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PubMed2003

The forager's dilemma: food sharing and food defense as risk-sensitive foraging options.

Although many variants of the hawk-dove game predict the frequency at which group foraging animals should compete aggressively, none of them can explain why a large number of group foraging animals share food clumps without any overt aggression. One reason for this shortcoming is that hawk-dove games typically consider only a single contest, while most group foraging situations involve opponents that interact repeatedly over discovered food clumps. The present iterated hawk-dove game predicts that in situations that are analogous to a prisoner's dilemma, animals should share the resources without aggression, provided that the number of simultaneously available food clumps is sufficiently large and the number of competitors is relatively small. However, given that the expected gain of an aggressive animal is more variable than the gain expected by nonaggressive individuals, the predicted effect of the number of food items in a clump-clump richness-depends on whether only the mean or both the mean and variability associated with payoffs are considered. More precisely, the deterministic game predicts that aggression should increase with clump richness, whereas the stochastic risk-sensitive game predicts that the frequency of encounters resulting in aggression should peak at intermediate clump richnesses or decrease with increasing clump richness if animals show sensitivity to the variance or coefficient of variation, respectively.

Animals↗

PubMed2003

The effects of enrichment on the dynamics of apparent competitive interactions in stage-structured systems.

In the absence of other limiting factors, assemblages in which species share a common, effective natural enemy are not expected to persist. Although a variety of mechanisms have been postulated to explain the coexistence of species that share natural enemies, the role of productivity gradients has not been explored in detail. Here, we examine how enrichment can affect the outcome of apparent competition. We develop a structured resource/consumer/natural enemy model in which the prey are exposed to attacks during a vulnerable life phase, the length of which depends on resource availability. With a single prey species, the model exhibits the "paradox of enrichment," with unstable dynamics at high levels of resource productivity. We extend this model to consider two prey species linked by a shared predator, each with their own distinct resource base. We derive invasion and stability conditions and examine how enrichment influences prey species exclusion and coexistence. Contrary to expectations from simpler, prey-dependent models, apparent competition is not necessarily strong at high productivity, and prey species coexistence may thus be more likely in enriched environments. Further, the coexistence of apparent competitors may be facilitated by unstable dynamics. These results contrast with the standard theory that apparent competition in productive environments leads to nonpersistent interactions and that coexistence of multispecies interactions is more likely under equilibrial conditions.

Animals↗

PubMed2004

Priority versus brute force: when should males begin guarding resources?

When should males begin guarding a resource when both resources and guarders vary in quality? This general problem applies, for example, to migrant birds occupying territories in the spring and to precopula in crustaceans where males grab females before they molt and become receptive. Previous work has produced conflicting predictions. Theory on migrant birds predicts that the strongest competitors should often arrive first, whereas some models of mate guarding have predicted that the strongest competitors wait and then simply usurp a female from a weaker competitor. We build a general model of resource guarding that allows varying the ease with which takeovers occur. The model is phrased in terms of mate-guarding crustaceans, but the same logic can be applied to other forms of resource acquisition where priority plays a role but takeovers might be possible too. The race to secure breeding positions can lead to strong competitors (large males) taking females earliest, even though this means accepting a lower-quality female. Paradoxically, this means that small males, which have fewer breeding opportunities, are more choosy than larger ones. Such solutions are found when takeovers are impossible. The easier the takeovers and the higher the rate of finding guarded resources, the more likely are solutions where guarding durations are short, where strong competitors begin guarding only just before breeding, and where they do this by usurping the resource. The relationship between an individual's competitive ability and its timing of resource acquisition can also be nonlinear if takeovers are moderately common; if this is the case, then males of intermediate size guard the longest.

Animals↗

PubMed1986

The intensity of competition versus its importance: an overlooked distinction and some implications.

The intensity of competition is a physiological concept, related directly to the well-being of individual organisms but only indirectly and conditionally to their fitness, and even more indirectly to the evolution of populations and the structure of communities. The importance of competition is primarily an ecological and evolutionary concept, related directly to the ecology and fitness of individuals but only indirectly to their physiological states. The intensity of competition is not necessarily correlated with the intensities of predation, disturbance, abiotic stress, or other ecological processes. The importance of competition is necessarily relative to the importances of other processes. Intensity refers primarily to the process of present competition, whereas importance refers primarily to the products of past competition. The distinction between the intensity and the importance of competition clarifies two long-standing ecological debates. Some ecologists have proposed that competition is greater in more stressful habitats, others the opposite, and still others that no such relationship exists. Evidence cited to refute or support these positions often confuses intensity and importance. Distinguishing between them focuses questions more sharply and indicates what sorts of new evidence should be sought. The more widely known debate over the prevalence of competition as an agent of community structure is a debate about the importance of competition, but evidence about the intensity of competition has often been used by both sides. We argue that intensity and importance need not be correlated, and so measurements of the intensity of competition are not directly relevant to this debate. This distinction also generates testable hypotheses and suggests directions for research. For example, we hypothesize that competition can be unimportant even if it is very intense: no such hypothesis is possible unless importance is distinguished from intensity. We discuss the application of these ideas to methods and theories used to study competition, ecological communities, and the evolution of competitive ability. We advocate a research approach that presumes multiple, interacting causes, including competition, affecting community structure, and we show how the distinction between intensity and importance helps to make this feasible.

Animals↗

PubMed1989

One selectionist's perspective.

Following introductory comments expressing doubts about the validity of genetic load and Haldane's "cost of natural selection," the role of selection (expressed as the average number of adult daughters per female) on gene frequencies in populations has been partitioned into population and time arenas. The population arena (a geometric plane) deals with the fitnesses of different genotypes under the many situations encountered by individual members of the population in a single generation; average fitnesses of carriers of various genotypes are obtained by calculating across these many situations. The population arena includes the point signifying that, on the average, each mother leaves one daughter as her replacement within the population. It is the plane within which evolutionarily significant norms of reaction exist. The time arena is also a (geometric) plane, one that is composed of the edge-on limit (average fitness) of each successive population arena. It does not include the effects of individual situations on relative fitnesses within each population arena; it encompasses only the temporal sequence of average relative fitnesses. Amino acid substitutions in proteins and base-pair substitutions in DNA are events of concern in the time arena; within the population arena, however, gene action (not merely gene structure) is a matter of considerable concern. Thus, the discussions of the 1950s and 1960s regarding genetic variation which were reasonable within the population arena seem less so within the time arena where structural, rather than functional, variation is stressed. The function-structure dichotomy is entangled with the neutralist-selectionist controversy.

Alleles↗

PubMed1990

The ecology and evolution of inducible defenses.

Inducible defenses are responses activated through a previous encounter with a consumer or competitor that confer some degree of resistance to subsequent attacks. While the importance of inducible resistance has long been known in host-parasite interactions, it is only recently that its importance has emerged in other natural systems. Although the structural defenses produced by invertebrates to their competitors and predators are by no means the same as an immune response triggered by parasites, these responses all share the properties of (1) specificity, (2) amplification and (3) memory. This review discusses the following ecological consequences and evolutionary causes of inducible defenses: (1) Inducible defenses render historical factors important in biological interactions and can affect the probability of individual survival and growth, as well as affect population dynamics of consumers in some circumstances. (2) Although the benefits of inducible defenses are often balanced by fitness costs, including reduced growth, reproductive output and survivorship, the role of costs and benefits in the evolution of inducible defenses is by no means clear. A more integrated approach would involve a multivariate analysis of the role of natural selection on the inducible characters of interest, their norms of reaction and correlated fitness characters. (3) The disproportionate representation of inducible, morphological defenses among clonal organisms may be due to both a higher rate of origination and enhanced selection to maintain these defenses in clonal taxa. (4) Inducible defenses should be most common when reliable cues are available, attacks by biological agents are unpredictable, and the fitness gains of defenses are balanced by the costs. An integrated approach to studying inducible defenses would thus combine mechanistic estimates of costs, population-level estimates of defense effectiveness, and genetic estimates of correlations between fitness and inducible characters. This will allow us to estimate rates of evolution in these phenotypically plastic threshold characters.

Animals↗

PubMed2004

Stoichiometrically explicit competition between grazers: species replacement, coexistence, and priority effects along resource supply gradients.

Assuming key trade-offs among interactors, several models (resource ratio, keystone predation, intraguild predation) predict changes in species composition over resource supply gradients. Ecological stoichiometry could also predict compositional shifts of grazers over gradients of nutrient and light supply through a mechanism involving (mis)matches between elemental body composition of grazers and plants. This hypothesis is explored here using a suite of two-grazer, one-plant models that incorporate three key components: plant production depends on light and nutrients, nutrient content of plants can vary, and homeostatic grazers can be carbon or nutrient limited. The results from this suite closely resemble the classical resource ratio model describing plant competition for two resources. Here, the models predict shifts of grazer composition along resource supply gradients if species trade off competitive abilities for plant carbon and nutrients. Given this trade-off, superior nutrient competitors should dominate low nutrient environments, and superior carbon competitors should dominate high nutrient environments. At intermediate nutrient supply, species can coexist at a stable equilibrium, or alternative stable states emerge, depending on how grazers impact their resources. These results depend on food web architecture, however. For instance, predators can alter or reduce possibilities for stoichiometry-mediated coexistence of grazers.

Animals↗

PubMed2004

Large species shifts triggered by small forces.

Changes in species composition of communities seem to proceed gradually at first sight, but remarkably rapid shifts are known to occur. Although disrupting disturbances seem an obvious explanation for such shifts, evidence for large disturbances is not always apparent. Here we show that complex communities tend to move through occasional catastrophic shifts in response to gradual environmental change or evolution. This tendency is caused by multiple attractors that may exist in such systems. We show that alternative attractors arise robustly in randomly generated multispecies models, especially if competition is symmetrical and if interspecific competition is allowed to exceed intraspecific competition. Inclusion of predators as a second trophic level did not alter the results greatly, although it reduced the probability of alternative attractors somewhat. These results suggest that alternative attractors may commonly arise from interactions between large numbers of species. Consequently, the response of complex communities to environmental change is expected to be characterized by hysteresis and sudden shifts. Some unexplained regime shifts observed in ecosystems could be related to alternative attractors arising from complex species interactions. Additionally, our results support the idea that ancient mass extinctions may partly be due to an intrinsic loss of stability of species configurations.

Animals↗

PubMed2004

Sexual selection: harem size and the variance in male reproductive success.

Sexual selection is potentially stronger than natural selection when the variance in male reproductive fitness exceeds all other components of fitness variance combined. However, measuring the variance in male reproductive fitness is difficult when nonmating males are absent, inconspicuous, or otherwise difficult to find. Omitting the nonmating males inflates estimates of average male reproductive success and diminishes the variance, leading to underestimates of the potential strength of sexual selection. We show that, in theory, the proportion of the total variance in male fitness owing to sexual selection is approximately equal to H, the mean harem size, as long as H is large and females are randomly distributed across mating males (i.e., Vharem=H). In this case, mean harem size not only provides an easy way to estimate the potential strength of sexual selection but also equals the opportunity for sexual selection, I(mates). In nature, however, females may be overdispersed with Vharem H. We show that H+(k-1) is a good measure of the opportunity for sexual selection, where k is the ratio Vharem/H. A review of mating system data reveals that in nature the median ratio for Vharem/H is 1.04, but as H increases, females tend to become more aggregated across mating males with V(harem) two to three times larger than H.

Animals↗

PubMed2004

Demographic theory of coral reef fish populations with stochastic recruitment: comparing sources of population regulation.

The effects of three forms of density-dependent regulation were explored in model coral reef fish populations: top-down (predation), bottom-up (competition for food), and pelagic (non-reef-based mechanisms) control. We describe the demographic responses of both biomass and numbers of adult fish, predicting the mean and the variance of temporal fluctuations resulting from stochastic recruitment of juveniles. We find that top-down control acts by suppressing variability of numbers of fish, which in turn suppresses the variability of biomass. Bottom-up control has no effect on fluctuations of numbers of fish, though it strongly reduces fluctuations of biomass. Because fecundity of fish is directly linked to body mass, the regulation of biomass tightly regulates reproductive output independently of the number of individuals in the population. Finally, populations under pelagic control experience bounded fluctuations of biomass and numbers directly proportional to the bounded fluctuations of recruitment. The demographic signatures predicted from both bottom-up and pelagic control are consistent with current evidence supporting the recruitment limitation hypothesis in reef fish ecology. We propose tests to discriminate the dominant mode of density-dependent regulation using qualitative trends in time series demographic data across environmental clines.

Animals↗

PubMed2004

Coexistence of cycling and dispersing consumer species: Armstrong and McGehee in space.

Two competing consumer species may coexist using a single homogeneous resource when the more efficient consumer--the one having the lowest equilibrium resource density--has a more nonlinear functional response that generates consumer-resource cycles. We extend this model of nonequilibrium coexistence, as proposed by Armstrong and McGehee, by putting the interaction into a spatial context using two frameworks: a spatially explicit individual-based model and a spatially implicit metapopulation model. We find that Armstrong and McGehee's mechanism of coexistence can operate in a spatial context. However, individual-based simulations suggest that decreased dispersal restricts coexistence in most cases, whereas differential equation models of metapopulations suggest that a low rate of dispersal between subpopulations often increases the coexistence region. This difference arises in part because of two potentially opposing effects on coexistence due to the asynchrony in the temporal dynamics at different locations. Asynchrony implies that the less efficient species is more likely to be favored in some spatial locations at any given time, which broadens the conditions for coexistence. On the other hand, asynchrony and dispersal can also reduce the amplitude of local population cycles, which restricts coexistence. The relative influence of these two effects depends on details of the population dynamics and the representation of space. Our results also demonstrate that coexistence via the Armstrong-McGehee mechanism can occur even when there is little variation in the global densities of either the consumers or the resource, suggesting that empirical studies of the mechanisms should measure densities on several spatial scales.

Animal Migration↗

PubMed2005

Transient dynamics limit the effectiveness of keystone predation in bringing about coexistence.

We analyze the transient dynamics of simple models of keystone predation, in which a predator preferentially consumes the dominant of two (or more) competing prey species. We show that coexistence is unlikely in many systems characterized both by successful invasion of either prey species into the food web that lacks it and by a stable equilibrium with high densities of all species. Invasion of the predator-resistant consumer species often causes the resident, more vulnerable prey to crash to such low densities that extinction would occur for many realistic population sizes. Subsequent transient cycles may entail very low densities of the predator or of the initially successful invader, which may also preclude coexistence of finite populations. Factors causing particularly low minimum densities during the transient cycles include biotic limiting resources for the prey, limited resource partitioning between the prey, a highly efficient predator with relatively slow dynamics, and a vulnerable prey whose population dynamics are rapid relative to the less vulnerable prey. Under these conditions, coexistence of competing prey via keystone predation often requires that the prey's competitive or antipredator characteristics fall within very narrow ranges. Similar transient crashes are likely to occur in other food webs and food web models.

Animals↗

PubMed2005

Scale-dependent feedback and regular spatial patterns in young mussel beds.

In the past decade, theoretical ecologists have emphasized that local interactions between predators and prey may invoke emergent spatial patterning at larger spatial scales. However, empirical evidence for the occurrence of emergent spatial patterning is scarce, which questions the relevance of the proposed mechanisms to ecological theory. We report on regular spatial patterns in young mussel beds on soft sediments in the Wadden Sea. We propose that scale-dependent feedback, resulting from short-range facilitation by mutual protection from waves and currents and long-range competition for algae, induces spatial self-organization, thereby providing a possible explanation for the observed patterning. The emergent self-organization affects the functioning of mussel bed ecosystems by enhancing productivity and resilience against disturbance. Moreover, self-organization allows mussels to persist at algal concentrations that would not permit survival of mussels in a homogeneous bed. Our results emphasize the importance of self-organization in affecting the emergent properties of natural systems at larger spatial scales.

Animals↗

PubMed2005

The evolution of phenotypic polymorphism: randomized strategies versus evolutionary branching.

A population is polymorphic when its members fall into two or more categories, referred to as alternative phenotypes. There are many kinds of phenotypic polymorphisms, with specialization in reproduction, feeding, dispersal, or protection from predators. An individual's phenotype might be randomly assigned during development, genetically determined, or set by environmental cues. These three possibilities correspond to a mixed strategy of development, a genetic polymorphism, and a conditional strategy. Using the perspective of adaptive dynamics, I develop a unifying evolutionary theory of systems of determination of alternative phenotypes, focusing on the relative possibilities for random versus genetic determination. The approach is an extension of the analysis of evolutionary branching in adaptive dynamics. It compares the possibility that there will be evolutionary branching, leading to genetic polymorphism, with the possibility that a mixed strategy evolves. The comparison is based on the strength of selection for the different outcomes. An interpretation of the resulting criterion is that genetic polymorphism is favored over random determination of the phenotype if an individual's heritable genotype is an adaptively advantageous cue for development. I argue that it can be helpful to regard genetic polymorphism as a special case of phenotypic plasticity.

Adaptation, Physiological↗

PubMed2005

Changes in community size affect the outcome of competition.

We examine the role of stochasticity and competitive ability in affecting competition between two species using models derived for population genetics. Just as changing population size affects the fixation of a new mutation, we show that changing the total number of competitors (i.e., community size) can alter the course of competitive exclusion across a wide range of initial starting densities of the two competing species. Shifts in competitive exclusion occur because changes in community size affect the relative importance of competitive ability and stochasticity in affecting the outcome of competition, potentially allowing inferior invaders to usurp superior residents. By shifting the role of stochasticity and competitive ability, any process that changes the total number of competitors in a habitat (e.g., disturbance, eutrophication, fragmentation, predation) may lead to shifts in competitive exclusion and the composition of communities.

Competitive Behavior↗

PubMed2006

Nonrandom mating preserves intrasexual polymorphism and stops population differentiation in sexual conflict.

Evolutionary conflict between the sexes is predicted to lead to sexual arms races in which male adaptations for acquiring mates ("offense" traits) are met by female counteradaptations--for example, to reduce mating rate ("defense" traits). Such coevolutionary chases may be perpetual. However, we show here that the coevolutionary process may also lead to a stable state in which multiple offense-defense trait pairs are maintained. This type of polymorphism below the species level is a result of sexual conflict in combination with nonrandom mating. Our results show that if nonrandom mating occurs with respect to male and female conflict traits, genetic correlations will act to stabilize the trait frequencies so that all morphs are maintained. We discuss the results in special relation to the evolution of female polymorphism in diving beetles and argue that the process we describe may be a general force that maintains polymorphism in other taxa as well.

Animals↗

PubMed2006

Intraspecific competitive divergence and convergence under assortative mating.

Ecologically driven sympatric speciation has received much attention recently. We investigate a multilocus model of a quantitative trait that is under frequency-dependent selection caused by intraspecific competition and acts as mating character for assortment. We identify the conditions that lead to the establishment of reproductively isolated clusters. This may be interpreted as evolutionary splitting or sympatric speciation. In our model, there are parameters that independently determine the strength of assortment, the costs for being choosy, and the strength of frequency-dependent natural selection. Sufficiently strong frequency dependence leads to disruptive selection on the phenotypes. The population consists of (sexual) haploid individuals. If frequency dependence is strong enough to induce disruptive selection and costs are absent or low, the result of evolution depends in a distinctive nonlinear way on the strength of assortment: under moderately strong assortment, less genetic variation is maintained than under weak or strong assortment, and sometimes there is none at all. Evolutionary splitting occurs only if frequency dependence and assortment are both strong enough and costs are low. Even then, the evolutionary outcome depends on the genetics and the initial conditions. The roles of the number of loci, of linkage, and of asymmetric selection are also explored.

Animals↗

PubMed2006

Inedible producers in food webs: controls on stoichiometric food quality and composition of grazers.

Ecological stoichiometry and food web theories focus on distinct mechanisms that shape communities. These mechanisms, however, likely interact in ways that neither theory alone addresses. To illustrate, we show how a model that tracks flow of energy and nutrients through two producers and two grazers reveals two indirect, interrelated roles for "neutrally inedible" producers. First, inedible producers can exert controls over the nutrient content of edible producers and indirectly influence whether grazers are nutrient or energy limited. Second, through these controls, inedible producers can shape community assembly by excluding grazers that are weak competitors for nutrients contained in edible producers. A mesocosm experiment revealed patterns consistent with both predictions: high abundances of inedible algae were accompanied by low phosphorus contents of edible algae and low abundances of the grazer Daphnia. Both lines of inference suggest that interactions between stoichiometry and plant heterogeneity may shape plankton communities.

Animal Nutritional Physiological Phenomena↗