different communities and can substantially affect their dynamics. In this article, I use mathematical models to test the hypothesis that adaptive plasticity in the timing of ontogenetic niche shifts has a stabilizing effect on consumer-resource dynamics. Adaptive plasticity allows consumers in one ontogenetic niche to perform an early shift to the next ontogenetic niche if the resource density of the first niche is low. The early shift will reduce predation by the consumer on the scarce resource. On the other hand, adaptive plasticity allows consumers to delay their shift to the next niche if the resource density of the first niche is high. The delayed shift will increase the predation on the abundant resource. As a result, the scarce resource will tend to increase, and the abundant resource will tend to decrease. This causes density-dependent negative feedback in the resource dynamics, which stabilizes the consumer-resource dynamics. To test this hypothesis, I compare three consumer-resource models differing in terms of mechanisms controlling the timing of the ontogenetic niche shift: the fixed-age model assumes that the age at which the ontogenetic niche shift occurs is fixed; the fixed-size model assumes that the size at the shift is fixed; and the adaptive plasticity model assumes that the timing of the shift is such that the individual fitness of the consumer is maximized. I show that only the adaptive plasticity model has a locally stable equilibrium and that the stabilizing effect is due to the density-dependent negative feedback in the resource dynamics. I discuss the ontogenetic niche shifts of lake fish in light of the obtained result.     

Disentangling niche and neutral influences on community assembly: assessing the performance of community phylogenetic structure tests

Patterns of phylogenetic relatedness within communities have been widely used to infer the importance of different ecological and evolutionary processes during community assembly, but little is known about the relative ability of community phylogenetics methods and null models to detect the signature of processes such as dispersal, competition and filtering under different models of trait evolution. Using a metacommunity simulation incorporating quantitative models of trait evolution and community assembly, I assessed the performance of different tests that have been used to measure community phylogenetic structure. All tests were sensitive to the relative phylogenetic signal in species metacommunity abundances and traits; methods that were most sensitive to the effects of niche-based processes on community structure were also more likely to find non-random patterns of community phylogenetic structure under dispersal assembly. When used with a null model that maintained species occurrence frequency in random communities, several metrics could detect niche-based assembly when there was strong phylogenetic signal in species traits, when multiple traits were involved in community assembly, and in the presence of environmental heterogeneity. Interpretations of the causes of community phylogenetic structure should be modified to account for the influence of dispersal.