Evolutionary games and two species population dynamics

Competition between species has long been modeled by population dynamics based on total numbers of each species. Recently, the evolution of strategy frequencies has been used successfully for competition models between individuals. In this paper, we illustrate that these two views of competition are compatible. It is shown that the rate of intra and interspecific competitions between individuals largely determines the population dynamics. Competition models over a single common resource and predator-prey models are developed from this individual competition approach. In particular, the equilibrium strategies in a co-evolving predator-prey system are shown to be more stable than the predicted strategy cycling of standard evolutionary game theory.     

Habitat selection and population regulation in temporally fluctuating environments

Understanding and predicting the distribution of organisms in heterogeneous environments lies at the heart of ecology, and the theory of density-dependent habitat selection (DDHS) provides ecologists with an inferential framework linking evolution and population dynamics. Current theory does not allow for temporal variation in habitat quality, a serious limitation when confronted with real ecological systems. We develop both a stochastic equivalent of the ideal free distribution to study how spatial patterns of habitat use depend on the magnitude and spatial correlation of environmental stochasticity and also a stochastic habitat selection rule. The emerging patterns are confronted with deterministic predictions based on isodar analysis, an established empirical approach to the analysis of habitat selection patterns. Our simulations highlight some consistent patterns of habitat use, indicating that it is possible to make inferences about the habitat selection process based on observed patterns of habitat use. However, isodar analysis gives results that are contingent on the magnitude and spatial correlation of environmental stochasticity. Hence, DDHS is better revealed by a measure of habitat selectivity than by empirical isodars. The detection of DDHS is but a small component of isodar theory, which remains an important conceptual framework for linking evolutionary strategies in behavior and population dynamics.     

On the relationship between niche and distribution

Applications of Hutchinson's n -dimensional niche concept are often focused on the role of interspecific competition in shaping species distribution patterns. In this paper, I discuss a variety of factors, in addition to competition, that influence the observed relationship between species distribution and the availability of suitable habitat. In particular, I show that Hutchinson's niche concept can be modified to incorporate the influences of niche width, habitat availability and dispersal, as well as interspecific competition per se . I introduce a simulation model called NICHE that embodies many of Hutchinson's original niche concepts and use this model to predict patterns of species distribution. The model may help to clarify how dispersal, niche size and competition interact, and under what conditions species might be common in unsuitable habitat or absent from suitable habitat. A brief review of the pertinent literature suggests that species are often absent from suitable habitat and present in unsuitable habitat, in ways predicted by theory. However, most tests of niche theory are hampered by inadequate consideration of what does and does not constitute suitable habitat. More conclusive evidence for these predictions will require rigorous determination of habitat suitability under field conditions. I suggest that to do this, ecologists must measure habitat specific demography and quantify how demographic parameters vary in response to temporal and spatial variation in measurable niche dimensions.     

Niche theory: New insights from an old paradigm

We apply a quantification of contemporary niche theory to several related topics of current interest: competition for alternative resources, the effect of consumers on those resources, and contemporary and evolutionary patterns at the community level attributable to the sequential colonization of resources and their consumers. Although the first topic is the traditional purview of niche theory, we believe that our model brings several issues into sharper focus than heretofore: The breadths of species' niches, overlap between them and the resulting interspecific effects all depend on the strategies by which competitors exploit heterogeneous environments. For our second application, we allow the alternative resources to be animate, and discover a positive relationship between predation and prey species diversity. Because we expect such a relationship for reasons that differ from those motivating the disturbance hypothesis, these ideas are easily distinguished. Finally, we consider the development of communities whose constituent populations behave according to the tenets of niche theory. Because successive trophic levels are colonized sequentially, patterns resulting from resource competition are replaced by those resulting from predation on consumers. For instance, consumer species-abundance distributions, which are uneven initially, become increasingly even during succession. Similar processes occur in evolutionary time, but the results have been reported as individual taxon cycles of invasion, accommodation and extinction or reinvasion. Niche theory suggests strategies and efficiencies of resource use that would facilitate invasion and prevent extinction. Though these last three are novel applications, the potential of this theory far exceeds them all. Moreover, the resulting insights augment the theory itself, increasing its general applicability. As a consequence of such feedback, niche theory may yet become a unifying concept in population biology.     

Isodars unveil asymmetric effects on habitat use caused by competition between two endangered species

In order for competing species to coexist, segregation on some ecological niche component is required and is often mediated by differential habitat use. When unequal competitors are involved, the dominant species tends to displace the subordinate one to its less preferred habitat. Here, we use habitat isodars, an approach which reflects evolutionary stable strategies of habitat selection, to evaluate whether interspecific competition between two competing species with distinct habitat preferences, the little bustard Tetrax tetrax and the great bustard Otis tarda, modulates their habitat use. Field data on these endangered species demonstrate that unequal competitors can coexist without completely segregating on their preferred habitats. The negatively sloped isodar of the subordinate little bustard unveils its competition with the dominant great bustard. Interference from great bustards in secondary cereal habitats reinforces use of preferred natural habitat by little bustards. Studies of density‐dependent habitat selection by a single‐species can thus aid in identifying the effects of competition on community composition, and guide the conservation of at‐risk species. Isodars, in particular, represent a promising method to gain clear knowledge on interspecific competition for species in which experimental manipulations are not feasible.     

Evolutionarily stable dispersal of a waterstrider in a temporally and spatially heterogeneous environment

Summary Evolutionary stable dispersal and wing muscle histolysis strategies are studied in the waterstriderGerris thoracicus. These strategies relate to spreading reproductive risk. Overwintering individuals have the choice of dispersing to either a brackish sea bay or a rock pool habitat. The former is reproductively more favorable than the latter during warm dry years and less favorable during cool wet years. After spring migration, individuals may histolyse their flight muscles and lay all their eggs in one pool or they may retain their flight ability and lay fewer eggs in total but spread them in several pools. We use a simple two-habitat model to examine the question of habitat dispersal. Our results indicate that, although the value of the evolutionary stable dispersal depends on the degree of variability in the environment and on the probability of local extinctions in either habitat, the population always disperses to both habitats as a consequence of density dependent growth. We use a more detailed multiple-rockpool habitat model to examine the question of wing muscle histolysis as a response to density dependence. Our results indicate that a wing muscle histolysis response to population density is an evolutionarily stable strategy when compared with the two alternatives of females always histolysing or never histolysing their flight muscles. The application of evolutionarily stable theory to stochastic problems presents a number of difficulties. We discuss these difficulties in the context of computing evolutionarily stable strategies for the problems at hand.     

Two gerbils of the Negev: A long-term investigation of optimal habitat selection and its consequences

Optimal foraging theory has entered a new phase. It is not so much tested as used. It helps behavioural ecologists discover the nature of the information in an animals brain. It helps population ecologists reveal coefficients of interaction and their patterns of density-dependent variation. And it helps community ecologists examine niche relationships. In our studies on two species of Negev desert gerbil, we have taken advantage of the second and third of these functions. Both these gerbils prefer semi-stabilized dune habitat, and both altered their selective use of this habitat and stabilized sand according to experimental changes we made in their populations. Their changes in selectivity agree with a type of optimal foraging theory called isoleg theory. Isoleg theories provide examples of dipswitch theories – bundles of articulated qualitative predictions – that are easier to falsify than single qualitative predictions. By linking behaviour to population dynamics through isoleg theory, we were able to use the behaviour of the gerbils to reveal the shapes of their competitive isoclines. These have the peculiar non-linear shapes predicted by optimal foraging theory. Finally, when owl predation threatens, the behaviour of Gerbillus allenbyi reveals the shape of their victim isocline. As has long been predicted by predation theory and laboratory experiments, it is unimodal.     

Disturbance regimes and life-history evolution

Disturbance regimes are ecologically important, but many of their evolutionary consequences are poorly understood. A model is developed here that combines the within- and among-season dynamics of disturbances with evolutionary life-history theory. "Disturbance regime" is defined in terms of disturbance timing, frequency, predictability, and severity. The model predicts the optimal body size and time at which organisms should abandon a disturbance-prone growth habitat by maturing and moving to a disturbance-free, nongrowth habitat. The effects of both coarse-grained (those affecting the entire population synchronously) and fine-grained disturbances (those occurring in a patch dynamics setting) are explored. Several predictions are congruent with previous theory. Infrequent or temporally unpredictable disturbances should have little effect on the evolution of life-history strategies, even though they may cause high mortality. Similar to seasonal time constraints on reproduction, disturbance regimes can synchronize metamorphosis within a population, resulting in a seasonal decline in body size at maturity. Other model predictions are novel. When disturbances cause high mortality, coarse-grained disturbances have a much stronger effect on life-history strategies than fine-grained disturbances, suggesting that population structure (relative to the scale of disturbance) plays a critical evolutionary role when disturbances are severe. When within-population variance in juvenile body size is high, two consecutive seasonal declines in body size at maturity can occur, the first associated with disturbance regime and the second associated with seasonal time constraints.     

The functional trophic role of lotic primary consumers: generalist versus specialist strategies

1. The relative extent of generalist or specialist resource use strategies is an important question in ecology. A community dominated by specialist strategies suggests a high level of interspecific competition for resources, resulting in the evolutionary development of isolating mechanisms between species (e.g. resource specialization to avoid and/or outcompete other species). A community dominated by generalist strategies suggests less interspecific competition for resources, allowing many taxa to utilize the same resources. In stream systems, generalist food habits are a common strategy among primary consumers, but little is known about resource assimilation strategies (resources incorporated into tissue growth). Published data indicate that generalist resource assimilation strategies may prevail in lotic systems as well.
2. Functional feeding groups (FFGs) are often used to infer resource assimilation among lotic macroinvertebrates (e.g. shredder-detritivore, scraper-herbivore). While these groupings are aptly used to describe invertebrate feeding modes and community structure, the use of FFGs to describe resource assimilation among lotic consumers is not appropriate. Sufficient data now exist to seriously question how accurately FFG assignments describe the processes of energy flow and material transfer between trophic levels in stream ecosystems.
3. Because FFGs may not accurately describe functional attributes in lotic systems, an alternative approach is needed. One approach is to determine the amount of secondary production that is derived from autochthonous (e.g. periphyton and algae) and allochthonous (e.g. detritus) resources directly. A simple model of community function based on this approach is presented. The model incorporates trophic generalists into measurement of consumer–resource energetics in lotic systems.     

Niche construction in the light of niche theory

Ecological niche construction, the process whereby an organism improves its environment to enhance its growth and persistence, is an important missing element of niche theory. Niche theory has mainly focused on niche-deteriorating processes, such as resource consumption, predation and competition, which have negative effects on population growth. Here, we integrate niche construction explicitly into modern niche theory. We use a graphical approach to analyse how a species' niche-improving impacts interplay with niche-deteriorating impacts to modify its response to the environment. In a model of two consumers that compete for one limiting resource and one predator, we show how niche construction modifies the traditional niche-deteriorating impacts of its agent or of competing species, and hence the potential for species coexistence. By altering the balance between intraspecific and interspecific competitive effects, niche construction can either generate net interspecific facilitation or strengthen interspecific competition. The adaptive benefit derived from niche construction also strongly affects the realized niche of a niche-constructing species.     

Competition on productivity gradients – what do we expect?

Many experimental studies have quantified how the effects of competition vary with habitat productivity, with the results often interpreted in terms of the ideas of Grime and Tilman. Unfortunately, these ideas are not relevant to many experiments, and so we develop an appropriate resource competition model and use this to explore the effects of habitat productivity on the intensity of competition. Several mechanisms influencing the productivity–competition intensity relationship are identified, and these mechanisms explored using two classic data sets. In both cases, there is good agreement between the model predictions and empirical patterns. Quantification of the mechanisms identified by the models will allow the development of a simple predictive theory linking measures of the intensity of competition with ecosystem‐level properties.     

Dispersal, migration, and offspring retention in saturated habitats

We examine the evolutionary stability of year-round residency in territorial populations, where breeding sites are a limiting resource. The model links individual life histories to the population-wide competition for territories and includes spatial variation in habitat quality as well as a potential parent-offspring conflict over territory ownership. The general form of the model makes it applicable to the evolution of dispersal, migration, partial migration, and delayed dispersal (offspring retention). We show that migration can be evolutionarily stable only if year-round residency in a given area would produce a sink population, where mortality exceeds reproduction. If this applies to a fraction of the breeding habitat only, partial migration is expected to evolve. In the context of delayed dispersal, habitat saturation has been argued to form an ecological constraint on independent breeding, which favors offspring retention and cooperative breeding. We show that habitat saturation must be considered as a dynamic outcome of birth, death, and dispersal rates in the population, rather than an externally determined constraint. Although delayed dispersal often associates with intense competition for territories, life-history traits have direct effects on stable dispersal strategies, which can often override the effect of habitat saturation. As an example, high survival of floaters selects against delayed dispersal, even though it increases the number of competitors for each breeding vacancy (the "habitat saturation factor"). High survival of territory owners, by contrast, generally favors natal philopatry. We also conclude that spatial variation in habitat quality only rarely selects for delayed dispersal. Within a population, however, offspring retention is more likely in high-quality territories.     

The functional trophic role of lotic primary consumers: generalist versus specialist strategies

1. The relative extent of generalist or specialist resource use strategies is an important question in ecology. A community dominated by specialist strategies suggests a high level of interspecific competition for resources, resulting in the evolutionary development of isolating mechanisms between species (e.g. resource specialization to avoid and/or outcompete other species). A community dominated by generalist strategies suggests less interspecific competition for resources, allowing many taxa to utilize the same resources. In stream systems, generalist food habits are a common strategy among primary consumers, but little is known about resource assimilation strategies (resources incorporated into tissue growth). Published data indicate that generalist resource assimilation strategies may prevail in lotic systems as well.
2. Functional feeding groups (FFGs) are often used to infer resource assimilation among lotic macroinvertebrates (e.g. shredder-detritivore, scraper-herbivore). While these groupings are aptly used to describe invertebrate feeding modes and community structure, the use of FFGs to describe resource assimilation among lotic consumers is not appropriate. Sufficient data now exist to seriously question how accurately FFG assignments describe the processes of energy flow and material transfer between trophic levels in stream ecosystems.
3. Because FFGs may not accurately describe functional attributes in lotic systems, an alternative approach is needed. One approach is to determine the amount of secondary production that is derived from autochthonous (e.g. periphyton and algae) and allochthonous (e.g. detritus) resources directly. A simple model of community function based on this approach is presented. The model incorporates trophic generalists into measurement of consumer–resource energetics in lotic systems.     

EVOLUTIONARY RESPONSES TO ENVIRONMENTAL CHANGES: HOW DOES COMPETITION AFFECT ADAPTATION?

The role and importance of ecological interactions for evolutionary responses to environmental changes is to large extent unknown. Here it is shown that interspecific competition may slow down rates of adaptation substantially and fundamentally change patterns of adaptation to long-term environmental changes. In the model investigated here, species compete for resources distributed along an ecological niche space. Environmental change is represented by a slowly moving resource maximum and evolutionary responses of single species are compared with responses of coalitions of two and three competing species. In scenarios with two and three species, species that are favored by increasing resource availability increase in equilibrium population size whereas disfavored species decline in size. Increased competition makes it less favorable for individuals of a disfavored species to occupy a niche close to the maximum and reduces the selection pressure for tracking the moving resource distribution. Individual-based simulations and an analysis using adaptive dynamics show that the combination of weaker selection pressure and reduced population size reduces the evolutionary rate of the disfavored species considerably. If the resource landscape moves stochastically, weak evolutionary responses cause large fluctuations in population size and thereby large extinction risk for competing species, whereas a single species subject to the same environmental variability may track the resource maximum closely and maintain a much more stable population size. Other studies have shown that competitive interactions may amplify changes in mean population sizes due to environmental changes and thereby increase extinction risks. This study accentuates the harmful role of competitive interactions by illustrating that they may also decrease rates of adaptation. The slowdown in evolutionary rates caused by competition may also contribute to explain low rates of morphological change in spite of large environmental fluctuations found in fossil records.     

Recruitment pulses induce cannibalistic giants in Arctic char

1. Recent theoretical studies on the population dynamic consequences of cannibalism have focused on mechanisms behind the emergence of large cannibals (giants) in size-structured populations. Theoretically, giants emerge when a strong recruiting cohort imposes competition induced mortality on stunted adults, but also provides a profitable resource for a few adults that accelerate in growth and reach giant sizes. 2. Here the effects of a recruitment pulse on the individual and population level in an allopatric Arctic char population have been studied over a 5-year period and these results were contrasted with theoretical model predictions for the conditions necessary for the emergence of cannibalistic giants. 3. The recruitment pulse had negative effects on invertebrate resource abundance, and the decrease in body condition and increase in mortality of adult char suggested that strong intercohort competition took place. 4. The frequency of cannibalism increased and a few char accelerated in growth and reached 'giant' sizes. 5. The main discrepancy between model predictions and field data was the apparently small effect the recruited cohort had on resources and adult char performance during their first summer. Instead, the effects became pronounced when the cohort was 1 year old. This mismatch between model predictions and field observations was suggested to be due to the low per capita fecundity in char and the restricted nearshore habitat use in young-of-the-year (YOY) char. 6. This study provides empirical evidence that the emergence of giants is associated with the breakthrough of a strong recruiting cohort and also suggests that the claimed stable char populations with large cannibals may instead be populations with dynamic size structure that results in intermittent breakthroughs of recruitment pulses, providing the conditions necessary for char to enter the cannibalistic niche. 7. The data suggest that increased recruit survival through restricted habitat use may destabilize dynamics and cause the emergence of giants. However, they also suggest that this does not necessarily develop into populations with bi-modal size structure in populations with low per capita fecundity and size- and density-dependent habitat use of recruiting cohorts.     

Heterogeneity in Primary Productivity Influences Competitive Interactions between Red Deer and Alpine Chamois

Habitat heterogeneity can promote coexistence between herbivores of different body size limited to different extents by resource quantity and quality. Red deer (Cervus elaphus) are known as superior competitors to smaller species with similar diets. We compared competitive interactions and habitat use between red deer and Alpine chamois (Rupicapra rupicapra) in two adjacent valleys in a strictly protected area in the Central Alps. Red deer density was higher in the valley with higher primary productivity. Only here was horn growth in kid and yearling chamois (as a measure for body condition) negatively correlated with red deer population size, suggesting interspecific competition, and chamois selected meadows with steeper slopes and lower productivity than available on average. Conversely, red deer selected meadows of high productivity, particularly in the poorer area. As these were located mainly at lower elevations, this led to strong altitudinal segregation between the two species here. Local differences in interspecific competition thus coincided with differences in habitat preference and–segregation between areas. This suggests that spatial habitat and resource heterogeneity at the scale of adjacent valleys can provide competition refuges for competitively inferior mountain ungulates which differ from their superior competitor in their metabolic requirements.     

Effects of habitat quality and size on extinction in experimental populations

Stochastic population theory makes clear predictions about the effects of reproductive potential and carrying capacity on characteristic time-scales of extinction. At the same time, the effects of habitat size and quality on reproduction and regulation have been hotly debated. To trace the causal relationships among these factors, we looked at the effects of habitat size and quality on extinction time in experimental populations of Daphnia magna. Replicate model systems representative of a broad-spectrum consumer foraging on a continuously supplied resource were established under crossed treatments of habitat size (two levels) and habitat quality (three levels) and monitored until eventual extinction of all populations. Using statistically derived estimates of key parameters, we related experimental treatments to persistence time through their effect on carrying capacity and the population growth rate. We found that carrying capacity and the intrinsic rate of increase were each influenced similarly by habitat size and quality, and that carrying capacity and the intrinsic rate of increase were in turn both correlated with time to population extinction. We expected habitat quality to have a greater influence on extinction. However, owing to an unexpected effect of habitat size on reproductive potential, habitat size and quality were similarly important for population persistence. These results support the idea that improving the population growth rate or carrying capacity will reduce extinction risk and demonstrate that both are possible by improving habitat quality or increasing habitat size.     

Competition and evolutionary stability of plants in a spatially structured habitat

We modelled the population dynamics of two types of plants with limited dispersal living in a lattice structured habitat. Each site of the square lattice model was either occupied by an individual or vacant. Each individual reproduced to its neighbors. We derived a criterion for the invasion of a rare type into a population composed of a resident type based on a pair-approximation method, in which the dynamics of both average densities and the nearest neighbor correlations were considered. Based on this invasibility criterion, we showed that, when there is a tradeoff between birth and death rates, the evolutionarily stable type is the one that has the highest ratio of birth rate to mortality. If these types are different species, they form segregated spatial patterns in the lattice model in which intraspecific competitive interactions occur more frequently than interspecific interactions. However, stable coexistence is not possible in the lattice model contrary to results from completely mixed population models. This clearly shows that the casual conclusion, based on traditional well mixed population models, that different species can coexist if intraspecific competition is stronger than interspecific competition, does not hold for spatially structured population models.