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Grazers and Diggers: Exploitation Competition and Coexistence among Foragers with Different Feeding Strategies on a Single Resource.

A mathematical model is presented that describes a system where two consumer species compete exploitatively for a single renewable resource. The resource is distributed in a patchy but homogeneous environment; that is, all patches are intrinsically identical. The two consumer species are referred to as diggers and grazers, where diggers deplete the resource within a patch to lower densities than grazers. We show that the two distinct feeding strategies can produce a heterogeneous resource distribution that enables their coexistence. Coexistence requires that grazers must either move faster than diggers between patches or convert the resources to population growth much more efficiently than diggers. The model shows that the functional form of resource renewal within a patch is also important for coexistence. These results contrast with theory that considers exploitation competition for a single resource when the resource is assumed to be well mixed throughout the system.

coexistence↗

Exploitative competition in the chemostat for two perfectly substitutable resources.

After formulating a general model involving two populations of microorganisms competing for two nonreproducing, growth-limiting resources in a chemostat, we focus on perfectly substitutable resources. León and Tumpson considered a model of perfectly substitutable resources in which the amount of each resource consumed is assumed to be independent of the concentration of the other resource. We extend their analysis and then consider a new model involving a class of response functions that takes into consideration the effects that the concentration of each resource has on the amount of the other resource consumed. This new model includes, as a special case, the model studied by Waltman, Hubbell, and Hsu in which Michaelis-Menten functional response for a single resource is generalized to two perfectly substitutable resources. Analytical methods are used to obtain information about the qualitative behavior of the models. The range of possible dynamics of model I of León and Tumpson and our new model is then compared. One surprising difference is that our model predicts that for certain parameter ranges it is possible that one of the species is unable to survive in the absence of a competitor even though there is a locally asymptotically stable coexistence equilibrium when a competitor is present. The dynamics of these models for perfectly substitutable resources are also compared with the dynamics of the classical growth and two-species competition models as well as models involving two perfectly complementary resources.

Cell Division↗

Evolution and intraspecific exploitative competition. II. A two-locus model for additive gene effects.

A two-locus model corresponding to the model of Christiansen and Loeschcke (1980, theoret, Popul. Biol. 18, 297-313) is analysed. The two loci each have two alleles, and the loci influences a character which determines the utilization of resources in a one-dimensional continuum. The analysis of the model is supported by numerical iterations of the recurrence equations. The previous prediction of high linkage disequilibrium for small allele contributions to the character and close linkage between the loci is confirmed. For larger allele contributions results comparable to those for the symmetric viabilities model are obtained. The model degenerates when the allele contributions at the two loci are equal, i.e., in the most symmetric situation. The results are discussed as the outcome of a balance between optimizing selection and disruptive selection. For small allele contributions the results are virtually independent of which genotype is most favoured by the optimizing aspect of the selective forces.

Alleles↗

The approximately ideal, more or less free distribution.

We present the minimum set of requirements necessary and sufficient to represent the foraging behaviour of an animal, and its utilisation of food, in order to explore the emergent properties of behaviour that allow animals to reduce their hunger. We present an individual-based model of foraging that provides a simple quantification of the requirements, which is sufficiently simple to yield some analytical results. Complex interactions beyond the scope of analysis have been explored through simulating animals foraging in regenerating patchy environments. In most cases the populations pass into equilibrium distributions which appear to be stable. The equilibria always approximate closely to the ideal free distribution, although typically with a small degree of undermatching. (Undermatching is the term applied to the departure from the ideal free distribution caused by a smaller proportion of the population than expected occupying areas with a higher than average regeneration rate). The model therefore implies that the distribution, hitherto accounted for in terms of ESSs may, in fact, be simply an effect of the animal's utilization of the food it collects to reduce its hunger. The model defines a specific feeling rate, v, the rate at which an animal can feed on a unit of food. This is a function of three parameters, v1, the specific feeding rate when alone, v(infinity), the rate, possibly zero, at which it can feed in the presence of an indefinitely large number of conspecifics, and n1/2, the number of conspecifics that cause v to take the value (v1+v(infinity)/2. Exploitation competition in the absence of interference is represented by setting v1 = v(infinity). Differences in competitive ability in exploitation have been represented by simulating animals with a range of values of v1, those with the larger values, feeding more rapidly, being the more effective competitors, and those with the lower values being the less effective. Interference competition is represented by setting v1 > v(infinity) and social facilitation by v1 < v(infinity). Individual differences in the strength of interaction are represented by different values of n(1/2). In competition, the animals with the larger values of n(1/2) are the more effective competitors: in facilitation, they are the less effective facilitators. The addition of physiological and behavioural detail makes very little alteration to the emergent equilibria, always close to the ideal free distribution, almost always showing undermatching.

Animals↗

Apparent competition through facilitation between Melanoides tuberculata and Biomphalaria glabrata and the control of schistosomiasis.

Interactions between two species that result in reduced growth rates for both and extinction of one of the species are generally considered cases of asymmetric interspecific competition. Exploitative or interference competition is the usual mechanism invoked. Here we describe another mechanism producing the same result, named apparent competition through facilitation (ACF), observed between Melanoides tuberculata and Biomphalaria glabrata populations. The superior competitor actually gives some benefit to the other species, whose population becomes unstable with progressively increasing oscillations, leading to extinction. A model of ACF using difference equations suggests initial dynamics distinct from traditional interspecific competition. The dynamics of two freshwater snails in the field and in laboratory experiments suggest ACF, and these relations should be considered in studies of schistosomiasis control. ACF could occur in natural populations, but might have gone undetected because the final result is similar to traditional interspecific competition.

Animals↗

Ideal free distributions when resources undergo population dynamics.

This study examines the influence of optimal patch choice by consumers on resource population dynamics and on consumer distribution in a two patch environment. The evolutionarily stable strategy which describes animal distributions across habitat patches is called the ideal free distribution (IFD) strategy. Two mechanisms that lead to the IFD are: (1) direct consumer competition such as interference, and (2) exploitative competition for resources. This article focuses on the second mechanism by assuming that resources undergo population dynamics while consumer abundance is fixed. Two models of resource growth are considered in detail: the exponential and the logistic. The corresponding consumer IFD is derived for each of these two models, assuming that consumers behave adaptively by moving to the patch which provides them with the highest fitness. This derivation does not require that resources are at an equilibrium, and it provides, for each resource density, the corresponding distribution of consumers. The article suggests that adaptive patch choice by consumers decreases between patch heterogeneity in resource levels and weakens the apparent competition between resources. The results for a single consumer population are extended for two competing consumer populations. The corresponding IFD is computed as a function of the two consumer densities. This allows for the analytical description of isolegs which are the boundary lines, in the two consumer density phase space, separating regions where qualitatively different habitat preferences are predicted.

Animals↗

The competition diallel and the exploitation and interference components of larval competition in Drosophila melanogaster.

A logistic model of the competition diallel is presented based on two linear parameters for the exploitation component of competition, namely the acquisition rate (f) and utilization efficiency (u), and one linear parameter for the interference component of competition (i). This interference component encompasses all phenomena that are uniquely related to duocultures, such as resource partitioning, mutual stimulation, inhibition and complementation. The model uses yield-density regression coefficients (c-values), but could be adapted to suit other variates that account for both competitor density and relative frequency. In Drosophila larval competition most interference is negative and depresses the performance of duocultures with respect to monocultures, over and above that expected from shared exploitation of a common resource. Even in the closely controlled competitive conditions of these experiments this interference accounts for a considerable proportion of the total variation. The isolation of a general, and therefore predictable, interference component may prove useful in agriculture when assessing the relative importance of mixture effects to the yield potential of different crops.

Animals↗

Interspecific combative interactions between wood-decaying basidiomycetes.

Competition is the most common type of interaction occurring between wood-decaying higher fungi. Since competition for nutrients in organic resources is effectively brought about by competition for space, the common division into interference and exploitation competition is not very appropriate. Fungal competition can be divided into primary resource capture (obtaining uncolonized resources) and secondary resource capture (combat to obtain resources already colonized by other fungi). Combative mechanisms include antagonism at a distance, hyphal interference, mycoparasitism and gross mycelial contact. Interactions can result in deadlock or replacement, and a hierarchy of combative ability can be discerned amongst fungi that inhabit particular resources, but within this hierarchy there exists intransitivity, modification of outcome by other species and abiotic variables. Interactions can dramatically alter mycelial function, and have potential as biological control agents of fungal pathogens of trees and in service timber.

Journal Article↗

When trade-offs interact: balance of terror enforces dominance discovery trade-off in a local ant assemblage.

1. Trade-offs underpin local species coexistence. Trade-offs between interference and exploitative competitive ability provie a mechanism for explaining species coexistence within guilds that exploit overlapping resources. 2. Omnivorous, leaf litter ants exploit a shared food base and occur in species-rich assemblages. In these assemblages, species that excel at usurping food items from other species are poor at finding food items first. In assemblages where some members are attacked by phorid fly parasitoids, host species face an additional trade-off between defending themselves against parasitic attack and maximizing their competitive abilities. Host species thus face two trade-offs that interact via the trait-mediated indirect interaction generated by phorid defence behaviour. 3. In this study we test for the existence of these trade-offs and evaluate the predictions of a model for how they interact in an assemblage of woodland ants in which two behaviourally dominant members are attacked by phorid fly parasitoids as they attempt to harvest food resources. 4. The major findings are that unparasitized species in the assemblage follow a dominance-discovery trade-off curve. When not subject to attack by phorid flies, host species violate that trade-off by finding resources too quickly for their level of behavioural dominance. In contrast, when attacked by their phorid parasitoids, the host species dominance drops such that they fall into the assemblage trade-off. 5. These results match the predictions of the balance of terror model, which derives the optimal host response to parasitism, indicating that the host species balance the competing fitness costs of reduced competitive dominance and loss of workers to parasitism. This result supports the view that understanding the structure of ecological communities requires incorporating the indirect effects created by trait plasticity.

Animals↗

Is the shape of the density-growth relationship for stream salmonids evidence for exploitative rather than interference competition?

1. Empirical studies show that average growth of stream-dwelling salmon and trout often declines with increasing density in a characteristic concave relationship. However, the mechanisms that generate negative density-growth relationships in populations in natural streams are not certain. 2. In a recent study, Imre, Grant & Cunjak (2005; Journal of Animal Ecology, 74, 508-516) argue that density-dependent growth due to exploitative competition for prey causes the negative density-growth relationships for stream salmonids. They argue that the concave shape of empirical density-growth relationships is consistent with a simple model of exploitative competition and not consistent with interference competition for space. 3. We use a simple model to show that competition for space can yield concave density-growth relationships consistent with the empirical pattern when individuals compete for foraging sites that vary spatially in quality and lower-quality sites predominate. Thus, the predictions of the exploitative competition and spatial competition models overlap. 4. The shape of the density-growth relationship does not differentiate between candidate mechanisms underlying density-dependent growth for stream salmonids. Our results highlight the general problem with determining the mechanism driving an ecological process from patterns in observational data within the context of linking population demographics to habitat structure and animal behaviour.

Animals↗

Begging signals and biparental care: nestling choice between parental feeding locations

The evolutionary conflict over the amount of resources transferred between a parent and its offspring may be resolved by honest signalling of 'need' by offspring and parental investment in relation to signalling level. In birds, biparental care is the norm and evidence that male and female parents differ in their investment pattern in individual offspring is growing. In an experiment on great tits, Parus major, we investigated how and why parents differ in food allocation when responding to similar chick signals, which supposedly uniquely reflect the chick's nutritional condition. Nestling hunger level was manipulated by food deprivation and hand-feeding. Subsequent filming revealed that parents fed from significantly different locations on the nest and thereby forced chicks to choose between them when competing for favourable positions. Deprived nestlings approached, and fed ones retreated (or were displaced by siblings) from, positions near the female. No such behaviour was observed towards the male. Females allocated more feeds than males to the food-deprived nestlings. The results are discussed in terms of nestling competition for access to 'begging patches'. By varying their 'begging patch' value, parents may exploit competitive inter-sibling dynamics to influence the outcome of competition among chick phenotypes (e.g. 'need', size, sex). Parent birds may thereby exert considerable control over the information content of chick begging behaviour. Copyright 1998 The Association for the Study of Animal Behaviour.

Journal Article↗

Bacterial competition in activated sludge: theoretical analysis of varying solids retention times on diversity.

A mechanistic model for activated sludge sewage treatment was developed to predict exploitative competition of six aerobic heterotrophic bacterial species competing for three essential resources. The central hypothesis of the model is that in a multispecies/limiting resource system the number of coexisting bacterial species, N, exceeds the number of limiting resources, K, available for them. The explanation for this is that for certain species combinations, the dynamics of the competition process generate oscillations in the abundances of species, and these oscillations allow the coexistence of greater number of species than the number of limiting resources ( N > K). This result is a direct contradiction of an existing activated sludge steady state competition theory, "the principle of competitive exclusion," which states that the competition process proceeds to equilibrium, allowing only N </= K species to coexist. The model was used to investigate the effect of varying solids retention times on the diversity of species using the conventional, completely mixed activated sludge configuration. The results of model simulations showed that for a certain range of solids retention times (2.28-5.66 days) the competition of six species for three essential resources produces oscillations within the structure of the bacterial community allowing for the sustained growth of more than three species on three resources.

Bacterial Physiological Phenomena↗

Role of nutrient bound of prey on the dynamics of predator-mediated competitive-coexistence.

The coexistence of competitive species with a shared predator is well established. The effect of 'food-value' on predator-prey dynamics has also received much attention. However, the study of a nutrient bound of prey, specifically on predator-mediated competitive-coexistence has not received much attention. Here we study the effects of the caloric content or a nutrient bound of prey on the dynamics of competitive-coexistence with the shared predator in a specific model. We propose and analyze a mathematical model for exploitative competition of two prey species with a shared predator. The change of dynamic stability due to the variation of a nutrient bound of each prey on predator-mediated dynamics is studied through extensive numerical experiments. Our analytical and numerical results demonstrate that variation in a nutrient bound promotes the switching of dynamics and may be treated as a driving force for the dynamics of competitive-coexistence with the shared predator.

Animal Nutritional Physiological Phenomena↗

Three-way coexistence in obligate mutualist-exploiter interactions: the potential role of competition.

Many mutualisms host "exploiter" species that consume the benefits provided by one or both mutualists without reciprocating. Exploiters have been widely assumed to destabilize mutualisms, yet they are common. We develop models to explore conditions for local coexistence of obligate plant/pollinating seed parasite mutualisms and nonpollinating exploiters. As the larvae of both pollinators and (at a later time) exploiters consume seeds, we examine the importance of intraspecific and (asymmetric) interspecific competition among and between pollinators and exploiters for achieving three-way coexistence. With weak intra- and interspecific competition, exploiters can invade the stable mutualism and coexist with the mutualists (either stably or with oscillations), provided the exploiters' intrinsic birthrate (b(E)) slightly exceeds that of the pollinators. At higher b(E), all three species go locally extinct. When facing strong interspecific competition, exploiters cannot invade and coexist with the mutualists if intraspecific competition in pollinators and exploiters is weak. However, strong intraspecific competition in pollinators and exploiters facilitates exploiter invasion and coexistence and greatly expands the range of b(E) over which stable coexistence occurs. Our results suggest that mutualist/exploiter coexistence may be more easily achieved than previously thought, thus highlighting the need for a better understanding of competition among and between mutualists and exploiters.

Animals↗

Interference competition set limits to the fundamental theorem of natural selection.

The relationship between Fisher's fundamental theorem of natural selection and the ecological environment of density regulation is examined. Using a linear model, it is shown that the theorem holds when density regulation is caused by exploitative competition and that the theorem fails with interference competition. In the latter case the theorem holds only at the limit of zero population density and/or at the limit where the competitively superior individuals cannot monopolize the resource. The results are discussed in relation to population dynamics and life history evolution, where evidence suggests that the level of interference competition in natural populations is so high that the fundamental theorem does not apply.

Animals↗

Habitat structure determines competition intensity and invasion success in gecko lizards.

Species diversity is correlated with structural complexity in many animal communities; however, experimental tests of the mechanisms underlying this important relationship are rare, especially in terrestrial communities. We manipulated physical features of the habitat of gecko lizards and measured the effect on exploitation competition for insects. Increasing both the dispersion of food resources and microhabitat topography dramatically reduced interspecific competition. Adding topographic structure reduced the advantages of the larger, faster, invasive species. Interindividual spacing decreased, but intraspecific agonistic interference increased in the more territorial, resident species. Human structural alterations of the environment facilitate invasion and competitive displacement in this system. Physical microhabitat structure can potentially affect species interactions through a variety of complex mechanisms.

Animals↗

Competitive exclusion and coexistence of species with complex life cycles.

Complex life cycles are life histories in which abrupt ontogenetic transformations and niche shifts occur at the transition between stages. The effects of this niche differentiation between stages on coexistence between species are investigated using a simple discrete model of two-stage populations. The model incorporates exploitation competition for limiting resources within stages, between stages, and between species. While species with simple life cycles can never coexist at equilibrium, stable coexistence is shown to be possible between species with complex life cycles provided that (1) one species is more efficient in resource utilization at low resource abundance in the larval stage while the other is more efficient at low resource abundance in the adult stage; and (2) each species is mainly limited by that stage which is less efficient at low resource abundance. Stable coexistence is somewhat easier between a species with a simple life cycle and one with a complex life cycle. It requires that (1) the species with the simple life cycle should not be decidedly more efficient than that with the complex life cycle in utilizing the resource on which it lives; and (2) the main resource limitation for the species with a complex life cycle should occur in that stage which escapes competition with the species with a simple life cycle. Lastly, a complex life cycle can offer a decisive competitive superiority over a simple life cycle in interspecific competition, which suggests that competition can be a driving force of the evolution of complex life cycles.

Animals↗