Population structure inhibits evolutionary diversification under competition for resources

A model is presented that explores how population structure affects the evolutionary outcome of ecological competition for resources. The model assumes that competition for resources occurs within groups of a finite number of individuals (interaction groups), and that limited dispersal of individuals between groups (according to Wright's island model of population structure) results in genetic structuring of the population. It is found that both finite-sized interaction groups and limited dispersal can have substantial effects on the evolution of resource exploitation strategies as compared to models with a single, infinitely large, well-mixed interaction group. Both effects, in general, tend to select for less aggressive competitive strategies. Moreover, both effects also tend to reduce the likelihood of the evolutionary diversification of resource exploitation strategies that often occurs in models of resource competition with infinite populations. The results are discussed in the context of theories of the evolutionary diversification of resource exploitation strategies and speciation.     

On the evolutionary dynamics of pathogens with direct and environmental transmission

A number of ecologically and economically important pathogens exhibit a complex transmission dynamics that involves distinct transmission modes. In this paper, we study the evolutionary dynamics of pathogens for which transmission includes direct host-to-host as well as indirect environmental transmission. Different routes of infection spread require specific adaptations of the parasite, which may result in conflicting selection pressures. Using the framework of Adaptive dynamics, we investigate how these conflicting selection pressures are resolved in the course of evolution and determine the conditions for evolutionary diversification of pathogen strains. We show that evolutionary branching and subsequent evolution of specialist strains occurs in wide parameter regions but evolutionary bistability and evolution of generalist pathogens are possible as well. Our analysis reveals that the relative contributions of direct and environmental transmission, as well as the underlying ecological dynamics, play a crucial role in shaping the course of pathogen evolution. Our findings may explain the coexistence of high and low virulence strains observed in several pathogenic organisms using different transmission modes (e.g., influenza viruses) and highlight the importance of considering ecological dynamics in virulence management.     

Evolution of condition-dependent dispersal under kin competition

Dispersers often differ in body condition from non-dispersers. The social dominance hypothesis explains dispersal of weak individuals, but it is not yet well understood why strong individuals, which could easily retain their natal site, are sometimes exposed to risky dispersal. Based on the model for dispersal under kin competition by Hamilton and May, we construct a model where dispersal propensity depends on body condition. We consider an annual species that inhabits a patchy environment with varying patch qualities. Offspring body condition corresponds to the quality of the natal patch and competitive ability increases with body condition. Our main general result balances the fitness benefit from not dispersing and retaining the natal patch and the benefit from dispersing and establishing somewhere else. We present four different examples for competition, which all hint that dispersal of strong individuals may be a common outcome under the assumptions of the present model. In three of the examples, the evolutionarily stable dispersal probability is an increasing function of body condition. However, we found an example where, counterintuitively, the evolutionarily stable dispersal probability is a non-monotone function of body condition such that both very weak and very strong individuals disperse with high probability but individuals of intermediate body condition do not disperse at all.     

Fitness-dependent dispersal in metapopulations and its consequences for persistence and synchrony

1. We present a novel metapopulation model where dispersal is fitness dependent: the strength of migration from a site is dependent on the expected reproductive fitness of individuals there. Furthermore, individuals continue to migrate until they reach a suitable habitat where their expected fitness is above a threshold value.
2. Fitness-dependent dispersal has a very strong stabilizing effect on population dynamics, even when the intrinsic dynamics of populations in the absence of dispersal exhibit complex high-amplitude oscillations. This stabilizing effect is much stronger than that of the density-independent dispersal normally considered in metapopulation models.
3. Even when fitness-dependent dispersal does not stabilize the dynamics in a formal sense, it generally leads to simplification, with complex or even chaotic fluctuations being reduced to simple cycles.
4. This form of dispersal also has a strong tendency to synchronize local population dynamics across the spatial extent of the metapopulation.
5. These conclusions are robust to the addition of strong stochasticity in the migration threshold.     

Protected polymorphism and evolutionary stability in pleiotropic models with trait-specific dominance

When alleles have pleiotropic effects on a number of quantitative traits, the degree of dominance between a pair of alleles can be different for each trait. Such trait-specific dominance has been studied previously in models for the maintenance of genetic variation by antagonistic effects of an allele on two fitness components. By generalizing these models to an arbitrary number of fitness components or other phenotypic traits with different degrees of dominance, I show that genetic polymorphism is generally impossible without antagonistic fitness effects of different traits and without trait-specific dominance. I also investigate dominance and pleiotropy from a more long-term evolutionary perspective, allowing for the study of general ecological scenarios, and I discuss the effects of trait-specific dominance on evolutionary stability criteria. When selection is mainly directional and only trait-specific dominance and antagonism cause the emergence of polymorphism, then these polymorphisms can be overtaken by single mutants again, such that they are probably short-lived on an evolutionary time scale. Near evolutionarily singular points where directional selection is absent, trait-specific dominance and overdominance facilitate the emergence of polymorphism and cause evolutionary divergence in some cases. An important outcome of these models is that trait-specific dominance allows for the emergence of genetic polymorphisms without a selective disadvantage for heterozygotes. This removes the scope for the evolution of assortative mate choice and affects dominance modification. Sympatric speciation by disruptive ecological selection requires this heterozygote disadvantage in order to evolve, and therefore it becomes less plausible if the emergence of genetic polymorphism usually occurs via trait-specific dominance and antagonistic effects.     

Variant evolutionary trees under phenotypic variance

Evolutionary branching, which is a coevolutionary phenomenon of the development of two or more distinctive traits from a single trait in a population, is the issue of recent studies on adaptive dynamics. In previous studies, it was revealed that trait variance is a minimum requirement for evolutionary branching, and that it does not play an important role in the formation of an evolutionary pattern of branching. Here we demonstrate that the trait evolution exhibits various evolutionary branching paths starting from an identical initial trait to different evolutional terminus traits as determined by only changing the assumption of trait variance. The key feature of this phenomenon is the topological configuration of equilibria and the initial point in the manifold of dimorphism from which dimorphic branches develop. This suggests that the existing monomorphic or polymorphic set in a population is not an unique inevitable consequence of an identical initial phenotype.     

On evolution under asymmetric competition

The evolutionary consequences of asymmetric competition between species are poorly understood in comparison with symmetric competition. A model for evolution of body size under asymmetric competition within and between species is described. The model links processes operating at the scale of the individual to that of macroscopic evolution through a stochastic mutation–selection process. Phase portraits of evolution in a phenotype space characteristically show character convergence and parallel character shifts, with character divergence being relatively uncommon. The asymptotic states of evolution depend very much on the properties of asymmetric competition. Given relatively weak asymmetries between species, a single equilibrium point exists; this is a local attractor, and its position is determined by the intra- and interspecific asymmetries. When the asymmetries are made stronger, several fixed points may come about, creating further equilibrium points which are local attractors. It is also possible for periodic attractors to occur; such attractors comprise Red Queen dynamics with phenotype values that continue to change without ever settling down to constant values. From certain initial conditions, evolution leading to extinction of one of the species is also a likely outcome.     

Long-term evolution of polygenic traits under frequency-dependent intraspecific competition

We analytically investigate the long-term evolution of a continuously varying quantitative character in a diploid population that is determined additively by a finite number of loci. The trait is under a mixture of frequency-dependent disruptive selection induced by intraspecific competition and frequency-independent stabilizing selection. Moreover, the trait is restricted to a finite range by constraints on the particular loci. Our investigations are based on explicit analytical results (provided by Bürger [2005. A multilocus analysis of intraspecific competition and stabilizing selection on a quantitative trait. J. Math. Biol. 50, 355-396]; Schneider [2006. A multilocus-multiallele analysis of frequency-dependent selection induced by intraspecific competition. J. Math. Biol. 52, 483-523]) on the short-term dynamics under the assumption of linkage equilibrium. We show that the population always reaches a long-term equilibrium (LTE), i.e., an equilibrium that is resistant against perturbations of mutations of sufficiently small effect. In general, several LTEs can coexist. They can be calculated explicitly, and we provide necessary and sufficient conditions for their existence. In the case that more than one LTE exists, we exemplify numerically that the evolutionary outcome depends crucially on the initial genetic architecture, on the joint distribution of mutational effects across loci, and on the particular realization of the mutation process. Therefore, long-term evolution cannot be predicted from the ecology alone. We further show that a partial order exists for the LTEs. The set of LTEs has a 'largest' element, an LTE which is reached during long-term evolution if the effects of the occurring mutant alleles are sufficiently large.     

Population dynamics and evolutionary processes: the manifold roles of habitat selection

Summary Any character that has a substantial effect on a species' distribution and abundance can exert a variety of indirect effects on evolutionary processes. It is suggested that an organism's capacity for habitat selection is just such a character. Habitat selection can constrain the selective environment experienced by a population. Habitat selection can also indirectly influence the relative importance of natural selection, drift, and gene flow, through its effect on population size and growth rate. In many circumstances (but not all), habitat selection increases population size and growth rate, and thereby makes selection in a local environment more effective than drift and gene flow.     

Ecological and evolutionary principles in immunology

Experimental immunology has given rise to detailed insights into how immune cells react to infectious agents and fight pathogens. At the same time, however, the interplay between infectious agents and immune responses can be viewed as an ecological system in vivo . This is characterized by complex interactions between species of immune cells and populations of pathogens. This review discusses how an understanding of the immune system can be aided by the application of ecological and evolutionary principles: competition, predation, and the evolution of viruses in vivo . These concepts can shed light onto important immunological concepts such as the correlates of efficient virus control, immunodominance, the relationship between viral evolution and the development of pathology, as well as the ability of the immune system to control immunosuppressive infections.     

Eco-evolutionary feedbacks,adaptive dynamics and evolutionary rescue theory

Adaptive dynamics theory has been devised to account for feedbacks between ecological and evolutionary processes. Doing so opens new dimensions to and raises new challenges about evolutionary rescue. Adaptive dynamics theory predicts that successive trait substitutions driven by eco-evolutionary feedbacks can gradually erode population size or growth rate, thus potentially raising the extinction risk. Even a single trait substitution can suffice to degrade population viability drastically at once and cause ‘evolutionary suicide’. In a changing environment, a population may track a viable evolutionary attractor that leads to evolutionary suicide, a phenomenon called ‘evolutionary trapping’. Evolutionary trapping and suicide are commonly observed in adaptive dynamics models in which the smooth variation of traits causes catastrophic changes in ecological state. In the face of trapping and suicide, evolutionary rescue requires that the population overcome evolutionary threats generated by the adaptive process itself. Evolutionary repellors play an important role in determining how variation in environmental conditions correlates with the occurrence of evolutionary trapping and suicide, and what evolutionary pathways rescue may follow. In contrast with standard predictions of evolutionary rescue theory, low genetic variation may attenuate the threat of evolutionary suicide and small population sizes may facilitate escape from evolutionary traps.     

Female competition and its evolutionary consequences in mammals

Following Darwin's original insights regarding sexual selection, studies of intrasexual competition have mainly focused on male competition for mates; by contrast, female reproductive competition has received less attention. Here, we review evidence that female mammals compete for both resources and mates in order to secure reproductive benefits. We describe how females compete for resources such as food, nest sites, and protection by means of dominance relationships, territoriality and inter‐group aggression, and by inhibiting the reproduction of other females. We also describe evidence that female mammals compete for mates and consider the ultimate causes of such behaviour, including competition for access to resources provided by mates, sperm limitation and prevention of future resource competition. Our review reveals female competition to be a potentially widespread and significant evolutionary selection pressure among mammals, particularly competition for resources among social species for which most evidence is currently available. We report that female competition is associated with many diverse adaptations, from overtly aggressive behaviour, weaponry, and conspicuous sexual signals to subtle and often complex social behaviour involving olfactory signalling, alliance formation, altruism and spite, and even cases where individuals appear to inhibit their own reproduction. Overall, despite some obvious parallels with male phenotypic traits favoured under sexual selection, it appears that fundamental differences in the reproductive strategies of the sexes (ultimately related to parental investment) commonly lead to contrasting competitive goals and adaptations. Because female adaptations for intrasexual competition are often less conspicuous than those of males, they are generally more challenging to study. In particular, since females often employ competitive strategies that directly influence not only the number but also the quality (survival and reproductive success) of their own offspring, as well as the relative reproductive success of others, a multigenerational view ideally is required to quantify the full extent of variation in female fitness resulting from intrasexual competition. Nonetheless, current evidence indicates that the reproductive success of female mammals can also be highly variable over shorter time scales, with significant reproductive skew related to competitive ability. Whether we choose to describe the outcome of female reproductive competition (competition for mates, for mates controlling resources, or for resources per se) as sexual selection depends on how sexual selection is defined. Considering sexual selection strictly as resulting from differential mating or fertilisation success, the role of female competition for the sperm of preferred (or competitively successful) males appears particularly worthy of more detailed investigation. Broader definitions of sexual selection have recently been proposed to encompass the impact on reproduction of competition for resources other than mates. Although the merits of such definitions are a matter of ongoing debate, our review highlights that understanding the evolutionary causes and consequences of female reproductive competition indeed requires a broader perspective than has traditionally been assumed. We conclude that future research in this field offers much exciting potential to address new and fundamentally important questions relating to social and mating‐system evolution.     

Evolution and persistence of obligate mutualists and exploiters: competition for partners and evolutionary immunization

Mutualisms are ubiquitous in nature, as is their exploitation by both conspecific and heterospecific cheaters. Yet, evolutionary theory predicts that cheating should be favoured by natural selection. Here, we show theoretically that asymmetrical competition for partners generally determines the evolutionary fate of obligate mutualisms facing exploitation by third-species invaders. When asymmetry in partner competition is relatively weak, mutualists may either exclude exploiters or coexist with them, in which case their co-evolutionary response to exploitation is usually benign. When asymmetry is strong, the mutualists evolve towards evolutionary attractors where they become extremely vulnerable to exploiter invasion. However, exploiter invasion at an early stage of the mutualism's history can deflect mutualists' co-evolutionary trajectories towards slightly different attractors that confer long-term stability against further exploitation. Thus, coexistence of mutualists and exploiters may often involve an historical effect whereby exploiters are co-opted early in mutualism history and provide lasting 'evolutionary immunization' against further invasion.     

Conditional dispersal under kin competition: extension of the Hamilton-May model to brood size-dependent dispersal

I consider a site-based model with contest competition among siblings, and assume that dispersal is conditional on the number of offspring in the natal site. Evolutionarily stable populations contain threshold dispersal strategies, which retain a certain number of offspring in the natal site and disperse the rest (if the actual number of offspring is less than the threshold, then all offspring are retained). Due to the discrete nature of the strategy set (the threshold must be integer), the ESS may not be unique or may not exist. In the latter case, two neighboring threshold strategies coexist in the evolutionarily stable population. Dispersal first decreases and then increases as a function of dispersal mortality, such that all but one offspring should be dispersed both when dispersal mortality is very small or very high. Population-level dispersal fractions are often similar to the unconditional ESS, but differ strongly when fecundity is small and dispersal mortality is high.     

Consequences of asymmetric competition between resident and invasive defoliators: A novel empirically based modelling approach

Invasive species can have profound effects on a resident community via indirect interactions among community members. While long periodic cycles in population dynamics can make the experimental observation of the indirect effects difficult, modelling the possible effects on an evolutionary time scale may provide the much needed information on the potential threats of the invasive species on the ecosystem. Using empirical data from a recent invasion in northernmost Fennoscandia, we applied adaptive dynamics theory and modelled the long term consequences of the invasion by the winter moth into the resident community. Specifically, we investigated the outcome of the observed short-term asymmetric preferences of generalist predators and specialist parasitoids on the long term population dynamics of the invasive winter moth and resident autumnal moth sharing these natural enemies. Our results indicate that coexistence after the invasion is possible. However, the outcome of the indirect interaction on the population dynamics of the moth species was variable and the dynamics might not be persistent on an evolutionary time scale. In addition, the indirect interactions between the two moth species via shared natural enemies were able to cause asynchrony in the population cycles corresponding to field observations from previous sympatric outbreak areas. Therefore, the invasion may cause drastic changes in the resident community, for example by prolonging outbreak periods of birch-feeding moths, increasing the average population densities of the moths or, alternatively, leading to extinction of the resident moth species or to equilibrium densities of the two, formerly cyclic, herbivores.     

Ecological and evolutionary consequences of selective interspecific information use

Recent work has shown that animals frequently use social information from individuals of their own species as well as from other species; however, the ecological and evolutionary consequences of this social information use remain poorly understood. Additionally, information users may be selective in their social information use, deciding from whom and how to use information, but this has been overlooked in an interspecific context. In particular, the intentional decision to reject a behaviour observed via social information has received less attention, although recent work has indicated its presence in various taxa. Based on existing literature, we explore in which circumstances selective interspecific information use may lead to different ecological and coevolutionary outcomes between two species, such as explaining observed co-occurrences of putative competitors. The initial ecological differences and the balance between the costs of competition and the benefits of social information use potentially determine whether selection may lead to trait divergence, convergence or coevolutionary arms race between two species. We propose that selective social information use, including adoption and rejection of behaviours, may have far-reaching fitness consequences, potentially leading to community-level eco-evolutionary outcomes. We argue that these consequences of selective interspecific information use may be much more widespread than has thus far been considered.     

Ecological and evolutionary consequences of niche construction for its agent

Niche construction can generate ecological and evolutionary feedbacks that have been underinvestigated so far. We present an eco-evolutionary model that incorporates the process of niche construction to reveal its effects on the ecology and evolution of the niche-constructing agent. We consider a simple plant-soil nutrient ecosystem in which plants have the ability to increase the input of inorganic nutrient as an example of positive niche construction. On an ecological time scale, the model shows that niche construction allows the persistence of plants under infertile soil conditions that would otherwise lead to their extinction. This expansion of plants' niche, however, requires a high enough rate of niche construction and a high enough initial plant biomass to fuel the positive ecological feedback between plants and their soil environment. On an evolutionary time scale, we consider that the rates of niche construction and nutrient uptake coevolve in plants while a trade-off constrains their values. Different evolutionary outcomes are possible depending on the shape of the trade-off. We show that niche construction results in an evolutionary feedback between plants and their soil environment such that plants partially regulate soil nutrient content. The direct benefit accruing to plants, however, plays a crucial role in the evolutionary advantage of niche construction.     

Joint evolution of dispersal and connectivity

Functional connectivity, the realized flow of individuals between the suitable sites of a heterogeneous landscape, is a prime determinant of the maintenance and evolution of populations in fragmented habitats. While a large body of literature examines the evolution of dispersal propensity, it is less known how evolution shapes functional connectivity via traits that influence the distribution of the dispersers. Here, we use a simple model to demonstrate that, in a heterogeneous environment with clustered and solitary sites (i.e., with variable structural connectivity), the evolutionarily stable population contains strains that are strongly differentiated in their pattern of connectivity (local vs. global dispersal), but not necessarily in the fraction of dispersed individuals. Also during evolutionary branching, selection is disruptive predominantly on the pattern of connectivity rather than on dispersal propensity itself. Our model predicts diversification along a hitherto neglected axis of dispersal strategies and highlights the role of the solitary sites—the more isolated and therefore seemingly less important patches of habitat—in maintaining global dispersal that keeps all sites connected.