Competition and the Effect of Spatial Resource Heterogeneity on Evolutionary Diversification

A model is presented to explore how the form of selection arising from competition for resources is affected by spatial resource heterogeneity. The model consists of a single species occupying two patches connected by migration, where the two patches can differ in the type of resources that they contain. The main goal is to determine the conditions under which competition for resources results in disruptive selection (i.e., selection favoring a polymorphism) since it is this form of selection that will give rise to the evolutionary diversification of resource exploitation strategies. In particular, comparing the conditions giving rise to disruptive selection when the two patches are identical to the conditions when they contain different resources reveals the effect of spatial resource heterogeneity. Results show that when the patches are identical, the conditions giving rise to disruptive selection are identical to those that give rise to character displacement in previous models. When the patches are different, the conditions giving rise to disruptive selection can be either more or less stringent depending upon demographic parameters such as the intrinsic rate of increase and the migration rate. Surprisingly, spatial resource heterogeneity can actually make forms of evolutionary diversification such as character displacement less likely. It is also found that results are dependent on how the resource exploitation strategies and the spatial resource heterogeneity affect the population dynamics. One robust conclusion however, is that spatial resource heterogeneity always has a disruptive effect when the migration rate between patches is low.     

Spatial structure of ecological opportunity drives adaptation in a bacterium

Abundant ecological opportunity is thought to drive adaptation and diversification. The presence of multiple opportunities leads to divergent selection, which can slow adaptation when niche-specific beneficial mutations have antagonistically pleiotropic effects. Alternately, competition for multiple opportunities can generate divergent selection, which leads to high rates of adaptive differentiation. Which outcome occurs may depend on the spatial structure of those ecological opportunities. In a mixture of resources, competition for multiple opportunities can drive divergent selection; however, if each resource is available in a spatially distinct patch, simultaneous competition for multiple opportunities cannot occur. We report the effects of the extent and spatial structure of ecological opportunity on the evolutionary dynamics of populations of Pseudomonas fluorescens over 1,000 generations. We varied the extent of ecological opportunity by varying the number of sugar resources (mannose, glucose, and xylose), and we varied spatial structure by providing resources in either mixtures or spatially distinct patches. We saw that a particularly novel resource (xylose) drove the rate of adaptation when provided in a mixture but had no effect on diversity. Instead, we saw the evolution of a single adaptive strategy that differed with respect to phenotype and degree of specialization, depending on both the extent and the spatial structure of ecological opportunity.     

Interspecific Competition Models Derived from Competition Among Individuals

This paper demonstrates how discrete-time models describing population dynamics of two competing species can be derived in a bottom-up manner by considering competition for resources among individuals and the spatial distribution of individuals. The competition type of each species is assumed to be either scramble, contest, or an intermediate between them. Individuals of two species are distributed over resource sites or patches following one of three distribution functions. According to the combination of competition types of the two species and the distribution of individuals, various interspecific competition models are derived. Furthermore, a general interspecific competition model that includes various competition models as special cases is derived for each distribution of individuals. Finally, this paper examines dynamics of some of the derived competition models and shows that the likelihood of coexistence of the two species varies greatly, depending on the type of spatial distribution of individuals.     

Nutrient addition does not increase the benefits of clonal integration in an invasive plant spreading from open patches into plant communities

• One benefit of clonal integration is that resource translocation between connected ramets enhances the growth of the ramets grown under stressful conditions, but whether such resource translocation reduces the performance of the ramets grown under favourable conditions has not produced consistent results. In this study, we tested the hypothesis that resource translocation to recipient ramets may reduce the performance of donor ramets when resources are limiting but not when resources are abundant.
• We grew Mikania micrantha stolon fragments (each consisting of two ramets, either connected or not connected) under spatially heterogeneous competition conditions such that the developmentally younger, distal ramets were grown in competition with a plant community and the developmentally older, proximal ramets were grown without competition. For half of the stolon fragments, slow‐release fertiliser pellets were applied to both the distal and proximal ramets.
• Under both the low and increased soil nutrient conditions, the biomass, leaf number and stolon length of the distal ramets were higher, and those of the proximal ramets were lower when the stolon internode was intact than when it was severed. For the whole clone, the biomass, leaf number and stolon length did not differ between the two connection treatments. Connection did not change the biomass of the plant communities competing with distal ramets of M. micrantha.
• Although clonal integration may promote the invasion of M. micrantha into plant communities, resource translocation to recipient ramets of M. micrantha will induce a cost to the donor ramets, even when resources are relatively abundant.

     

Departure from naturalized to invasive stage: a disturbance-induced mechanism and associated interacting factors

Aims Within a habitat of multiple plant species, increased resource availabilities and altered species abundances following disturbances create opportunities for exotic species to successfully establish and subsequently naturalize into its non-native environment. Such post-disturbance changes in abiotic and biotic environments may also promote a naturalized exotic species (or invading species) to become invasive through rapid colonization of the habitat sites by reducing the extent and size of resident plant species. By combining species life history traits with that of the disturbance-induced changes in habitat characteristics, we aimed to determine those interacting factors and associated mechanism allowing an exotic invasion to start off.Methods We used a modified version of the classic competition–colonization (CC) model which was formulated first by Hastings (1980) and studied later by Tilman (1994) to explain spatial coexistence of multiple species. Within this model framework, recruitment-limited spatial competition has explicitly been linked with interspecific resource competition without altering the basic assumptions and structure of the original CC model.Important findings The model results showed that at a constant rate of resource supply, invading species can stably coexist with native species via trade-offs between species competitive ability and colonizing ability. On the other hand, the model predicted that with a fluctuating resource condition, invading species can successfully invade a habitat following continuous reductions in the size and extent of native species. Whether or not invading species holds competitive superiority over the native species for limiting resource, we showed that there exists a range of variation in available resource that allows an exotic invasion to start off in post-disturbance habitat. The associated disturbance-induced mechanism promoting invading species to become invasive has been identified. It states that occurrences of disturbances such as fire or clear-cutting influence variation in resource availability, and in addition open up many vacant microsites; given these disturbance-induced changes, invading species with a higher rate of propagule production and with a higher survival rate of adults particularly in low-resource condition recruits microsites at faster rate relative to native competitor species, and with a given range of variation in resource availabilities, it maintains continued expansions following reductions in size and extent of native species. Moreover, we identified those interacting factors and their specific roles that drive this mechanism. These factors include propagule supply, variable resource level and vacant microsite availability. Increased availability of vacant microsites following disturbances creates an opportunity for rapid colonization. Given this opportunity, higher number of propagules supplied by the invading species enhances the rate of colonization success, whereas the resource variation within a range of given thresholds maintains enhanced colonization rate of the invading species while it depresses native competitor species. Owing to the each factor's invasion regulatory ability, controlling one or all of them may have strong negative impact on the occurrence of exotic invasion.     

Two-Species Migration and Clustering in Two-Dimensional Domains

We extend two-species models of individual aggregation or clustering to two-dimensional spatial domains, allowing for more realistic movement of the populations compared with one spatial dimension. We assume that the domain is bounded and that there is no flux into or out of the domain. The motion of the species is along fitness gradients which allow the species to seek out a resource. In the case of competition, species which exploit the resource alone will disperse while avoiding one another. In the case where one of the species is a predator or generalist predator which exploits the other species, that species will tend to move toward the prey species, while the prey will tend to avoid the predator. We focus on three primary types of interspecies interactions: competition, generalist predator–prey, and predator–prey. We discuss the existence and stability of uniform steady states. While transient behaviors including clustering and colony formation occur, our stability results and numerical evidence lead us to believe that the long-time behavior of these models is dominated by spatially homogeneous steady states when the spatial domain is convex. Motivated by this, we investigate heterogeneous resources and hazards and demonstrate how the advective dispersal of species in these environments leads to asymptotic steady states that retain spatial aggregation or clustering in regions of resource abundance and away from hazards or regions or resource scarcity.     

Determinants of resource use in lizard assemblages from the semiarid Caatinga,Brazil

Nonsessile animals could partition the use of resources in different axes, reducing the effects of competition and allowing coexistence. Here, we investigated the spatial and trophic niche dimensions in four lizard assemblages in the Neotropical semiarid Caatinga to investigate the determinants of resource use and the extent to which lizards partition their niches. We sampled each lizard assemblage once, for 10 days, in the dry season of 2017 and 2018. In two lizard assemblages, we detected nonrandom niche overlap patterns that were higher or lower than expected by chance. The high niche overlap patterns suggest that either there is intense current competition for available microhabitats or an abundance of microhabitats. The lower niche overlap may be influenced by the presence of species adapted to sandy habitats (psammophilous), suggesting that spatial partitioning detected has historical basis, which is supported by the pPCA results and by the lack of patterns in the realized niche distribution of species across niche space. We detected trophic niche partitioning in three lizard assemblages. In one assemblage, we discovered random spatial and trophic niche overlap patterns, revealing that competition is not a determining factor in the structure of that assemblage. In fact, phylogenetic effects were predominantly the main determinants of resource use in the four studied lizard assemblages. Arid and semiarid habitats cover about one third of land surface of the world. Comparisons between our findings and those from other regions of the world may aid identify general trends in the lizard ecology of dry environments.     

Advantages and disadvantages of interference‐competitive ability and resource‐use efficiency when invading established communities

Invaders into established communities must overcome low resource availability. To establish, invaders must either appropriate resources from existing individuals through interference competition or efficiently use the small amount of resource that remains. Although both strategies may be important, they are rarely considered together and, in particular, resource‐use efficiency is often ignored in systems dominated by interference competition. To identify the traits that confer invasion success, we experimentally invaded resource patches in established communities with multiple species from two functional groups that differ in interference competitive ability and resource‐use efficiency. In contrast to previous assessments, we show that resource‐use efficiency can facilitate invasion in systems dominated by interference competition. Furthermore, large resource requirements can be a liability when establishing because interference competition is inherently costly and so cannot fully compensate for limitations in the primary resource. However, we also show that there is a tradeoff in performance among functional groups between small and large resource gaps. Our results suggest we modify the way we view and manage species invasion in systems dominated by interference competition.     

Effects of spatial aggregation on competition,complementarity and resource use

Abstract The spatial distributions of most species are aggregated to varying degrees. A limited number of studies have examined the effects of spatial aggregation on interspecific and intraspecific interactions, generally finding that spatial aggregation can enhance coexistence between species by reducing the capacity for interspecific competition. Less well studied are the effects of spatial aggregation on complementarity (i.e. differences in resource use strategies) and resource use. Our primary hypothesis was that spatial aggregation reduces the complementarity between species owing to: (i) less interspecific interactions as a result of spatial separation; and (ii) less differences between species as a result of phenotypic plasticity. We further postulate that these negative effects of spatial aggregation on complementarity will reduce resource use by the community. Here we test these hypotheses in a pot experiment in which we applied three levels of spatial aggregation to three sets of two‐species mixtures of herbaceous perennial plant species from native grasslands of south‐eastern Australia. Both root and shoot biomass were significantly affected by spatial aggregation, although the nature of these affects depended upon the species involved, and the relative strengths of interspecific versus intraspecific competition. Complementarity between species in the distribution of their green leaves decreased significantly as spatial aggregation increased for one of the species mixtures, providing some evidence in support of our hypothesis that aggregation reduces complementarity through phenotypic plasticity. Spatial aggregation also altered light interception and use of soil moisture resources, although these effects were dependent on the species involved. We suggest that clear effects of spatial aggregation on complementarity and resource use may be obscured by the idiosyncratic way in which neighbour identity influences plant growth and hence plant size, limiting the ability to generalize, at the community level, any underlying effects of spatial pattern on ecological process.     

Trade-off between reciprocal mutualists: local resource availability-oriented interaction in fig/fig wasp mutualism

1. The mechanisms that prevent competition (conflict) between the recipient and co-operative actor in co-operative systems remain one of the greatest problems for evolutionary biology. Previous hypotheses suggest that self-restraint, dispersal or spatial constraints can prevent direct competition for local resources or any other common resources, thereby maintaining stable co-operation interactions. In this study, we use the obligate fig-fig-wasp mutualism to examine whether the above mechanisms can maintain stable co-operation sufficiently between figs and fig wasps. 2. Our data on obligate co-operation between figs (Ficus racemosa Linn.) and fig wasps (Ceratoslen fusciceps Mayr) show that the number of viable seeds of figs is positively correlated with the number of pollinator offspring when the number of vacant female flowers is high while the foundress number is low (two foundresses). Meanwhile, they are negatively correlated when the number of vacant female flowers is low and the number of foundresses is increased manually (eight foundresses). The correlation coefficient between viable seeds and wasp offspring (galls) depends on vacant female flower availability. 3. Our data suggest that the interaction between figs and fig wasps is conditional, and that they co-operate when local resource availability is plentiful but are in conflict when local resource availability is limited. The self-restraint, dispersal and spatial heterogeneity previously hypothesized in maintaining stable co-operation cannot sufficiently prevent the symbionts from utilizing more local resources at the expense of the recipients. The conflict, which can disrupt the co-operation interaction, exists after the local resource is saturated with symbionts. The repression of symbiont increase, therefore repressing the utilization of local resources in the conflict period, is crucial in the maintenance and evolution of co-operation.     

Intraspecific variation in male mating strategies in an African ground squirrel (Xerus inauris)

Male mating strategies respond to female availability such that variation in resources that affect spatial distribution can also alter cost–benefit tradeoffs within a population. In arid‐adapted species, rainfall alters reproduction, behavior, morphology, and population density such that populations differing in resource availability may also differ in successful reproductive strategies. Here, we compare two populations of Cape ground squirrels (Xerus inauris), a sub‐Saharan species with year‐round breeding and intense mating competition. Unlike most mammals where males resort to aggressive interactions over females, male X. inauris are tolerant of one another, relying instead on other nonaggressive pre‐ and postcopulatory strategies to determine reproductive success. Our findings suggest that differences in resource availability affect female distribution, which ultimately leads to intraspecific variation in male reproductive tactics and sexual morphology. Sperm competition, assessed by reproductive morphometrics, was more pronounced in our high resource site where females were distributed evenly across the landscape, whereas dominance seemed to be an important determinant of success in our low resource site where females were more aggregated. Both sites had similar mating intensities, and most males did not sire any offspring. However, our low resource site had a higher variance in fertilization success with fewer males siring multiple offspring compared with our high resource site where more individuals were successful. Our results lend support to resource models where variations in female spatial distribution attributed to environmental resources ultimately impact male reproductive behaviors and morphology.     

Putting competition strategies into ideal free distribution models: habitat selection as a tug of war

When resources are patchily distributed in an environment, behavioral ecologists frequently turn to ideal free distribution (IFD) models to predict the spatial distribution of organisms. In these models, predictions about distributions depend upon two key factors: the quality of habitat patches and the nature of competition between consumers. Surprisingly, however, no IFD models have explored the possibility that consumers modulate their competitive efforts in an evolutionarily stable manner. Instead, previous models assume that resource acquisition ability and competition are fixed within species or within phenotypes. We explored the consequences of adaptive modulation of competitive effort by incorporating tug-of-war theory into payoff equations from the two main classes of IFD models (continuous input (CI) and interference). In the models we develop, individuals can increase their share of the resources available in a patch, but do so at the costs of increased resource expenditures and increased negative interactions with conspecifics. We show how such models can provide new hypotheses to explain what are thought to be deviations from IFDs (e.g., the frequent observation of fewer animals than predicted in "good" patches of habitat). We also detail straightforward predictions made uniquely by the models we develop, and we outline experimental tests that will distinguish among alternatives.     

Optimal cell size for resource uptake in fluids: a new facet of resource competition

Planktonic microorganisms are affected by various size-dependent processes both from the bottom up and from the top down. We developed a simple resource-consumer model to explore how size-dependent resource uptake and resource loss influence the growth of, and competition between, planktonic microorganisms. We considered three steps of resource uptake: diffusive transport of resource molecules, uptake by membrane transporters, and cellular enzymatic catalysis, and we investigated optimal cell size when one, two, or three of those steps limit resource uptake. Optimal cell size depends negatively on the size of resource molecules when resource uptake is limited by diffusive transport and membrane uptake. When competing for two resources of different molecular sizes, two different-sized consumers can coexist if the inputs of resources and sizes of consumers are correctly chosen. The model suggests that mixtures of various-sized resources can promote coexistence and size diversity of microorganisms even if the availability of one element, such as carbon, nitrogen, or phosphorus, limits the whole community. Model predictions include that bacteria grown on maltose or polysaccharides should be smaller compared with those grown on glucose under carbon limitation. Our results suggest that size of resource molecules can be an important factor in microbial resource competition in aquatic environments.     

Evolution of resource specialisation in competitive metacommunities

Spatial environmental heterogeneity coupled with dispersal can promote ecological persistence of diverse metacommunities. Does this premise hold when metacommunities evolve? Using a two‐resource competition model, we studied the evolution of resource‐uptake specialisation as a function of resource type (substitutable to essential) and shape of the trade‐off between resource uptake affinities (generalist‐ to specialist‐favouring). In spatially homogeneous environments, evolutionarily stable coexistence of consumers is only possible for sufficiently substitutable resources and specialist‐favouring trade‐offs. Remarkably, these same conditions yield comparatively low diversity in heterogeneous environments, because they promote sympatric evolution of two opposite resource specialists that, together, monopolise the two resources everywhere. Consumer diversity is instead maximised for intermediate trade‐offs and clearly substitutable or clearly essential resources, where evolved metacommunities are characterised by contrasting selection regimes. Taken together, our results present new insights into resource‐competition‐mediated evolutionarily stable diversity in homogeneous and heterogeneous environments, which should be applicable to a wide range of systems.     

Multiple resource limitation: nonequilibrium coexistence of species in a competition model using a synthesizing unit

During the last two decades, the simple view of resource limitation by a single resource has been changed due to the realization that co-limitation by multiple resources is often an important determinant of species growth. Hence, the multiple resource limitation hypothesis needs to be taken into account, when communities of species competing for resources are considered. We present a multiple species–multiple resource competition model which is based on the concept of synthesizing unit to formulate the growth rates of species competing for interactive essential resources. Using this model, we demonstrate that a more mechanistic explanation of interactive effects of co-limitation may lead to the known complex dynamics including nonequilibrium states as oscillations and chaos. We compare our findings with earlier investigations on biological mechanisms that can predict the outcome of multispecies competition. Moreover, we show that this model yields a periodic state where more species than limiting complementary resources can coexist (supersaturation) in a homogeneous environment. We identify two novel mechanisms, how such a state can emerge: a transcritical bifurcation of a limit cycle and a transition from a heteroclinic cycle. Furthermore, we demonstrate the robustness of the phenomenon of supersaturation when the environmental conditions are varied.     

How Gaussian competition leads to lumpy or uniform species distributions

A central model in theoretical ecology considers the competition of a range of species for a broad spectrum of resources. Recent studies have shown that essentially two different outcomes are possible. Either the species surviving competition are more or less uniformly distributed over the resource spectrum, or their distribution is “lumped” (or “clumped”), consisting of clusters of species with similar resource use that are separated by gaps in resource space. Which of these outcomes will occur crucially depends on the competition kernel, which reflects the shape of the resource utilization pattern of the competing species. Most models considered in the literature assume a Gaussian competition kernel. This is unfortunate, since predictions based on such a Gaussian assumption are not robust. In fact, Gaussian kernels are a border case scenario, and slight deviations from this function can lead to either uniform or lumped species distributions. Here, we illustrate the non-robustness of the Gaussian assumption by simulating different implementations of the standard competition model with constant carrying capacity. In this scenario, lumped species distributions can come about by secondary ecological or evolutionary mechanisms or by details of the numerical implementation of the model. We analyze the origin of this sensitivity and discuss it in the context of recent applications of the model.     

Direct competition results from strong competition for limited resource

We study a model of competition for resource through a chemostat-type model where species consume the common resource that is constantly supplied. We assume that the species and resources are characterized by a continuous trait. As already proved, this model, although more complicated than the usual Lotka–Volterra direct competition model, describes competitive interactions leading to concentrated distributions of species in continuous trait space. Here we assume a very fast dynamics for the supply of the resource and a fast dynamics for death and uptake rates. In this regime we show that factors that are independent of the resource competition become as important as the competition efficiency and that the direct competition model is a good approximation of the chemostat. Assuming these two timescales allows us to establish a mathematically rigorous proof showing that our resource-competition model with continuous traits converges to a direct competition model. We also show that the two timescales assumption is required to mathematically justify the corresponding classic result on a model consisting of only finite number of species and resources (MacArthur in, Theor Popul Biol 1:1–11, 1970). This is performed through asymptotic analysis, introducing different scales for the resource renewal rate and the uptake rate. The mathematical difficulty relies in a possible initial layer for the resource dynamics. The chemostat model comes with a global convex Lyapunov functional. We show that the particular form of the competition kernel derived from the uptake kernel, satisfies a positivity property which is known to be necessary for the direct competition model to enjoy the related Lyapunov functional.     

Starve a competitor: evolution of luxury consumption as a competitive strategy

Organisms are often observed to acquire an excess of non-limiting resources, a process known as luxury consumption. Luxury consumption has been largely treated as a bet hedging strategy for temporal variation in resource supply, but may also function as a competitive strategy. We incorporate luxury resource consumption into a derivation of the classic resource ratio model for competition between terrestrial plant, and explore its consequences for population dynamics and competition. We show that luxury consumption reduces the potential for coexistence between two species competing for two resources. Furthermore, we demonstrate that luxury consumption can be selected for because of the competitive advantage that luxury consumers gain. Luxury consumption evolves when competition for resources is local rather than global, there is potential for coexistence between the two species and the competitive environment remains stable over a sufficient period of time to allow selection to act. The evolutionary outcome can be either extinction of one of the competing species or coexistence of the two species with maximum luxury consumption. The potential for selection to favor luxury consumption is well predicted by the competitive outcome between individuals of the two species with and without luxury consumption.     

High competition with low similarity and low competition with high similarity: exploitative and apparent competition in consumer-resource systems

This article investigates the relationship between the similarity of resource capture abilities and the amount of competition between two consumer species that exploit common resources. Most of the analysis is based on a consumer-resource model introduced by Robert MacArthur. Contrary to many statements in the literature and in textbooks, measures of competition may decrease as similarity increases and may be greatest when similarity of the two species' sets of resource capture rates is very low. High competition with low similarity may occur whether competition is measured by a competition coefficient near equilibrium or is measured by the proportional increase in a species' population density when its competitor is removed. However, these two measures may differ considerably and may change in opposite directions with a given change in similarity. The general conditions required for such counterintuitive relationships between similarity and competition are that the consumer species have relatively low resource requirements for successful reproduction and that the resources be self-reproducing. These same conditions also frequently lead to exclusion of one or more resources via apparent competition, and this is always true of MacArthur's model. A variety of other models of competition are analyzed, and circumstances most likely to produce large competitive effects with little overlap are identified.