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Population responses to environmental change depend on both the ecological interactions between species and the evolutionary responses of all species. In this study, we explore how evolution in prey, predators, or both species affect the responses of predator populations to a sustained increase in mortality. We use an eco-evolutionary predator–prey model to explore how evolution alters the predator extinction threshold (defined as the minimum mortality rate that prevents population growth at low predator densities) and predator hydra effects (increased predator abundance in response to increased mortality). Our analysis identifies how evolutionary responses of prey and predators individually affect the predator extinction threshold and hydra effects, and how those effects are altered by interactions between the evolutionary responses. Based on our theoretical results, we predict that it is common in natural systems for evolutionary responses in one or both species to allow predators to persist at higher mortality rates than would be possible in the absence of evolution (i.e., evolution increases the predator mortality extinction threshold). We also predict that evolution-driven hydra effects occur in a minority of natural systems, but are not rare. We revisited published eco-evolutionary models and found that evolution causes hydra effects and increases the predator extinction threshold in many studies, but those effects have been overlooked. We discuss the implications of these results for species conservation, predicting population responses to environmental change, and the possibility of evolutionary rescue.

Populations that experience severe stress may avoid extinction through adaptation by natural selection. This process is called evolutionary rescue and has been studied under different names in medicine, agriculture, and conservation biology. It is a component of the emerging field of eco-evolutionary dynamics, which investigates how the ecological attributes of species may evolve rapidly under strong selection. Its distinguishing feature is to combine the evolutionary concept of relative fitness with the ecological concept of absolute fitness in a synthetic theory of persistent adaptation. The likelihood of rescue will depend both on attributes of the population, particularly abundance and variation, and on properties of the environment, particularly the rate and severity of deterioration. Medical interventions (e.g., the administration of antibiotics), agricultural practices (e.g., the application of pesticides), and population ecology (e.g., the effects of species introductions) provide numerous examples of evolutionary rescue. The general theory of rescue has been tested in laboratory experiments with microbes, in which experimental evolution shows how different treatments affect the frequency of rescue. Overall, these experiments have supported the predictions of general theory: In particular, abundance, variation, and dispersal have pronounced and repeatable effects on the rescue of populations and communities. Extending these laboratory results to the field is a major task for future research.

Theory suggests that evolutionary changes in phenotypic plasticity could either hinder or facilitate evolutionary rescue in a changing climate. Nevertheless, the actual role of evolving plasticity in the responses of natural populations to climate change remains unresolved. Direct observations of evolutionary change in nature are rare, making it difficult to assess the relative contributions of changes in trait means versus changes in plasticity to climate change responses. To address this gap, this review explores several proxies that can be used to understand evolving plasticity in the context of climate change, including space for time substitutions, experimental evolution and tests for genomic divergence at environmentally responsive loci. Comparisons among populations indicate a prominent role for divergence in environmentally responsive traits in local adaptation to climatic gradients. Moreover, genomic comparisons among such populations have identified pervasive divergence in the regulatory regions of environmentally responsive loci. Taken together, these lines of evidence suggest that divergence in plasticity plays a prominent role in adaptation to climatic gradients over space, indicating that evolving plasticity is also likely to play a key role in adaptive responses to climate change through time. This suggests that genetic variation in plastic responses to the environment (G × E) might be an important predictor of species' vulnerabilities to climate-driven decline or extinction. This article is part of the theme issue ‘The role of plasticity in phenotypic adaptation to rapid environmental change’.

Local adaptations can determine the potential of populations to respond to environmental changes, yet adaptive genetic variation is commonly ignored in models forecasting species vulnerability and biogeographical shifts under future climate change. Here we integrate genomic and ecological modeling approaches to identify genetic adaptations associated with climate in two cryptic forest bats. We then incorporate this information directly into forecasts of range changes under future climate change and assessment of population persistence through the spread of climate-adaptive genetic variation (evolutionary rescue potential). Considering climate-adaptive potential reduced range loss projections, suggesting that failure to account for intraspecific variability can result in overestimation of future losses. On the other hand, range overlap between species was projected to increase, indicating that interspecific competition is likely to play an important role in limiting species' future ranges. We show that although evolutionary rescue is possible, it depends on a population's adaptive capacity and connectivity. Hence, we stress the importance of incorporating genomic data and landscape connectivity in climate change vulnerability assessments and conservation management.

Assisted gene flow (AGF) between populations has the potential to mitigate maladaptation due to climate change. However, AGF may cause outbreeding depression (especially if source and recipient populations have been long isolated) and may disrupt local adaptation to nonclimatic factors. Selection should eliminate extrinsic outbreeding depression due to adaptive differences in large populations, and simulations suggest that, within a few generations, evolution should resolve mild intrinsic outbreeding depression due to epistasis. To weigh the risks of AGF against those of maladaptation due to climate change, we need to know the species' extent of local adaptation to climate and other environmental factors, as well as its pattern of gene flow. AGF should be a powerful tool for managing foundation and resource-producing species with large populations and broad ranges that show signs of historical adaptation to local climatic conditions.

Adaptive phenotypic plasticity provides a mechanism of developmental rescue in novel and rapidly changing environments. Understanding the underlying mechanism of plasticity is important for predicting both the likelihood that a developmental response is adaptive and associated life-history trade-offs that could influence patterns of subsequent evolutionary rescue. Although evolved developmental switches may move organisms toward a new adaptive peak in a novel environment, such mechanisms often result in maladaptive responses. The induction of generalized physiological mechanisms in new environments is relatively more likely to result in adaptive responses to factors such as novel toxins, heat stress, or pathogens. Developmental selection forms of plasticity, which rely on within-individual selective processes, such as shaping of tissue architecture, trial-and-error learning, or acquired immunity, are particularly likely to result in adaptive plasticity in a novel environment. However, both the induction of plastic responses and the ability to be plastic through developmental selection come with significant costs, resulting in delays in reproduction, increased individual investment, and reduced fecundity. Thus, we might expect complex interactions between plastic responses that allow survival in novel environments and subsequent evolutionary responses at the population level.

Populations subject to severe stress may be rescued by natural selection, but its operation is restricted by ecological and genetic constraints. The cost of natural selection expresses the limited capacity of a population to sustain the load of mortality or sterility required for effective selection. Genostasis expresses the lack of variation that prevents many populations from adapting to stress. While the role of relative fitness in adaptation is well understood, evolutionary rescue emphasizes the need to recognize explicitly the importance of absolute fitness. Permanent adaptation requires a range of genetic variation in absolute fitness that is broad enough to provide a few extreme types capable of sustained growth under a stress that would cause extinction if they were not present. This principle implies that population size is an important determinant of rescue. The overall number of individuals exposed to selection will be greater when the population declines gradually under a constant stress, or is progressively challenged by gradually increasing stress. In gradually deteriorating environments, survival at lethal stress may be procured by prior adaptation to sublethal stress through genetic correlation. Neither the standing genetic variation of small populations nor the mutation supply of large populations, however, may be sufficient to provide evolutionary rescue for most populations.

Thousands of small populations are at increased risk of extinction because genetics and evolutionary biology are not well‐integrated into conservation planning–a major lost opportunity for effective actions. We propose that if the risk of outbreeding depression is low, the default should be to evaluate restoration of gene flow to small inbred populations of diploid outbreeding organisms that were isolated by human activities within the last 500 years, rather than inaction. We outline the elements of a scientific‐based genetic management policy for fragmented populations of plants and animals, and discuss the reasons why the current default policy is, inappropriately, inaction.

The research framework of eco‐evolutionary dynamics is increasing in popularity, as revealed by a steady stream of review articles and a recent and influential book, but primary empirical research is lagging behind. Moreover, the few empirical case studies demonstrating eco‐evolutionary dynamics might not be entirely representative. Much current research on eco‐evolutionary dynamics is focused on how ecological interactions lead to natural selection on phenotypic traits (“eco‐evo”), and in turn how the evolutionary change in such traits feed back on ecological dynamics (“evo‐eco”). A key feature of eco‐evolutionary dynamics is thus a feedback loop between ecology (e.g., population dynamics) and evolution (i.e., genetic change). In contrast to previous research on eco‐evolutionary dynamics driven by natural selection, the role of eco‐evolutionary feedbacks in sexual selection and sexual conflict is largely unknown. Here, I review theory and the limited empirical evidence in this area and identify some promising future lines of research. I update a past review on contemporary evolution of secondary sexual traits in natural populations and formulate six explicit and rigorous criteria for contemporary evolution of secondary sexual traits by natural or sexual selection or sexual conflict. I then discuss the other key prediction of eco‐evolutionary dynamics (i.e., evolution by sexual selection or sexual conflict shapes ecological dynamics). My overview reveals that our current knowledge in this area is limited and mainly come from theoretical models and laboratory experiments. A major challenge in eco‐evolutionary dynamics is therefore to link ecological and population dynamics with sexual selection and sexual conflict. This is not an easy task but might be possible with carefully chosen study systems and methods. A plain language summary is available for this article. Plain Language Summary

Why is drug resistance common and vaccine resistance rare? Drugs and vaccines both impose substantial pressure on pathogen populations to evolve resistance and indeed, drug resistance typically emerges soon after the introduction of a drug. But vaccine resistance has only rarely emerged. Using well-established principles of population genetics and evolutionary ecology, we argue that two key differences between vaccines and drugs explain why vaccines have so far proved more robust against evolution than drugs. First, vaccines tend to work prophylactically while drugs tend to work therapeutically. Second, vaccines tend to induce immune responses against multiple targets on a pathogen while drugs tend to target very few. Consequently, pathogen populations generate less variation for vaccine resistance than they do for drug resistance, and selection has fewer opportunities to act on that variation. When vaccine resistance has evolved, these generalities have been violated. With careful forethought, it may be possible to identify vaccines at risk of failure even before they are introduced.