Patch choice and risk: relative competitive ability is context dependent

The relative abilities of individual cichlids, Tilapia zillii to obtain food under scramble competition was highly repeatable between trials using a single input source, regardless of whether the input was constant or variable. However, when given a choice between two patches differing only in their temporal variability in input about an identical mean, an individual's rank based on intake in one patch was uncorrelated with either its intake in the other patch or its intake in the single-patch trials. In the two-patch trials, certain individuals both spent more time in food patches and visited patches more often than others, and overall the fish spent more time in the constant rate patch than the variable patch, leading to more items being consumed from the constant rate patch. We discuss possible causes and consequences of this dependence of relative competitive ability on the context of the foraging situation. Copyright 1999 The Association for the Study of Animal Behaviour.     

Temporal resource variability and the habitat-matching rule

Summary The ideal free distribution of competitors in a heterogeneous environment often predicts habitat matching, where the equilibrium number of consumers in a patch is proportional to resource abundance in that patch. We model the interaction between habitat matching and temporal variation in resource abundance. In one patch the rate of resource input follows a Markov chain; a second patch does not vary temporally. We predict patch use by scaling transition rates in the variable patch to the time that consumers require to respond to changes in rates of resource input. If consumers respond very quickly, habitat matching tracks temporal variability. If resource input fluctuates faster than consumers respond, habitat matching averages over the equilibrium of the Markov chain. Tracking and averaging produce the same mean resource consumption for individuals, but long-term mean occupation of the patches differs. When habitat matching tracks temporal variability in resources, consumer density in the variable patch has a lower mean and a higher variance than when habitat matching reflects only average rates of resource input.We tested our model by feeding free-living mallard ducks (Anas platyrynchos) at two artificial patches. The foragers' behavior satisfied the quantitative predictions of the model in each of two experiments.     

Aggregated distribution of resources creates competition refuges for rainforest dung beetles

This study examines the effects of resource distribution on colonisation, intra- and interspecific aggregation, and the occurrence of low-density, competition refuges for tropical dung beetles. In field experiments from central Peru, using dung pats (resource patches) of different volumes, the numbers of interacting species and total beetle biomass at individual pats increased with increasing pat volume. In two of three separate experiments (including an experiment that also varied patch density), this represented a decrease in the biomass of beetles per unit volume (biomass-density) at larger patches. The numbers of interacting tunneller species and tunneller biomass-density were also related to the distance between pats (patch density) in one of two experiments with constant numbers of pats. Closely positioned pats had generally fewer interacting species and a lower biomass-density of beetles. For the most abundant Dichotomius species, interspecific associations increased as distances increased between dung pats. The numbers of interacting species and biomass-density declined linearly under the combined effects of increasing patch density and local patch abundance in 25 m² plots. In experimentally placed grids with large numbers of pats, colonisation of pats at the edge of the grids was generally higher than at the centre of the grids for tunnellers and Eurysternus spp. but not for ball rollers; however, at least tunnellers did not readjust to avoid patches with high densities of competitors. These results indicate that an aggregated distribution of dung and natural variability in patch size contribute to species coexistence by creating low-density refuges for weaker competitors.     

Habitat assessment by parasitoids: consequences for population distribution

The ideal free distribution (IFD) is a stable distribution ofcompetitors among resource patches. For equally efficient competitors,equilibrium is reached when the per capita rate of intake equalizesacross patches. The seminal version of the IFD assumes omniscience,but populations may still converge toward the equilibrium providedthat competitors 1) accurately assess their environment by learningand 2) remain for an optimal (rate-maximizing) time on eachencountered patch. In the companion article (Tentelier C, DesouhantE, Fauvergue X. 2006. Habitat assessment by parasitoids: mechanismsfor patch time allocation. Behav Ecol. Forthcoming), it is shownthat the parasitoid wasp Lysiphlebus testaceipes adapts itsexploitation of aphid host colonies based on previous experience,in a manner consistent with these two conditions. We thereforepredicted that a randomly distributed population of initiallynaive wasps should converge toward the IFD. We tested this predictionby introducing 1300 L. testaceipes females into a 110-m² greenhousecontaining 40 host patches. Just after introduction, the parasitoidrate of gain was positively affected by host number and negativelyaffected by parasitoid number but, as predicted, these effectsvanished in the course of the experiment. Six hours after introduction,the expected rate of gain reached a constant. Surprisingly,this passage through equilibrium was not accompanied by a decreasein the coefficient of variation among gain rates or by a shiftfrom a random to an aggregated distribution of parasitoids.These results challenge our understanding of the link betweenindividual behavior and population distribution.     

An evolutionary explanation of the aggregation model of species coexistence

In ecology, the 'aggregation model of coexistence' provides a powerful concept to explain the unexpectedly high species richness of insects on ephemeral resources like dung pats, fruits, etc. It suggests that females aggregate their eggs across resource patches, which leads to an increased intraspecific competition within occupied patches and a relatively large number of patches that remain unoccupied. This provides competitor-free patches for heterospecifics, facilitating species coexistence. At first glance, deliberately causing competition among the females' own offspring and leaving resources to heterospecific competitors seems altruistic and incompatible with individual fitness maximization, raising the question of how natural selection operates in favour of egg aggregation on ephemeral resource patches. Allee effects that lead to fitness maxima at intermediate egg densities have been suggested, but not yet detected. Using drosophilid flies on decaying fruits as a study system, we demonstrate a hump-shaped relationship between egg density and individual survival probability, with maximum survivorship at intermediate densities. This pattern clearly selects for egg aggregation and resolves the possible conflict between the ecological concept of species coexistence on ephemeral resources and evolutionary theory.     

Competition among plant species that interact with their environment at different spatial scales

Clonal plants that are physiologically integrated might perceive and interact with their environment at a coarser resolution than smaller, non-clonal competitors. We develop models to explore the implications of such scale asymmetries when species compete for multiple depletable resources that are heterogeneously distributed in space across two patches. Species are either 'non-integrators', whose growth in each patch depends on resource levels in that patch alone, or 'integrators', whose growth is equal between patches and depends on average resource levels across patches. Integration carried both benefits and costs. It tended to be advantageous in poorer patches, where the integrators drew resources down further than the non-integrators (more easily excluding competitors) and might persist by using resources from richer adjacent patches. Integration tended to be disadvantageous in richer patches, where integrators did not draw resources down as far (creating an opportunity for competitors) and could be excluded due to the cost of supporting growth in poorer adjacent patches. Complementarity between patches (each rich in a separate resource) favoured integrators. Integration created new opportunities for local coexistence, and for delayed susceptibility of patches to invasion, but eliminated some opportunities for regional coexistence. Implications for the interpretations of species' zero net growth isoclines and Rs are also discussed.     

Transition from a random to an ideal free to an ideal despotic distribution: the effect of individual difference in growth

Individual differences in growth can lead to a monopolistic form of food competition. We studied the long-term transition in the mode of competition and the distribution of individuals between food patches of the cloned salmonid fish, Oncorhynchus masou ishikawae, in the laboratory. This transition was accompanied by growth depensation, i.e., the increase over time in the variance of size between individuals resulting from the differences in individual growth rates. The 120-cm experimental tanks were divided into two compartments (patches) between which an opaque partition was placed. Fish were able to move freely between the patches and therefore were able to assess the patch quality using long-term memory, but they were not able to see the food input in the other patch directly. The distribution between the two food patches, the amount of food gained, and the growth and the agonistic behavior of four groups of six individuals were observed over 4 weeks. We found that (1) within-group variation in body weight increased with time; (2) on average, the better patch was used by more individuals than predicted by a random distribution but fewer individuals than predicted by an ideal free distribution, and (3) the distribution and pattern of resource use by the fish changed over the 4-week experimental period from a random distribution to an ideal free distribution and finally to an ideal despotic distribution. We suggest that growth depensation causes the long-term change in the spatial distribution and pattern of resource use by competitors. Received: December 19, 2000 / Accepted: March 19, 2001     

Choosing where to feed: the influence of competition on feeding behaviour of cod, Gadus morhua L.

Groups of cod, Gadus morhua L, presented with two feeding patches with a food abundance ratio of 2:1, distributed themselves between the patches in a ratio of 2.5:1. This is slightly higher than the 2:1 ratio predicted by the ideal free distribution theory. Large differences were observed in competitive ability between individual fish. A strong correlation was found between feeding success of individuals and time spent in a feeding patch. The more successful competitors caught about 2.5 times as many food items in the rich patch as in the poor patch. The less successful competitors caught an equal amount of food in both patches. All competitors, however, spent significantly more time in the rich patch. These results suggest that hunting success is the most important factor in assessing patch quality. However, it is not the only parameter which cod use in deciding where to feed.     

Consumer movement through differentially subsidized habitats creates a spatial food web with unexpected results

Energy and nutrient flow between habitats, or allochthonous input, can have a significant impact on food web dynamics. Previous theory demonstrated that resource abundance decreases in habitats where consumers are subsidized. Here we examine the effect of subsidies that are available in localized parts of a habitat (such as near the shore in a marine‐subsidized terrestrial ecosystem) with a two‐patch model in which consumers move between patches, resources are stationary, and consumers receive the subsidy in only one of the two patches. In contrast to previous theory, our results show that subsidized consumers can increase resource abundance, though only in the subsidized patch. Furthermore, the total resource population responds positively to increasing consumer movement. These results demonstrate the importance of spatial heterogeneity in food web dynamics and the need for further examination of the role of space in multispecies trophic webs.     

Persistence and periodic orbits of a three-competitor model with refuges.

We consider a model composed of four patches. One patch has three competing species forming a heteroclinic cycle within the patch. The remaining patches are refuges for the three competitors, and each species can diffuse between the competitive patch and its refuge. It is proved that the model can be made persistent by the introduction of the refuges for the competitors even if the isolated competitive patch has an attracting heteroclinic cycle. Further it is shown that Hopf bifurcation is possible when we change the value of the diffusion constant and periodic orbits may exist in a specific case.     

Spatial heterogeneity of biomass turnover has contrasting effects on synchrony and stability in trophic metacommunities

Spatial heterogeneity is a fundamental feature of ecosystems, and ecologists have identified it as a factor promoting the stability of population dynamics. In particular, differences in interaction strengths and resource supply between patches generate an asymmetry of biomass turnover with a fast and a slow patch coupled by a mobile predator. Here, we demonstrate that asymmetry leads to opposite stability patterns in metacommunities receiving localized perturbations depending on the characteristics of the perturbed patch. Perturbing prey in the fast patch synchronizes the dynamics of prey biomass between the two patches and destabilizes predator dynamics by increasing the predator's temporal variability. Conversely, perturbing prey in the slow patch decreases the synchrony of the prey's dynamics and stabilizes predator dynamics. Our results have implications for conservation ecology and suggest reinforcing protection policies in fast patches to dampen the effects of perturbations and promote the stability of population dynamics at the regional scale.     

Competition in a group of equal foragers

Abstract Using techniques from renewal process theory, we build a stochastic model for gain accumulation in a group of equal competitors foraging in a patchy environment. The model for gain of the individuals is based on the waiting times between subsequent prey encounters by the group. These waiting times depend on the number of foragers in the group. A single parameter of this dependency encompasses a variety of foraging scenarios, from co-operation to scramble. With constant patch size, correlations between gains of any pair of foragers are negative. This dependency is most intense in small groups. Increased variation in patch size makes correlations in gains between group members positive irrespective of the group size. For a solitary forager, variance in gain approaches zero with increasing time in the patch. For an individual member in a group, variance grows monotonically. Thus, depending on the patch departure rule controlling the time to be spent in the patch, solitary foragers may have a smaller variance in gain than members in a group. As solitary foragers also potentially harvest all prey in the patch, it is hard to believe that grouping behavior would evolve solely on the basis of foraging.     

Contributions of high- and low-quality patches to a metapopulation with stochastic disturbance

Studies of time-invariant matrix metapopulation models indicate that metapopulation growth rate is usually more sensitive to the vital rates of individuals in high-quality (i.e., good) patches than in low-quality (i.e., bad) patches. This suggests that, given a choice, management efforts should focus on good rather than bad patches. Here, we examine the sensitivity of metapopulation growth rate for a two-patch matrix metapopulation model with and without stochastic disturbance and found cases where managers can more efficiently increase metapopulation growth rate by focusing efforts on the bad patch. In our model, net reproductive rate differs between the two patches so that in the absence of dispersal, one patch is high quality and the other low quality. Disturbance, when present, reduces net reproductive rate with equal frequency and intensity in both patches. The stochastic disturbance model gives qualitatively similar results to the deterministic model. In most cases, metapopulation growth rate was elastic to changes in net reproductive rate of individuals in the good patch than the bad patch. However, when the majority of individuals are located in the bad patch, metapopulation growth rate can be most elastic to net reproductive rate in the bad patch. We expand the model to include two stages and parameterize the patches using data for the softshell clam, Mya arenaria. With a two-stage demographic model, the elasticities of metapopulation growth rate to parameters in the bad patch increase, while elasticities to the same parameters in the good patch decrease. Metapopulation growth rate is most elastic to adult survival in the population of the good patch for all scenarios we examine. If the majority of the metapopulation is located in the bad patch, the elasticity to parameters of that population increase but do not surpass elasticity to parameters in the good patch. This model can be expanded to include additional patches, multiple stages, stochastic dispersal, and complex demography.     

Spatial ecology of multiple parasitoids of a patchily‐distributed host: implications for species coexistence

1. The coexistence of multiple species sharing similar but spatially fragmented resources (e.g. parasitoids sharing a host species) may depend on their relative competitive and dispersal abilities, or on fine‐scale resource partitioning. Four generalist and one specialist parasitoid species associated with the holly leaf miner, Phytomyza ilicis, in a woodland network of 127 holly trees were investigated. 2. To understand coexistence and persistence of these potential competitors, patterns of occurrence in relation to patch size and isolation, vertical stratum within patches, and incidence and abundance of potential competitors were documented. Field experiments creating empty habitat patches suggested that dispersal rather than local demographic processes determines abundance and incidence. 3. Parasitoids showed species‐specific responses to patch properties, with the incidence of species determined mostly by patch size. Parasitism rates were less clearly related to patch characteristics, but parasitism rates for most species were lower in patches where the numerically dominant parasitoid species, Chrysocharis gemma, was present. No evidence of vertical stratification was found in species composition or abundance within patches, making it unlikely that coexistence is enhanced by fine‐scale resource division. 4. Overall, the patterns detected may be attributed to the distribution of C. gemma and differences in species' ecology other than dispersal ability. The life history of C. gemma may allow it to pre‐emptively exploit a large fraction of the available hosts, avoiding direct competition with other parasitoids. In contrast, direct competition is more likely among the pupal parasitoids Cyrtogaster vulgaris, Chrysocharis pubicornis, and Sphegigaster flavicornis which have a similar biology and phenology. For these species, coexistence may be facilitated by contrasting incidence in relation to patch size and isolation.     

The ideal free distribution when the resource is variable

On the basis of the ideal free distribution (IFD) model, twostochastic models that incorporate the uncertainty of the informationused for decision making were considered to investigate theeffects of the variability in the resource supply rate on theIFD under continuous input conditions. In the uncertain-informationmodel, competitors cannot trace the variation of the supplyrate and use the expectation of the supply rate or previouspayoffs for decision making. Both submodels predict matchingof means, in which the average number of competitors for eachpatch is proportional to the average supply rate in the patch.In the perfect-information model, competitors continuously knowand trace the environment conditions. Numerical predictionsdepend on the relative size of the resource variance betweenpatches. When the resource variance in the good patch is sufficientlylarger than that in the poor patch, it predicts undermatchingof means; when the variance of the supply rate for each patchis small and proportional to the average of the supply ratein the patch, it predicts matching of means; and when the resourcevariance in the poor patch is larger than (or equal to) thatin the good patch, it predicts overmatching of means. Theseresults indicate the importance of clarifying the assumptionon the uncertainty in information for decision making and thetype of the resource variance for the test of the IFD underconditions where the resource supply rate is stochastic.     

Indirect effects of heterospecific interactions on progeny size through maternal stress

Maternal effects are increasingly being recognized as an important pre-natal source of life history variation in the next generation. The present study uses a field experiment to explore the influence of heterospecific interactions on the reproductive output and offspring characteristics of a common Indo-Pacific damselfish, Pomacentrus amboinensis . On the Great Barrier Reef pairs of breeding P. amboinensis were placed on isolated patch reefs and to half of the pairs resource competitors (other planktivorous damselfishes), and predators of eggs and juveniles were added. Females inhabiting patches with heterospecifics had more aggressive interactions and higher levels of the stress hormone, cortisol. Neither the number of clutches nor number of eggs produced differed among treatments. The size of larvae at hatching was found to be reduced as a result of the stress associated with increased interactions with heterospecific and the transfer of cortisol to offspring. This stress-associated mechanism appears to be an important and directional source of life history variability, but the individual nature of the maternal response is likely to result in a conclusion of a diversified bet hedging reproductive strategy when viewed at the local population level. These findings highlight the complex determinants of individual success and the important role of parental well-being in the population dynamics of the next generation.     

Larval competition causes the difference in male ejaculate expenditure in Callosobruchus maculatus

The seed beetle Callosobruchus maculatus larvae exhibit two types of resource competition: scramble, in which a resource is shared, and contest, in which the resource is monopolized. This difference in larval behavior results in different adult densities. Under contest competition, adult density remains constant regardless of larval density, but under scramble competition, adult density increases with larval density. This in turn affects mating frequency during adulthood, and thus, the intensity of sexual selection operating on males. In this study, we examined the relationship between larval competition types and male reproductive investment in mating. We assessed the male ejaculate expenditure per mating across geographic strains of C. maculatus. The male investment (ejaculate expenditure) increased with the degree of scramble competition and decreased with the degree of contest competition. We therefore suggest that males experience different selective pressures depending on the type of larval competition: scramble type males are selected for increased reproductive investment.     

Physiological Ecology Meets the Ideal-free Distribution: Predicting the Distribution of Size-structured Fish Populations Across Temperature Gradients

We describe a habitat selection model that predicts the distribution of size-structured groups of fish in a habitat where food availability and water temperature vary spatially. This model is formed by combining a physiological model of fish growth with the logic of ideal free distribution (IFD) theory. In this model we assume that individuals scramble compete for resources, that relative competitive abilities of fish vary with body size, and that individuals select patches that maximize their growth rate. This model overcomes limitations in currently existing physiological and IFD-based models of habitat selection. This is because existing physiological models do not take into account the fact that the amount of food consumed by a fish in a patch will depend on the number of competitors there (something that IFD theory addresses), while traditional IFD models do not take into account the fact that fish are likely to choose patches based on potential growth rate rather than gross food intake (something that physiological models address). Our model takes advantage of the complementary strengths of these two approaches to overcome these weaknesses. Reassuringly, our model reproduces the predictions of its two constituent models under the simple conditions where they apply. When there is no competition for resources it mimics the physiological model of habitat selection, and when there is competition but no temperature variation between patches it mimics either the simple IFD model or the IFD model for unequal competitors. However, when there are both competition and temperature differences between patches our model makes different predictions. It predicts that input-matching between the resource renewal rate and the number of fish (or competitive units) in a patch, the hallmark of IFD models, will be the exception rather than the rule. It also makes the novel prediction that temperature based size-segregation will be common, and that the strength and direction of this segregation will depend on per capita resource renewal rates and the manner in which competitive weight scales with body size. Size-segregation should become more pronounced as per capita resource abundance falls. A larger fish/cooler water pattern is predicted when competitive ability increases more slowly than maximum ration with body size, and a smaller fish/cooler water pattern is predicted when competitive ability increases more rapidly than maximum ration with body size.     

The ideal free distribution of clonal plant's ramets among patches in a heterogeneous environment

The plastic response of clonal plant to different patch quality is not always the same and the degree is different too. So the result of this kind of foraging behaviour is different. In order to make clear whether the ramtes stay in favourable patches and get the quantitative relationship between the ramets distribution among patches and the available resource amount in heterogeneous environment, we develop a theoretical work under ideal free distribution (IFD) theory framework by neglecting some morphological plasticity of the spacer in this article. The results of our general model show that the ramet distribution should obey input matching rule at equilibrium. That means the ratio of ramet number in different patches should be equal to the ratio of available resource amount in these patches. We also use the simulation to predict the distribution pattern under history mattering. The results show that the initial ramets number has significant influence on the final distribution: over matching and under matching both can occur. More initial ramets in favourable patch result in over matching and more initial ramets in unfavourable patch result in under matching. The degree of the deviation from input matching rule is great when the difference of patches is small. These results prove that ideal free distribution theory works the same with animals. The ramets can stay in favourable patches sometimes in spite of the plasticity of the spacer, and the distribution depends on both patch quality and the history factors. But these results are true only when the functional response is type II.