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.     

外来植物成功入侵的生物学特征

外来植物的入侵能力与其性状之间的关系是入侵生态学中的基本问题之一.成功的入侵种常常能占据多样化的生境,具有较强的适应性、繁殖力和散布力.表型可塑性和遗传分化是外来入侵植物对生境异质性的两种适应策略;散布体多态型和散布途径多种化,使外来入侵植物迅速占领入侵生境,并进行远距离扩散;无性生殖和有性生殖并存,并根据生境和入侵阶段权衡的繁育对策不仅使入侵种群大面积暴发成为可能,而且直接影响散布机制并对种群遗传结构具有调节作用.高效的资源利用性竞争,加之以化感作用为基础的干扰性竞争使植物更具入侵性.     

不同栖息地状态下物种竞争模式及模拟研究与应用

物种竞争是影响生态系统演化的重要生态过程之一.而物种在受人类影响出现不同程度毁坏的栖息地上的演化又是非常复杂的,因此研究物种演化对栖息地毁坏的响应是非常必要的.在Tilman研究工作的基础上,将竞争系数引入集合种群动力模式,建立了多物种集合种群竞争共存的数学模型,并对5-物种集合种群在不同栖息地状态下的竞争动态进行了计算机模拟研究.结果表明：（1）不同结构的群落（q值不同）,物种之间的竞争排斥作用强度不同,优势物种明显的群落,物种之间的排斥强度大;（2）随着栖息地毁坏程度的增加,对优势物种的负面影响逐渐减小,而对弱势物种的负面影响逐渐增加;（3）随着栖息地恢复幅度的增加,优势物种和弱势物种之间的竞争越强烈,优势物种受到的竞争排斥加大,而弱势物种逐渐变强,出现了强者变弱、弱者变强的格局;（4）物种竞争排斥与共存受迁移扩散能力和竞争能力影响很大,竞争共存的条件是其竞争能力与扩散能力呈非线性负相关关系;（5）竞争共存的物种的强弱序列发生了变化.     

Evolution of resident bird breeding phenology in a landscape with heterogeneous resource phenology and carryover effects

It is generally expected that, in environments with pronounced seasonal resource peaks, birds’ reproductive success will be maximised when nestlings’ peak food demand coincides with the timing of high food availability. However in certain birds that stay resident over winter, earlier breeding leads juveniles to join the winter flock earlier, which by the prior residence effect increases their success in breeding territory competition. This trade-off between reproduction and competition may explain why, in certain species, breeding phenology is earlier and asynchronous with the resource. This study extends a previous model of the evolution of breeding phenology in a single habitat type to a landscape with two habitat types: ‘early’ and ‘late’ resource phenology. The offspring’s natal habitat type has a carryover effect upon their competitive ability regardless of which habitat type they settle in to potentially breed. We find that, when the difference in resource phenology between habitats is small (weak carryover effect), breeding phenology in the late habitat evolves to occur earlier and more asynchronously than in the early habitat, to compensate for the competitive disadvantage to juveniles raised there. However if the difference is large (strong carryover effect), then the reproductive cost of earlier breeding outweighs the benefit of the compensation, so instead breeding phenology in the late habitat evolves to become more synchronous with the resource. Recruitment is generally asymmetric, from early to late habitat type. However if the early habitat is less frequent in the landscape or produces fewer offspring, then the asymmetry is reduced, and if there is some natal habitat-type fidelity, then recruitment can have an insular pattern, i.e. most recruits to each habitat type come from that same habitat type. We detail the different scenarios in which the different recruitment patterns are predicted, and we propose that they have implications for local adaptation.     

Exact compensation of stream drift as an evolutionarily stable strategy

The colonization cycle hypothesis predicts that adults of stream-dwelling insects preferentially disperse in the upstream direction in order to compensate for larval drift. Upstream biased dispersal has indeed been shown in many, albeit not all, natural populations. Based on a recently published analysis, we develop a simple stochastic model for the competition of genotypes with different dispersal strategies in a stream habitat. By means of an invasion analysis, we show that exact compensation of larval drift by upstream biased adult dispersal is an evolutionarily stable strategy. Exact compensation means that, on average, the net movement of individuals from birth to the time of reproduction is zero. At the population level, we show that, in general, upstream biased dispersal is not necessary for persistence, unless the reproductive rate is very low. Under all conditions, however, populations of exact compensators attain highest sizes or persistence times, respectively. Although selection pressure towards exact compensation is arguably very general in populations subject to stream drift, trade-offs or constraints might change the outcome of selection. Therefore, the analysis presented in this paper has to be viewed as a null model for optimal dispersal behavior in stream habitats.     

Rapid changes in phenotype distribution during range expansion in a migratory bird

The capacity of species to track changing environmental conditions is a key component of population and range changes in response to environmental change. High levels of local adaptation may constrain expansion into new locations, while the relative fitness of dispersing individuals will influence subsequent population growth. However, opportunities to explore such processes are rare, particularly at scales relevant to species-based conservation strategies. Icelandic black-tailed godwits, Limosa limosa islandica, have expanded their range throughout Iceland over the last century. We show that current male morphology varies strongly in relation to the timing of colonization across Iceland, with small males being absent from recently occupied areas. Smaller males are also proportionately more abundant on habitats and sites with higher breeding success and relative abundance of females. This population-wide spatial structuring of male morphology is most likely to result from female preferences for small males and better-quality habitats increasing both small-male fitness and the dispersal probability of larger males into poorer-quality habitats. Such eco-evolutionary feedbacks may be a key driver of rates of population growth and range expansion and contraction.     

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.     

Sex-biased dispersal and adaptation to marginal habitats

If gene flow occurs through both sexes but only females contribute to population growth, adaptation to marginal (sink) habitats should be differentially affected by male versus female dispersal. Here I address this problem with two models. First, I consider the fate of a rare allele that improves fitness in the marginal habitat but reduces fitness in the core (source) habitat. Then I study the evolution of a polygenic character mediating a trade-off in fitness between the habitats. Both approaches led to qualitatively similar predictions. The effect of a difference in the dispersal rate between the sexes depends on the degree to which immigration from the core habitat boosts the reproductive output from the marginal habitat. This boost is slight if the marginal habitat is able to sustain well a population without immigration. In that case, both female- and male-biased dispersal is more favorable for adaptation to marginal habitats than equal dispersal of both sexes (assuming that the dispersal rate averaged over the sexes is kept constant). In contrast, if the marginal habitat is an absolute sink unable to sustain a population without immigration, the conditions for adaptation to that habitat are least favorable under highly male-biased dispersal and most favorable under highly female-biased dispersal. Under some circumstances, high average (male+female) dispersal is more favorable than low dispersal. Thus, gene flow should not be seen solely as thwarting adaptation to marginal habitats. The results are interpreted in terms of how male and female dispersal affects the relative rate of gene flow from the source to the sink habitat and in the opposite direction. This study predicts that ecological niches of taxa with female-biased dispersal should tend to be broader and more evolutionarily flexible.     

Fine-scale adaptation in a clonal sea anemone

Local adaptation in response to fine-scale spatial heterogeneity is well documented in terrestrial ecosystems. In contrast, in marine environments local adaptation has rarely been documented or rigorously explored. This may reflect real or anticipated effects of genetic homogenization, resulting from widespread dispersal in the sea. However, evolutionary theory predicts that for the many benthic species with complex life histories that include both sexual and asexual phases, each parental habitat patch should become dominated by the fittest and most competitive clones. In this study we used genotypic mapping to show that within headlands, clones of the sea anemone Actinia tenebrosa show restricted distributions to specific habitats despite the potential for more widespread dispersal. On these same shores we used reciprocal transplant experiments that revealed strikingly better performance of clones within their natal rather than foreign habitats as judged by survivorship, asexual fecundity, and growth. These findings highlight the importance of selection for fine-scale environmental adaptation in marine taxa and imply that the genotypic structure of populations reflects extensive periods of interclonal competition and site-specific selection.     

Matching habitat choice promotes species persistence under climate change

Species may survive under contemporary climate change by either shifting their range or adapting locally to the warmer conditions. Theoretical and empirical studies recently underlined that dispersal, the central mechanism behind these responses, may depend on the match between an individuals’ phenotype and local environment. Such matching habitat choice is expected to induce an adaptive gene flow, but it now remains to be studied whether this local process could promote species’ responses to climate change. Here, we investigate this by developing an individual‐based model including either random dispersal or temperature‐dependent matching habitat choice. We monitored population composition and distribution through space and time under climate change. Relative to random dispersal, matching habitat choice induced an adaptive gene flow that lessened spatial range loss during climate warming by improving populations’ viability within the range (i.e. limiting range fragmentation) and by facilitating colonization of new habitats at the cold margin. The model even predicted range contraction under random dispersal but range expansion under optimal matching habitat choice. These benefits of matching habitat choice for population persistence mostly resulted from adaptive immigration decision and were greater for populations with larger dispersal distance and higher emigration probability. We also found that environmental stochasticity resulted in suboptimal matching habitat choice, decreasing the benefits of this dispersal mode under climate change. However population persistence was still better under suboptimal matching habitat choice than under random dispersal. Our results highlight the urgent need to implement more realistic mechanisms of dispersal such as matching habitat choice into models predicting the impacts of ongoing climate change on biodiversity.     

Effects of territory competition and climate change on timing of arrival to breeding grounds: a game-theory approach

Phenology is an important part of life history that is gaining increased attention because of recent climate change. We use game theory to model phenological adaptation in migratory birds that compete for territories at their breeding grounds. We investigate how the evolutionarily stable strategy (ESS) for the timing of arrival is affected by changes in the onset of spring, the timing of the resource peak, and the season length. We compare the ESS mean arrival date with the environmental optimum, that is, the mean arrival date that maximizes fitness in the absence of competition. When competition is strong, the ESS mean arrival date responds less than the environmental optimum to shifts in the resource peak but more to changes in the onset of spring. Increased season length may not necessarily affect the environmental optimum but can still advance the ESS mean arrival date. Conversely, shifting a narrow resource distribution may change the environmental optimum without affecting the ESS mean arrival date. The ESS mean arrival date and the environmental optimum may even shift in different directions. Hence, treating phenology as an evolutionary game rather than an optimization problem fundamentally changes what we predict to be an adaptive response to environmental changes.     

Eco-evolutionary consequences of habitat warming and fragmentation in communities

Eco-evolutionary dynamics can mediate species and community responses to habitat warming and fragmentation, two of the largest threats to biodiversity and ecosystems. The eco-evolutionary consequences of warming and fragmentation are typically studied independently, hindering our understanding of their simultaneous impacts. Here, we provide a new perspective rooted in trade-offs among traits for understanding their eco-evolutionary consequences. On the one hand, temperature influences traits related to metabolism, such as resource acquisition and activity levels. Such traits are also likely to have trade-offs with other energetically costly traits, like antipredator defences or dispersal. On the other hand, fragmentation can influence a variety of traits (e.g. dispersal) through its effects on the spatial environment experienced by individuals, as well as properties of populations, such as genetic structure. The combined effects of warming and fragmentation on communities should thus reflect their collective impact on traits of individuals and populations, as well as trade-offs at multiple trophic levels, leading to unexpected dynamics when effects are not additive and when evolutionary responses modulate them. Here, we provide a road map to navigate this complexity. First, we review single-species responses to warming and fragmentation. Second, we focus on consumer–resource interactions, considering how eco-evolutionary dynamics can arise in response to warming, fragmentation, and their interaction. Third, we illustrate our perspective with several example scenarios in which trait trade-offs could result in significant eco-evolutionary dynamics. Specifically, we consider the possible eco-evolutionary consequences of (i) evolution in thermal performance of a species involved in a consumer–resource interaction, (ii) ecological or evolutionary changes to encounter and attack rates of consumers, and (iii) changes to top consumer body size in tri-trophic food chains. In these scenarios, we present a number of novel, sometimes counter-intuitive, potential outcomes. Some of these expectations contrast with those solely based on ecological dynamics, for example, evolutionary responses in unexpected directions for resource species or unanticipated population declines in top consumers. Finally, we identify several unanswered questions about the conditions most likely to yield strong eco-evolutionary dynamics, how better to incorporate the role of trade-offs among traits, and the role of eco-evolutionary dynamics in governing responses to warming in fragmented communities.     

Impacts of dispersal on rapid adaptation and dynamic stability of Daphnia in fluctuating environments

Prior ecological research has shown that spatial processes can enhance the temporal stability of populations in fluctuating environments. Less explored is the effect of dispersal on rapid adaptation and its concomitant impact on population dynamics. For asexually reproducing populations, theory predicts that dispersal in fluctuating environments can facilitate asynchrony among clones and enhance stability by reducing temporal variability of total population abundance. This effect is predicted when clones exhibit heritable variation in environmental optima and when fluctuations occur asynchronously among patches. We tested this in the field using artificial ponds and metapopulations composed of a diverse assemblage of Daphnia pulex clones. We directly manipulated dispersal presence/absence and environmental fluctuations in the form of nutrient pulses. Consistent with predictions, dispersal enhanced temporal asynchrony among clones in the presence of nutrient pulses; this in turn stabilized population dynamics. This effect only emerged when patches experienced spatially asynchronous nutrient pulses (dispersal had no effect when patches were synchronously pulsed). Clonal asynchrony was driven by strong positive selection for a single clone that exhibited a performance advantage under conditions of low resource availability. Our work highlights the importance of dispersal as a driver of eco-evolutionary dynamics and population stability in variable environments.     

Predicting range shifts under global change: the balance between local adaptation and dispersal

Bioclimate envelope models (BEMs) have often been criticized as being too simplistic due to e.g. not incorporating effects of biotic interactions or evolutionary adaptation. However, BEMs are widely applied and have proven to be often useful. Here we investigate, under which conditions evolution of dispersal, local adaptation or interspecific competition may be of minor importance for forecasting future range shifts. Therefore we use individual‐based simulations of metapopulations under climate change living in spatial temperature gradients. Scenarios incorporate single‐species systems or systems with competing species, respectively. Dispersal rate is evolving and adaptation to local conditions may also evolve in some scenarios. Results show that in single‐species scenarios excluding evolutionary adaptation, species either follow optimal habitat conditions or go extinct if habitat connectivity is too low. These simulations are in close accordance to predictions from BEMs. Including evolutionary adaptation qualitatively changes these results. In the absence of competing species the species either completely invades the world or goes extinct. With competitors, results strongly depend on habitat fragmentation. For highly connected habitats the range border may shift as predicted by BEMs, for intermediate connectivity it will lag behind, while species will go extinct if fragmentation is too high. Our results indicate that (simple) BEMs may work well if habitats are well connected and species will not encounter many difficulties in dispersing to new sites. Selection in this case may promote evolution of even higher dispersal activities. We thus show that the presence of biotic interactions may be ignored for predictions of range shifts when high dispersal can be expected.     

The effects of migration load,selfing, inbreeding depression,and the genetics of adaptation on autotetraploid versus diploid establishment in peripheral habitats

The distribution and abundance of polyploids has intrigued biologists since their discovery in the early 20th century. A pattern in nature that may give insight to processes that shape the distribution and abundance of polyploids is that polyploid populations are sometimes associated with peripheral habitats within the range of a species of mixed ploidy. Here, adaptation and competition of a diploid versus an autotetraploid population in a peripheral habitat are examined theoretically. It is shown that a nascent autotetraploid population adapts to and outcompetes a diploid population in the periphery when the rate of gamete dispersal is high, and when the mode of gene action is recessive for moderate to high rates of selfing. With additive or dominant modes of gene action, the conditions for an autotetraploid to outcompete a diploid in the periphery appear determined more by the rate of selfing and less by gamete dispersal. All of these results are based on empirical work that suggests inbreeding depression is higher in diploids versus autotetraploids. Generally, the results indicate that, although autotetraploids incur minority cytotype exclusion, diploids face burdens themselves. In the case of adaptation to a peripheral habitat, this burden is migration load from gamete and propagule dispersal.     

Ecological genetics of Bromus tectorum

Summary By incorporating demographic analyses of fitness components (e.g., survival and reproduction) within a reciprocal sowing design, we tested for 3 consecutive years whether local adaptation has occurred in the alien grass Bromus tectorum (cheatgrass) within 7 habitats along an environmental gradient from arid steppe to subalpine forest in the Intermontain Region of western North America. Patterns of emergence and survival were strongly influenced by the local environment. In terms of survival, expression of significant local adaptation in Tsuga heterophylla habitat varied among years. In contrast, relative differences in flowering time among seed sources were stable across sites and years. Populations from the arid steppe were the earliest to flower; flowering was latest in populations from the mesic Tsuga heterophylla habitat. In terms of net reproductive rate, evidence for local adaptation in B. tectorum was obtained in populations from habitats representing environmental extremes: an arid, saline site dominated by the shrub Sarcobatus vermiculatus and clearings within the cool, mesic Tsuga heterophylla forest habitat. Unlike the plants introduced from other sites, members of the resident population at the Sarcobatus site flowered and produced seeds before soil water became limiting. In contrast, net reproductive rates in other habitats were sometimes the lowest for populations in their home site. This lack of an advantage for local populations within more environmentally moderate sites suggests that limited dispersal may restrict the rate at which superior genotypes are introduced into a particular site.     

The ideal free distribution: a review and synthesis of the game-theoretic perspective

The Ideal Free Distribution (IFD), introduced by Fretwell and Lucas in [Fretwell, D.S., Lucas, H.L., 1970. On territorial behavior and other factors influencing habitat distribution in birds. Acta Biotheoretica 19, 16-32] to predict how a single species will distribute itself among several patches, is often cited as an example of an evolutionarily stable strategy (ESS). By defining the strategies and payoffs for habitat selection, this article puts the IFD concept in a more general game-theoretic setting of the “habitat selection game”. Within this game-theoretic framework, the article focuses on recent progress in the following directions: (1) studying evolutionarily stable dispersal rates and corresponding dispersal dynamics; (2) extending the concept when population numbers are not fixed but undergo population dynamics; (3) generalizing the IFD to multiple species.For a single species, the article briefly reviews existing results. It also develops a new perspective for Parker’s matching principle, showing that this can be viewed as the IFD of the habitat selection game that models consumer behavior in several resource patches and analyzing complications involved when the model includes resource dynamics as well. For two species, the article first demonstrates that the connection between IFD and ESS is now more delicate by pointing out pitfalls that arise when applying several existing game-theoretic approaches to these habitat selection games. However, by providing a new detailed analysis of dispersal dynamics for predator-prey or competitive interactions in two habitats, it also pinpoints one approach that shows much promise in this general setting, the so-called “two-species ESS”. The consequences of this concept are shown to be related to recent studies of population dynamics combined with individual dispersal and are explored for more species or more patches.     

Dispersal: risk spreading versus local adaptation

I investigate how risk spreading in stochastic environments and adaptation to permanent properties of local habitats interplay in the simultaneous evolution of dispersal and habitat specialization. In a simple two-patch model, I find many types of locally evolutionarily stable attractors of dispersal and of a trait involved in habitat specialization, including a single habitat specialist and a coalition of two specialists with low dispersal, a generalist with high dispersal, and several types of dispersal polymorphisms. In general, only one attractor is a global evolutionarily stable strategy (ESS). In addition to the ESS analysis, I also present some examples of the dynamics of evolution that exhibit adaptive diversification by evolutionary branching.     

Game theory sheds new light on ecological responses to current climate change when phenology is historically mismatched

Phenological changes are well documented biological effects of current climate change but their adaptive value and demographic consequences are poorly known. Game theoretical models have shown that deviating from the fitness-maximising phenology can be evolutionary stable under frequency-dependent selection. We study eco-evolutionary responses to climate change when the historical phenology is mismatched in this way. For illustration we model adaptation of arrival dates in migratory birds that compete for territories at their breeding grounds. We simulate climate change by shifting the timing and the length of the favourable season for breeding. We show that initial trends in changes of population densities can be either reinforced or counteracted during the ensuing evolutionary adaptation. We find in total seven qualitatively different population trajectories during the transition to a new evolutionary equilibrium. This surprising diversity of eco-evolutionary responses provides adaptive explanations to the observed variation in phenological responses to recent climate change.     

Reproductive asynchrony increases with environmental disturbance

While it is widely recognized that the manner in which organisms adjust their timing of reproduction reflects evolutionary strategies aimed at minimizing offspring mortality or maximizing reproductive output, the conditions under which the evolutionarily stable strategy involves synchronous or asynchronous reproduction is a matter of considerable discord. A recent theoretical model predicts that whether a population displays reproductive synchrony or asynchrony will depend on the relative scales of intrinsic regulation and environmental disturbance experienced by reproducing individuals. This model predicts that, under conditions of negligible competition and large-scale environmental perturbation, evolution of a single mixed strategy will result in asynchronous reproduction. We tested this prediction using empirical data on large-scale climatic fluctuation and the annual timing of reproduction by three species of flowering plants covering 1300-population-years and four degrees of latitude in Norway. In agreement with model predictions, within populations of all three species reproductive asynchrony increased with the magnitude of large-scale climatic perturbation, but bore no relation to the strength of local density dependence. These results suggest that mixed evolutionarily stable strategies can arise from the interplay of combinations of agents of selection and the scale at which they operate; hence it is fruitless to associate synchronous versus asynchronous timing with particular single factors like climate, competition, or predation.