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Harvested marine fish stocks often show a rapid and substantial decline in the age and size at maturation. Such changes can arise from multiple processes including fisheries‐induced evolution, phenotypic plasticity, and responses to environmental factors other than harvest. The relative importance of these processes could differ systematically between marine and freshwater systems. We tested for temporal shifts in the mean and within‐cohort variability of age‐ and size‐based maturation probabilities of female yellow perch (Perca flavescens Mitchill) from four management units (MUs) in Lake Erie. Lake Erie yellow perch have been commercially harvested for more than a century, and age and size at maturation have varied since sampling began in the 1980s. Our analysis compared probabilistic maturation reaction norms (PMRNs) for cohorts when abundance was lower and harvest higher (1993–1998) to cohorts when abundance was higher and harvest lower (2005–2010). PMRNs have been used in previous studies to detect signs of evolutionary change in response to harvest. Maturation size threshold increased between the early and late cohorts, and the increases were statistically significant for the youngest age in the western MU1 and for older ages in the eastern MU3. Maturation envelope widths, a measure of the variability in maturation among individuals in a cohort, also increased between early and late cohorts in the western MUs where harvest was highest. The highest rates of change in size at maturation for a given age were as large or larger than rates reported for harvested marine fishes where declines in age and size at maturation have been observed. Contrary to the general observation of earlier maturation evolving in harvested stocks, female yellow perch in Lake Erie may be rapidly evolving delayed maturation since harvest was relaxed in the late 1990s, providing a rare example of possible evolutionary recovery.

Harvested marine fish stocks often show a rapid and substantial decline in the age and size at maturation. Such changes can arise from multiple processes including fisheries‐induced evolution, phenotypic plasticity, and responses to environmental factors other than harvest. The relative importance of these processes could differ systematically between marine and freshwater systems. We tested for temporal shifts in the mean and within‐cohort variability of age‐ and size‐based maturation probabilities of female yellow perch (Perca flavescens Mitchill) from four management units (MUs) in Lake Erie. Lake Erie yellow perch have been commercially harvested for more than a century, and age and size at maturation have varied since sampling began in the 1980s. Our analysis compared probabilistic maturation reaction norms (PMRNs) for cohorts when abundance was lower and harvest higher (1993–1998) to cohorts when abundance was higher and harvest lower (2005–2010). PMRNs have been used in previous studies to detect signs of evolutionary change in response to harvest. Maturation size threshold increased between the early and late cohorts, and the increases were statistically significant for the youngest age in the western MU1 and for older ages in the eastern MU3. Maturation envelope widths, a measure of the variability in maturation among individuals in a cohort, also increased between early and late cohorts in the western MUs where harvest was highest. The highest rates of change in size at maturation for a given age were as large or larger than rates reported for harvested marine fishes where declines in age and size at maturation have been observed. Contrary to the general observation of earlier maturation evolving in harvested stocks, female yellow perch in Lake Erie may be rapidly evolving delayed maturation since harvest was relaxed in the late 1990s, providing a rare example of possible evolutionary recovery.

The relative roles of density dependence and life history evolution in contributing to rapid fisheries-induced trait changes remain debated. In the 1930s, northeast Arctic cod (Gadus morhua), currently the world’s largest cod stock, experienced a shift from a traditional spawning-ground fishery to an industrial trawl fishery with elevated exploitation in the stock’s feeding grounds. Since then, age and length at maturation have declined dramatically, a trend paralleled in other exploited stocks worldwide. These trends can be explained by demographic truncation of the population’s age structure, phenotypic plasticity in maturation arising through density-dependent growth, fisheries-induced evolution favoring faster-growing or earlier-maturing fish, or a combination of these processes. Here, we use a multitrait eco-evolutionary model to assess the capacity of these processes to reproduce 74 y of historical data on age and length at maturation in northeast Arctic cod, while mimicking the stock’s historical harvesting regime. Our results show that model predictions critically depend on the assumed density dependence of growth: when this is weak, life history evolution might be necessary to prevent stock collapse, whereas when a stronger density dependence estimated from recent data is used, the role of evolution in explaining fisheries-induced trait changes is diminished. Our integrative analysis of density-dependent growth, multitrait evolution, and stock-specific time series data underscores the importance of jointly considering evolutionary and ecological processes, enabling a more comprehensive perspective on empirically observed stock dynamics than previous studies could provide.

The relative roles of density dependence and life history evolution in contributing to rapid fisheries-induced trait changes remain debated. In the 1930s, northeast Arctic cod (Gadus morhua), currently the world’s largest cod stock, experienced a shift from a traditional spawning-ground fishery to an industrial trawl fishery with elevated exploitation in the stock’s feeding grounds. Since then, age and length at maturation have declined dramatically, a trend paralleled in other exploited stocks worldwide. These trends can be explained by demographic truncation of the population’s age structure, phenotypic plasticity in maturation arising through density-dependent growth, fisheries-induced evolution favoring faster-growing or earlier-maturing fish, or a combination of these processes. Here, we use a multitrait eco-evolutionary model to assess the capacity of these processes to reproduce 74 y of historical data on age and length at maturation in northeast Arctic cod, while mimicking the stock’s historical harvesting regime. Our results show that model predictions critically depend on the assumed density dependence of growth: when this is weak, life history evolution might be necessary to prevent stock collapse, whereas when a stronger density dependence estimated from recent data is used, the role of evolution in explaining fisheries-induced trait changes is diminished. Our integrative analysis of density-dependent growth, multitrait evolution, and stock-specific time series data underscores the importance of jointly considering evolutionary and ecological processes, enabling a more comprehensive perspective on empirically observed stock dynamics than previous studies could provide.

The relative roles of density dependence and life history evolution in contributing to rapid fisheries-induced trait changes remain debated. In the 1930s, northeast Arctic cod (Gadus morhua), currently the world’s largest cod stock, experienced a shift from a traditional spawning-ground fishery to an industrial trawl fishery with elevated exploitation in the stock’s feeding grounds. Since then, age and length at maturation have declined dramatically, a trend paralleled in other exploited stocks worldwide. These trends can be explained by demographic truncation of the population’s age structure, phenotypic plasticity in maturation arising through density-dependent growth, fisheries-induced evolution favoring faster-growing or earlier-maturing fish, or a combination of these processes. Here, we use a multitrait eco-evolutionary model to assess the capacity of these processes to reproduce 74 y of historical data on age and length at maturation in northeast Arctic cod, while mimicking the stock’s historical harvesting regime. Our results show that model predictions critically depend on the assumed density dependence of growth: when this is weak, life history evolution might be necessary to prevent stock collapse, whereas when a stronger density dependence estimated from recent data is used, the role of evolution in explaining fisheries-induced trait changes is diminished. Our integrative analysis of density-dependent growth, multitrait evolution, and stock-specific time series data underscores the importance of jointly considering evolutionary and ecological processes, enabling a more comprehensive perspective on empirically observed stock dynamics than previous studies could provide.

Managing fisheries resources to maintain healthy ecosystems is one of the main goals of the ecosystem approach to fisheries (EAF). While a number of international treaties call for the implementation of EAF, there are still gaps in the underlying methodology. One aspect that has received substantial scientific attention recently is fisheries-induced evolution (FIE). Increasing evidence indicates that intensive fishing has the potential to exert strong directional selection on life-history traits, behaviour, physiology, and morphology of exploited fish. Of particular concern is that reversing evolutionary responses to fishing can be much more difficult than reversing demographic or phenotypically plastic responses. Furthermore, like climate change, multiple agents cause FIE, with effects accumulating over time. Consequently, FIE may alter the utility derived from fish stocks, which in turn can modify the monetary value living aquatic resources provide to society. Quantifying and predicting the evolutionary effects of fishing is therefore important for both ecological and economic reasons. An important reason this is not happening is the lack of an appropriate assessment framework. We therefore describe the evolutionary impact assessment (EvoIA) as a structured approach for assessing the evolutionary consequences of fishing and evaluating the predicted evolutionary outcomes of alternative management options. EvoIA can contribute to EAF by clarifying how evolution may alter stock properties and ecological relations, support the precautionary approach to fisheries management by addressing a previously overlooked source of uncertainty and risk, and thus contribute to sustainable fisheries.

Managing fisheries resources to maintain healthy ecosystems is one of the main goals of the ecosystem approach to fisheries (EAF). While a number of international treaties call for the implementation of EAF, there are still gaps in the underlying methodology. One aspect that has received substantial scientific attention recently is fisheries-induced evolution (FIE). Increasing evidence indicates that intensive fishing has the potential to exert strong directional selection on life-history traits, behaviour, physiology, and morphology of exploited fish. Of particular concern is that reversing evolutionary responses to fishing can be much more difficult than reversing demographic or phenotypically plastic responses. Furthermore, like climate change, multiple agents cause FIE, with effects accumulating over time. Consequently, FIE may alter the utility derived from fish stocks, which in turn can modify the monetary value living aquatic resources provide to society. Quantifying and predicting the evolutionary effects of fishing is therefore important for both ecological and economic reasons. An important reason this is not happening is the lack of an appropriate assessment framework. We therefore describe the evolutionary impact assessment (EvoIA) as a structured approach for assessing the evolutionary consequences of fishing and evaluating the predicted evolutionary outcomes of alternative management options. EvoIA can contribute to EAF by clarifying how evolution may alter stock properties and ecological relations, support the precautionary approach to fisheries management by addressing a previously overlooked source of uncertainty and risk, and thus contribute to sustainable fisheries.

Managing fisheries resources to maintain healthy ecosystems is one of the main goals of the ecosystem approach to fisheries (EAF). While a number of international treaties call for the implementation of EAF, there are still gaps in the underlying methodology. One aspect that has received substantial scientific attention recently is fisheries-induced evolution (FIE). Increasing evidence indicates that intensive fishing has the potential to exert strong directional selection on life-history traits, behaviour, physiology, and morphology of exploited fish. Of particular concern is that reversing evolutionary responses to fishing can be much more difficult than reversing demographic or phenotypically plastic responses. Furthermore, like climate change, multiple agents cause FIE, with effects accumulating over time. Consequently, FIE may alter the utility derived from fish stocks, which in turn can modify the monetary value living aquatic resources provide to society. Quantifying and predicting the evolutionary effects of fishing is therefore important for both ecological and economic reasons. An important reason this is not happening is the lack of an appropriate assessment framework. We therefore describe the evolutionary impact assessment (EvoIA) as a structured approach for assessing the evolutionary consequences of fishing and evaluating the predicted evolutionary outcomes of alternative management options. EvoIA can contribute to EAF by clarifying how evolution may alter stock properties and ecological relations, support the precautionary approach to fisheries management by addressing a previously overlooked source of uncertainty and risk, and thus contribute to sustainable fisheries.

Fish stocks experiencing high fishing mortality show a tendency to mature earlier and at a smaller size, which may have a genetic component and therefore long-lasting economic and biological effects. To date, the economic effects of such ecoevolutionary dynamics have not been empirically investigated. Using 70 y of data, we develop a bioeconomic model for Northeast Arctic cod to compare the economic yield in a model in which life-history traits can vary only through phenotypic plasticity with a model in which, in addition, genetic changes can occur. We find that evolutionary changes toward faster growth and earlier maturation occur consistently even if a stock is optimally managed. However, if a stock is managed optimally, the evolutionary changes actually increase economic yield because faster growth and earlier maturation raise the stock’s productivity. The optimal fishing mortality is almost identical for the evolutionary and nonevolutionary model and substantially lower than what it has been historically. Therefore, the costs of ignoring evolution under optimal management regimes are negligible. However, if fishing mortality is as high as it has been historically, evolutionary changes may result in economic losses, but only if the fishery is selecting for medium-sized individuals. Because evolution facilitates growth, the fish are younger and still immature when they are susceptible to getting caught, which outweighs the increase in productivity due to fish spawning at an earlier age.

Fish stocks experiencing high fishing mortality show a tendency to mature earlier and at a smaller size, which may have a genetic component and therefore long-lasting economic and biological effects. To date, the economic effects of such ecoevolutionary dynamics have not been empirically investigated. Using 70 y of data, we develop a bioeconomic model for Northeast Arctic cod to compare the economic yield in a model in which life-history traits can vary only through phenotypic plasticity with a model in which, in addition, genetic changes can occur. We find that evolutionary changes toward faster growth and earlier maturation occur consistently even if a stock is optimally managed. However, if a stock is managed optimally, the evolutionary changes actually increase economic yield because faster growth and earlier maturation raise the stock's productivity. The optimal fishing mortality is almost identical for the evolutionary and nonevolutionary model and substantially lower than what it has been historically. Therefore, the costs of ignoring evolution under optimal management regimes are negligible. However, if fishing mortality is as high as it has been historically, evolutionary changes may result in economic losses, but only if the fishery is selecting for medium-sized individuals. Because evolution facilitates growth, the fish are younger and still immature when they are susceptible to getting caught, which outweighs the increase in productivity due to fish spawning at an earlier age.

Biological reference points are important tools for fisheries management. Reference points are not static, but may change when a population's environment or the population itself changes. Fisheries-induced evolution is one mechanism that can alter population characteristics, leading to "shifting" reference points by modifying the underlying biological processes or by changing the perception of a fishery system. The former causes changes in "true" reference points, whereas the latter is caused by changes in the yardsticks used to quantify a system's status. Unaccounted shifts of either kind imply that reference points gradually lose their intended meaning. This can lead to increased precaution, which is safe, but potentially costly. Shifts can also occur in more perilous directions, such that actual risks are greater than anticipated. Our qualitative analysis suggests that all commonly used reference points are susceptible to shifting through fisheries-induced evolution, including the limit and "precautionary" reference points for spawning-stock biomass, B_lim and B_pa, and the target reference point for fishing mortality, F_0.1. Our findings call for increased awareness of fisheries-induced changes and highlight the value of always basing reference points on adequately updated information, to capture all changes in the biological processes that drive fish population dynamics.

Harvest control rules (HCRs) have been implemented for many fisheries worldwide. However, in most instances, those HCRs are not based on the explicit feedbacks between stock properties and economic considerations. This paper develops a bio-economic model that evaluates the HCR adopted in 2004 by the Joint Norwegian-Russian Fishery Commission to manage the world's largest cod stock, Northeast Arctic cod (NEA). The model considered here is biologically and economically detailed, and is the first to compare the performance of the stock's current HCR with that of alternative HCRs derived with optimality criteria. In particular, HCRs are optimized for economic objectives including fleet profits, economic welfare, and total yield and the merging properties are analyzed. The performance of these optimal HCRs was compared with the currently used HCR. This paper show that the current HCR does in fact comes very close to maximizing profits. Furthermore, the results reveal that the HCR that maximizes profits is the most precautionary one among the considered HCRs. Finally, the HCR that maximizes yield leads to unprecautionary low levels of biomass. In these ways, the implementation of the HCR for NEA cod can be viewed as a success story that may provide valuable lessons for other fisheries.

Harvest control rules (HCRs) have been implemented for many fisheries worldwide. However, in most instances, those HCRs are not based on the explicit feedbacks between stock properties and economic considerations. This paper develops a bio-economic model that evaluates the HCR adopted in 2004 by the Joint Norwegian-Russian Fishery Commission to manage the world's largest cod stock, Northeast Arctic cod (NEA). The model considered here is biologically and economically detailed, and is the first to compare the performance of the stock's current HCR with that of alternative HCRs derived with optimality criteria. In particular, HCRs are optimized for economic objectives including fleet profits, economic welfare, and total yield and the merging properties are analyzed. The performance of these optimal HCRs was compared with the currently used HCR. This paper show that the current HCR does in fact comes very close to maximizing profits. Furthermore, the results reveal that the HCR that maximizes profits is the most precautionary one among the considered HCRs. Finally, the HCR that maximizes yield leads to unprecautionary low levels of biomass. In these ways, the implementation of the HCR for NEA cod can be viewed as a success story that may provide valuable lessons for other fisheries.

Fish stocks experiencing high fishing mortality show a tendency to mature earlier and at a smaller size, which may have a genetic component and therefore long-lasting economic and biological effects. To date, the economic effects of such ecoevolutionary dynamics have not been empirically investigated. Using 70 y of data, we develop a bioeconomic model for Northeast Arctic cod to compare the economic yield in a model in which life-history traits can vary only through phenotypic plasticity with a model in which, in addition, genetic changes can occur. We find that evolutionary changes toward faster growth and earlier maturation occur consistently even if a stock is optimally managed. However, if a stock is managed optimally, the evolutionary changes actually increase economic yield because faster growth and earlier maturation raise the stock's productivity. The optimal fishing mortality is almost identical for the evolutionary and nonevolutionary model and substantially lower than what it has been historically. Therefore, the costs of ignoring evolution under optimal management regimes are negligible. However, if fishing mortality is as high as it has been historically, evolutionary changes may result in economic losses, but only if the fishery is selecting for medium-sized individuals. Because evolution facilitates growth, the fish are younger and still immature when they are susceptible to getting caught, which outweighs the increase in productivity due to fish spawning at an earlier age.

Biological reference points are important tools for fisheries management. Reference points are not static, but may change when a population's environment or the population itself changes. Fisheries-induced evolution is one mechanism that can alter population characteristics, leading to "shifting" reference points by modifying the underlying biological processes or by changing the perception of a fishery system. The former causes changes in "true" reference points, whereas the latter is caused by changes in the yardsticks used to quantify a system's status. Unaccounted shifts of either kind imply that reference points gradually lose their intended meaning. This can lead to increased precaution, which is safe, but potentially costly. Shifts can also occur in more perilous directions, such that actual risks are greater than anticipated. Our qualitative analysis suggests that all commonly used reference points are susceptible to shifting through fisheries-induced evolution, including the limit and "precautionary" reference points for spawning-stock biomass, B_lim and B_pa, and the target reference point for fishing mortality, F_0.1. Our findings call for increased awareness of fisheries-induced changes and highlight the value of always basing reference points on adequately updated information, to capture all changes in the biological processes that drive fish population dynamics.

Fish stocks experiencing high fishing mortality show a tendency to mature earlier and at a smaller size, which may have a genetic component and therefore long-lasting economic and biological effects. To date, the economic effects of such ecoevolutionary dynamics have not been empirically investigated. Using 70 y of data, we develop a bioeconomic model for Northeast Arctic cod to compare the economic yield in a model in which life-history traits can vary only through phenotypic plasticity with a model in which, in addition, genetic changes can occur. We find that evolutionary changes toward faster growth and earlier maturation occur consistently even if a stock is optimally managed. However, if a stock is managed optimally, the evolutionary changes actually increase economic yield because faster growth and earlier maturation raise the stock's productivity. The optimal fishing mortality is almost identical for the evolutionary and nonevolutionary model and substantially lower than what it has been historically. Therefore, the costs of ignoring evolution under optimal management regimes are negligible. However, if fishing mortality is as high as it has been historically, evolutionary changes may result in economic losses, but only if the fishery is selecting for medium-sized individuals. Because evolution facilitates growth, the fish are younger and still immature when they are susceptible to getting caught, which outweighs the increase in productivity due to fish spawning at an earlier age.

Harvest control rules (HCRs) have been implemented for many fisheries worldwide. However, in most instances, those HCRs are not based on the explicit feedbacks between stock properties and economic considerations. This paper develops a bio-economic model that evaluates the HCR adopted in 2004 by the Joint Norwegian-Russian Fishery Commission to manage the world's largest cod stock, Northeast Arctic cod (NEA). The model considered here is biologically and economically detailed, and is the firt to compare the performance of the stock's current HCR with that of alternative HCRs derived with optimality criteria. In particular, HCRs are optimized for economic objectives including fleet profit, economic welfare, and total yield and the emerging properties are analyzed. The performance of these optimal HCRs was compared with the currently used HCR. This paper show that the current HCR does in fact comes very close to maximizing profits. Furthermore, the results reveal that the HCR that maximizes profits is the most precautionary one among the considered HCRs. Finally, the HCR that maximizes yield leads to un-precautionary low levels of biomass. In these ways, the implementation of the HCR for NEA cod can be viewed as a success story that may provide valuable lessons for other fishries.

The interest in fishing-induced life-history evolution has been growing in the last decade, in part because of the increasing number of studies suggesting evolutionary changes in life-history traits, and the potential ecological and economic consequences these changes may have. Among the traits that could evolve in response to fishing, growth has lately received attention. However, critical reading of the literature on growth evolution in fish reveals conceptual confusion about the nature of growth itself as an evolving trait, and about the different ways fishing can affect growth and size-at-age of fish, both on ecological and on evolutionary time-scales. It is important to separate the advantages of being big and the costs of growing to a large size, particularly when studying life-history evolution. In this review, we explore the selection pressures on growth and the resultant evolution of growth from a mechanistic viewpoint. We define important concepts and outline the processes that must be accounted for before observed phenotypic changes can be ascribed to growth evolution. When listing traits that could be traded-off with growth rate, we group the mechanisms into those affecting resource acquisition and those governing resource allocation. We summarize potential effects of fishing on traits related to growth and discuss methods for detecting evolution of growth. We also challenge the prevailing expectation that fishing-induced evolution should always lead to slower growth.

Harvest control rules (HCRs) have been implemented for many fisheries worldwide. However, in most instances, those HCRs are not based on the explicit feedbacks between stock properties and economic considerations. This paper develops a bio-economic model that evaluates the HCR adopted in 2004 by the Joint Norwegian-Russian Fishery Commission to manage the world's largest cod stock, Northeast Arctic cod (NEA). The model considered here is biologically and economically detailed, and is the firt to compare the performance of the stock's current HCR with that of alternative HCRs derived with optimality criteria. In particular, HCRs are optimized for economic objectives including fleet profit, economic welfare, and total yield and the emerging properties are analyzed. The performance of these optimal HCRs was compared with the currently used HCR. This paper show that the current HCR does in fact comes very close to maximizing profits. Furthermore, the results reveal that the HCR that maximizes profits is the most precautionary one among the considered HCRs. Finally, the HCR that maximizes yield leads to un-precautionary low levels of biomass. In these ways, the implementation of the HCR for NEA cod can be viewed as a success story that may provide valuable lessons for other fishries.

The interest in fishing-induced life-history evolution has been growing in the last decade, in part because of the increasing number of studies suggesting evolutionary changes in life-history traits, and the potential ecological and economic consequences these changes may have. Among the traits that could evolve in response to fishing, growth has lately received attention. However, critical reading of the literature on growth evolution in fish reveals conceptual confusion about the nature of growth itself as an evolving trait, and about the different ways fishing can affect growth and size-at-age of fish, both on ecological and on evolutionary time-scales. It is important to separate the advantages of being big and the costs of growing to a large size, particularly when studying life-history evolution. In this review, we explore the selection pressures on growth and the resultant evolution of growth from a mechanistic viewpoint. We define important concepts and outline the processes that must be accounted for before observed phenotypic changes can be ascribed to growth evolution. When listing traits that could be traded-off with growth rate, we group the mechanisms into those affecting resource acquisition and those governing resource allocation. We summarize potential effects of fishing on traits related to growth and discuss methods for detecting evolution of growth. We also challenge the prevailing expectation that fishing-induced evolution should always lead to slower growth.

Fish stocks experiencing high fishing mortality show a tendency to mature earlier and at a smaller size, which may have a genetic component and therefore long-lasting economic and biological effects. To date, the economic effects of such ecoevolutionary dynamics have not been empirically investigated. Using 70 y of data, we develop a bioeconomic model for Northeast Arctic cod to compare the economic yield in a model in which life-history traits can vary only through phenotypic plasticity with a model in which, in addition, genetic changes can occur. We find that evolutionary changes toward faster growth and earlier maturation occur consistently even if a stock is optimally managed. However, if a stock is managed optimally, the evolutionary changes actually increase economic yield because faster growth and earlier maturation raise the stock's productivity. The optimal fishing mortality is almost identical for the evolutionary and nonevolutionary model and substantially lower than what it has been historically. Therefore, the costs of ignoring evolution under optimal management regimes are negligible. However, if fishing mortality is as high as it has been historically, evolutionary changes may result in economic losses, but only if the fishery is selecting for medium-sized individuals. Because evolution facilitates growth, the fish are younger and still immature when they are susceptible to getting caught, which outweighs the increase in productivity due to fish spawning at an earlier age.

Fish stocks experiencing high fishing mortality show a tendency to mature earlier and at a smaller size, which may have a genetic component and therefore long-lasting economic and biological effects. To date, the economic effects of such ecoevolutionary dynamics have not been empirically investigated. Using 70 y of data, we develop a bioeconomic model for Northeast Arctic cod to compare the economic yield in a model in which life-history traits can vary only through phenotypic plasticity with a model in which, in addition, genetic changes can occur. We find that evolutionary changes toward faster growth and earlier maturation occur consistently even if a stock is optimally managed. However, if a stock is managed optimally, the evolutionary changes actually increase economic yield because faster growth and earlier maturation raise the stock's productivity. The optimal fishing mortality is almost identical for the evolutionary and nonevolutionary model and substantially lower than what it has been historically. Therefore, the costs of ignoring evolution under optimal management regimes are negligible. However, if fishing mortality is as high as it has been historically, evolutionary changes may result in economic losses, but only if the fishery is selecting for medium-sized individuals. Because evolution facilitates growth, the fish are younger and still immature when they are susceptible to getting caught, which outweighs the increase in productivity due to fish spawning at an earlier age.

Fish stocks experiencing high fishing mortality show a tendency to mature earlier and at a smaller size, which may have a genetic component and therefore long-lasting economic and biological effects. To date, the economic effects of such ecoevolutionary dynamics have not been empirically investigated. Using 70 y of data, we develop a bioeconomic model for Northeast Arctic cod to compare the economic yield in a model in which life-history traits can vary only through phenotypic plasticity with a model in which, in addition, genetic changes can occur. We find that evolutionary changes toward faster growth and earlier maturation occur consistently even if a stock is optimally managed. However, if a stock is managed optimally, the evolutionary changes actually increase economic yield because faster growth and earlier maturation raise the stock's productivity. The optimal fishing mortality is almost identical for the evolutionary and nonevolutionary model and substantially lower than what it has been historically. Therefore, the costs of ignoring evolution under optimal management regimes are negligible. However, if fishing mortality is as high as it has been historically, evolutionary changes may result in economic losses, but only if the fishery is selecting for medium-sized individuals. Because evolution facilitates growth, the fish are younger and still immature when they are susceptible to getting caught, which outweighs the increase in productivity due to fish spawning at an earlier age.

There is increasing evidence that fishing may cause rapid contemporary evolution in freshwater and marine fish populations. This has led to growing concern about the possible consequences such evolutionary change might have for aquatic ecosystems and the utility of those ecosystems to society. This special issue contains contributions from a symposium on fisheries-induced evolution held at the American Fisheries Society Annual Meeting in August 2008. Contributions include primary studies and reviews of field-based and experimental evidence, and several theoretical modeling studies advancing life-history theory and investigating potential management options. In this introduction we review the state of research in the field, discuss current controversies, and identify contributions made by the papers in this issue to the knowledge of fisheries-induced evolution. We end by suggesting directions for future research.

We present eco-genetic modeling as a flexible tool for exploring the course and rates of multi-trait life-history evolution in natural populations. We build on existing modeling approaches by combining features that facilitate studying the ecological and evolutionary dynamics of realistically structured populations. In particular, the joint consideration of age and size structure enables the analysis of phenotypically plastic populations with more than a single growth trajectory, and ecological feedback is readily included in the form of density dependence and frequency dependence. Stochasticity and life-history trade-offs can also be implemented. Critically, eco-genetic models permit the incorporation of salient genetic detail such as a population's genetic variances and covariances and the corresponding heritabilities, as well as the probabilistic inheritance and phenotypic expression of quantitative traits. These inclusions are crucial for predicting rates of evolutionary change on both contemporary and longer timescales. An eco-genetic model can be tightly coupled with empirical data and therefore may have considerable practical relevance, in terms of generating testable predictions and evaluating alternative management measures. To illustrate the utility of these models we present as an example an eco-genetic model used to study harvest-induced evolution of multiple traits in Atlantic cod. The predictions of our model (most notably that harvesting induces a genetic reduction in age and size at maturation, an increase or decrease in growth capacity depending on the minimum-length limit, and an increase in reproductive investment) are corroborated by patterns observed in wild populations. The predicted genetic changes occur together with plastic changes that could phenotypically mask the former. Importantly, our analysis predicts that evolutionary changes show little signs of reversal following harvest moratorium. This illustrates how predictions offered by eco-g

We present eco-genetic modeling as a flexible tool for exploring the course and rates of multi-trait life-history evolution in natural populations. We build on existing modeling approaches by combining features that facilitate studying the ecological and evolutionary dynamics of realistically structured populations. In particular, the joint consideration of age and size structure enables the analysis of phenotypically plastic populations with more than a single growth trajectory, and ecological feedback is readily included in the form of density dependence and frequency dependence. Stochasticity and life-history trade-offs can also be implemented. Critically, eco-genetic models permit the incorporation of salient genetic detail such as a population's genetic variances and covariances and the corresponding heritabilities, as well as the probabilistic inheritance and phenotypic expression of quantitative traits. These inclusions are crucial for predicting rates of evolutionary change on both contemporary and longer timescales. An eco-genetic model can be tightly coupled with empirical data and therefore may have considerable practical relevance, in terms of generating testable predictions and evaluating alternative management measures. To illustrate the utility of these models we present as an example an eco-genetic model used to study harvest-induced evolution of multiple traits in Atlantic cod. The predictions of our model (most notably that harvesting induces a genetic reduction in age and size at maturation, an increase or decrease in growth capacity depending on the minimum-length limit, and an increase in reproductive investment) are corroborated by patterns observed in wild populations. The predicted genetic changes occur together with plastic changes that could phenotypically mask the former. Importantly, our analysis predicts that evolutionary changes show little signs of reversal following harvest moratorium. This illustrates how predictions offered by eco-g

Worldwide depletion of fish stocks has led fisheries managers to become increasingly concerned about rebuilding and recovery planning. To succeed, factors affecting recovery dynamics need to be understood, including the role of fisheries-induced evolution. Here we investigate a stock's response to fishing followed by a harvest moratorium by analyzing an individual-based evolutionary model parameterized for Atlantic cod Gadus morhua from its northern range, representative of long-lived, late-maturing species. The model allows evolution of life-history processes including maturation, reproduction, and growth. It also incorporates environmental variability, phenotypic plasticity, and density-dependent feedbacks. Fisheries-induced evolution affects recovery in several ways. The first decades of recovery were dominated by demographic and density-dependent processes. Biomass rebuilding was only lightly influenced by fisheries-induced evolution, whereas other stock characteristics such as maturation age, spawning stock biomass, and recruitment were substantially affected, recovering to new demographic equilibria below their preharvest levels. This is because genetic traits took thousands of years to evolve back to preharvest levels, indicating that natural selection driving recovery of these traits is weaker than fisheries-induced selection was. Our result strengthen the case for proactive management of fisheries-induced evolution, as the restoration of genetic traits altered by fishing is slow and may even be impractical.

Evolutionary effects of fishing can have unwanted consequences diminishing a fishery's value and sustainability. Reserves, or no-take areas, have been proposed as a management tool for reducing fisheries-induced selection, but their effectiveness for migratory species has remained unexplored. Here we develop an eco-genetic model to predict the effects of marine reserves on fisheries-induced evolution under migration. To represent a stock that undergoes an annual migration between feeding and spawning grounds, we draw model parameters from Atlantic cod (Gadus morhua) in the northern part of its range. Our analysis leads to the following conclusions: (i) a reserve in stock's feeding grounds, protecting immature and mature fish alike, reduces fisheries-induced evolution, even though protected and unprotected population components mix on the spawning grounds; (ii) in contrast, a reserve in a stock's spawning grounds, protecting only mature fish, has little mitigating effects on fisheries-induced evolution and can sometimes even exacerbate its magnitude; (iii) evolutionary changes that are already underway may be difficult to reverse with a reserve; (iv) directly after a reserve is created or enlarged, most reserve scenarios result in yield losses; and (v) timescale is very important: short-term yield losses immediately after a reserve's creation can give way to long-term gains.

There is increasing evidence that fishing may cause rapid contemporary evolution in freshwater and marine fish populations. This has led to growing concern about the possible consequences such evolutionary change might have for aquatic ecosystems and the utility of those ecosystems to society. This special issue contains contributions from a symposium on fisheries-induced evolution held at the American Fisheries Society Annual Meeting in August 2008. Contributions include primary studies and reviews of field-based and experimental evidence, and several theoretical modeling studies advancing life-history theory and investigating potential management options. In this introduction we review the state of research in the field, discuss current controversies, and identify contributions made by the papers in this issue to the knowledge of fisheries-induced evolution. We end by suggesting directions for future research.

There is increasing evidence that fishing may cause rapid contemporary evolution in freshwater and marine fish populations. This has led to growing concern about the possible consequences such evolutionary change might have for aquatic ecosystems and the utility of those ecosystems to society. This special issue contains contributions from a symposium on fisheries-induced evolution held at the American Fisheries Society Annual Meeting in August 2008. Contributions include primary studies and reviews of field-based and experimental evidence, and several theoretical modeling studies advancing life-history theory and investigating potential management options. In this introduction we review the state of research in the field, discuss current controversies, and identify contributions made by the papers in this issue to the knowledge of fisheries-induced evolution. We end by suggesting directions for future research.

Worldwide depletion of fish stocks has led fisheries managers to become increasingly concerned about rebuilding and recovery planning. To succeed, factors affecting recovery dynamics need to be understood, including the role of fisheries-induced evolution. Here we investigate a stock's response to fishing followed by a harvest moratorium by analyzing an individual-based evolutionary model parameterized for Atlantic cod Gadus morhua from its northern range, representative of long-lived, late-maturing species. The model allows evolution of life-history processes including maturation, reproduction, and growth. It also incorporates environmental variability, phenotypic plasticity, and density-dependent feedbacks. Fisheries-induced evolution affects recovery in several ways. The first decades of recovery were dominated by demographic and density-dependent processes. Biomass rebuilding was only lightly influenced by fisheries-induced evolution, whereas other stock characteristics such as maturation age, spawning stock biomass, and recruitment were substantially affected, recovering to new demographic equilibria below their preharvest levels. This is because genetic traits took thousands of years to evolve back to preharvest levels, indicating that natural selection driving recovery of these traits is weaker than fisheries-induced selection was. Our result strengthen the case for proactive management of fisheries-induced evolution, as the restoration of genetic traits altered by fishing is slow and may even be impractical.

Evolutionary effects of fishing can have unwanted consequences diminishing a fishery's value and sustainability. Reserves, or no-take areas, have been proposed as a management tool for reducing fisheries-induced selection, but their effectiveness for migratory species has remained unexplored. Here we develop an eco-genetic model to predict the effects of marine reserves on fisheries-induced evolution under migration. To represent a stock that undergoes an annual migration between feeding and spawning grounds, we draw model parameters from Atlantic cod (Gadus morhua) in the northern part of its range. Our analysis leads to the following conclusions: (i) a reserve in stock's feeding grounds, protecting immature and mature fish alike, reduces fisheries-induced evolution, even though protected and unprotected population components mix on the spawning grounds; (ii) in contrast, a reserve in a stock's spawning grounds, protecting only mature fish, has little mitigating effects on fisheries-induced evolution and can sometimes even exacerbate its magnitude; (iii) evolutionary changes that are already underway may be difficult to reverse with a reserve; (iv) directly after a reserve is created or enlarged, most reserve scenarios result in yield losses; and (v) timescale is very important: short-term yield losses immediately after a reserve's creation can give way to long-term gains.

Worldwide depletion of fish stocks has led fisheries managers to become increasingly concerned about rebuilding and recovery planning. To succeed, factors affecting recovery dynamics need to be understood, including the role of fisheries-induced evolution. Here we investigate a stock's response to fishing followed by a harvest moratorium by analyzing an individual-based evolutionary model parameterized for Atlantic cod Gadus morhua from its northern range, representative of long-lived, late-maturing species. The model allows evolution of life-history processes including maturation, reproduction, and growth. It also incorporates environmental variability, phenotypic plasticity, and density-dependent feedbacks. Fisheries-induced evolution affects recovery in several ways. The first decades of recovery were dominated by demographic and density-dependent processes. Biomass rebuilding was only lightly influenced by fisheries-induced evolution, whereas other stock characteristics such as maturation age, spawning stock biomass, and recruitment were substantially affected, recovering to new demographic equilibria below their preharvest levels. This is because genetic traits took thousands of years to evolve back to preharvest levels, indicating that natural selection driving recovery of these traits is weaker than fisheries-induced selection was. Our result strengthen the case for proactive management of fisheries-induced evolution, as the restoration of genetic traits altered by fishing is slow and may even be impractical.

Evolutionary effects of fishing can have unwanted consequences diminishing a fishery's value and sustainability. Reserves, or no-take areas, have been proposed as a management tool for reducing fisheries-induced selection, but their effectiveness for migratory species has remained unexplored. Here we develop an eco-genetic model to predict the effects of marine reserves on fisheries-induced evolution under migration. To represent a stock that undergoes an annual migration between feeding and spawning grounds, we draw model parameters from Atlantic cod (Gadus morhua) in the northern part of its range. Our analysis leads to the following conclusions: (i) a reserve in stock's feeding grounds, protecting immature and mature fish alike, reduces fisheries-induced evolution, even though protected and unprotected population components mix on the spawning grounds; (ii) in contrast, a reserve in a stock's spawning grounds, protecting only mature fish, has little mitigating effects on fisheries-induced evolution and can sometimes even exacerbate its magnitude; (iii) evolutionary changes that are already underway may be difficult to reverse with a reserve; (iv) directly after a reserve is created or enlarged, most reserve scenarios result in yield losses; and (v) timescale is very important: short-term yield losses immediately after a reserve's creation can give way to long-term gains.

Although contemporary trends indicative of evolutionary change have been detected in the life-history traits of exploited populations, it is not known to what extent fishing influences the evolution of alternative life-history tactics in migratory species such as salmonids. Here, we build a model to predict the evolution of anadromy and residency in an exploited population of brook charr, "Salvelinus fontinalis". Our model allows for both phenotypic plasticity and genetic change in the age and size at migration by including migration reaction norms. Using this migration reaction norm, resulting in a decrease in average probabilities of migration with increasing harvest rate. Moreover, we show that differences in natural mortalities in freshwater greatly influence the magnitude and rate of evolutionary change. The fishing-induced changes in migration predicted by our model alter population abundances and reproductive output and should be accounted for in the sustainable management of salmonids.

Although contemporary trends indicative of evolutionary change have been detected in the lifehistory traits of exploited populations, it is not known to what extent fishing influences the evolution of alternative life-history tactics in migratory species such as salmonids. Here, we build a model to predict the evolution of anadromy and residency in an exploited population of brook charr, 'Salvelinus fontinalis'. Our model allows for both phenotypic plasticity and genetic change in the age and size at migration by including migration reaction norms. Using this model, we predict that fishing of anadromous individuals over the course of 100 years causes evolution in the migration reaction norm, resulting in a decrease in average probabilities of migration with increasing harvest rate. Moreover, we show that differences in natural mortalities in freshwater greatly influence the magnitude and rate of evolutionary change. The fishing-induced changes in migration predicted by our model alter population abundances and reproductive output and should be accounted for in the sustainable management of salmonids.

Although contemporary trends indicative of evolutionary change have been detected in the life-history traits of exploited populations, it is not known to what extent fishing influences the evolution of alternative life-history tactics in migratory species such as salmonids. Here, we build a model to predict the evolution of anadromy and residency in an exploited population of brook charr, "Salvelinus fontinalis". Our model allows for both phenotypic plasticity and genetic change in the age and size at migration by including migration reaction norms. Using this migration reaction norm, resulting in a decrease in average probabilities of migration with increasing harvest rate. Moreover, we show that differences in natural mortalities in freshwater greatly influence the magnitude and rate of evolutionary change. The fishing-induced changes in migration predicted by our model alter population abundances and reproductive output and should be accounted for in the sustainable management of salmonids.