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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↗

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↗

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↗

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↗

Parasite-mediated and direct competition in a two-host shared macroparasite system.

This paper investigates the local dynamical behaviour of a deterministic model describing two host species experiencing three forms of competition: direct competition, apparent competition mediated by macroparasites, and intra-specific (density-dependent) competition. The problem of algebraic intractability is sidestepped by adopting a geometric approach, in which an array of maps is constructed in parameter space, each structured by bifurcation surfaces which mark qualitative changes in system behaviour. The maps provide both a succinct and a comprehensive overview of the stability and feasibility structure of the system equilibria, from which can be deduced the possible modes of local dynamical behaviour. A detailed examination of these maps shows that (i) the system is highly sensitive to the effect of infection on fecundity with synchronous sustained cycles readily generated by Hopf bifurcations; (ii) for a broad range of parameter values, pertinent to actual biological systems, apparent competition mediated by macroparasites is sufficient, on its own, to explain host exclusion; (iii) direct competition reinforces parasite-mediated competition to expand the host exclusion region; and (iv) the condition for host exclusion can be expressed simply in a form which holds for both micro- and macroparasite models and which involves just two key indices, measuring tolerance to the infection and the strength of direct competition. The techniques used in this paper are not restricted to the analysis of host-parasite systems but can be applied to a wide range of nonlinear population models. They are therefore as relevant to the analysis of such general issues as exploitative competition and trophic interactions as they are to specific epidemiological problems.

Animals↗

Modelling the immune response to malaria with ecological concepts: short-term behaviour against long-term equilibrium.

A model for the human immune response to the malaria parasite Plasmodium falciparum is used to analyse the dynamics of an infection within an individual patient. Previous models either looked at competition between two parasite genotypes or at one parasite clone and the immune response to it. This model describes the course of an infection caused by the blood stages of two parasite genotypes differing in reproductive rate and in the immune response they elicit. The interactions between the genotypes can be interpreted as exploitative competition for red blood cells. Interactions between omnipotent immune cells and parasites resemble a predator-prey relation. In analysing these kinds of models, classical theoretical ecology usually deals with long-term behaviours, i.e. looks for equilibria and conditions for coexistence. However, especially in endemic regions with ongoing transmission, an equilibrium state of infections is unlikely. When reinfections with another parasite genotype were considered, the short-term dynamics of the infection changed dramatically, depending on which genotype was first, when the second one appeared, and what kind of immune response was elicited. If the slow development of immunity to malaria really is due to its genotype specificity, the effects of superinfections will be of great importance.

Animals↗

Co-existence of congeneric species of acanthocephala: Acanthocephalus lucii and A. anguillae in eels Anguilla anguilla in Ireland.

A population of eels Anguilla anguilla from Lough Derg, R. Shannon, Ireland, harbouring infections of both Acanthocephalus lucii and A. anguillae was studied over three years. Both parasite species had the same intermediate host and eels appeared to be the only definitive host for A. anguillae. Throughout the whole period, A. lucii was the dominant parasite, was over-dispersed throughout the eel population and most frequently occurred as a single species infection. A. anguillae was far less common, its dispersion was close to random at most times and it almost invariably occurred as a mixed species infection. The proportions of the two species remained fairly constant over the period. Despite some indication of site selection in the intestine, the distribution of both species overlapped considerably and there was no evidence of competitive displacement of one species by the other or of resource partitioning in space. The life-histories of both species were similar: they infected eels, bred and were lost from fish at the same time of year and there was no indication of resource partitioning in time. Congeneric species of acanthocephalans can thus co-exist in apparently stable equilibrium in fish as predicted and without any evidence of interactions, but it is still considered that exploitation competition between the species may be occurring in eels.

Acanthocephala↗

The population dynamics of communities of parasitic helminths.

This paper considers the dynamics of a host (animal) species that would grow exponentially in the absence of parasitism, and a community of parasite species that may regulate this growth. The model consists of a single differential equation for the host and one for each of the parasite species. This level of simplicity is achieved by assuming that each parasite species has a negative binomial distribution within the host population, with either zero covariance between the species (exploitation competition), or a specified covariance structure (interference competition). Conditions on the model parameters that determine the abundance of the different species are formulated, as are conditions that determine when a parasite species can invade a community and when a species is likely to be squeezed out. The results show that highly aggregated parasite species are more likely to coexist, but are less able to regulate their host population. A negative correlation between the distributions of the parasite species enhances both their ability to coexist and their ability to regulate the host population. The results of this analysis apply more generally to other systems where communities of exploiter species coexist on discretely distributed hosts, for example, insects on plants.

Animals↗

Screening for toxic effects on interspecies interactions: a mechanistic or an empirical approach?

The use of empirical and mechanistic approaches are possible in the development of tests to screen for a substance's potential to affect interspecies interactions. The advantages and disadvantages of the two approaches are discussed. An experimental study is presented, in which an empirical and a mechanistic screening test for effects on exploitative competition between bacterial species were established and perturbed with nalidixic acid. Comparison of test results indicates that the mechanistic test was faster, cheaper, more sensitive, and more quantitative. The empirical test attained similar sensitivity and quantification only if the dynamics of the competition event was continuously monitored; requiring even greater cost and time.

Bacteria↗

Cost and benefits of lizard thermoregulation.

Lizards thermoregulate by behavioral and physiological adjustments. The resultant control over metabolic processes is generally assumed to be beneficial. However, these thermoregulatory adjustments have associated costs which, if extensive, make thermoregulation impractical. We extend this idea into an abstract mathematical, cost-benefit model of thermoregulation in lizards. Investigation of the model leads to a set of predictions which includes: (1) the physiologically optimal temperature is not always the ecologically optimal temperature; (2) thermoregulation is beneficial only when associated costs are low; (3) thermal specialists will normally thermoregulate more carefully than thermal generalists unless costs are high; and (4) lizards will thermoregulate more carefully if productivity of the habitat is increased or if exploitation competition is reduced. Data on lizards, where available, generally agree with these predicitions.

Animals↗

Theoretical estimates of consumable food and probability of acquiring food in larvae of Chrysomya putoria (Diptera: Calliphoridae).

An indirect estimate of consumable food and probability of acquiring food in a blowfly species, Chrysomya putoria, is presented. This alternative procedure combines three distinct models to estimate consumable food in the context of the exploitative competition experienced by immature individuals in blowfly populations. The relevant parameters are derived from data for pupal weight and survival and estimates of density-independent larval mortality in twenty different larval densities. As part of this procedure, the probability of acquiring food per unit of time and the time taken to exhaust the food supply are also calculated. The procedure employed here may be valuable for estimations in insects whose immature stages develop inside the food substrate, where it is difficult to partial out confounding effects such as separation of faeces. This procedure also has the advantage of taking into account the population dynamics of immatures living under crowded conditions, which are particularly characteristic of blowflies and other insects as well.

Animals↗