Erin S. Dunlop

Implications of fisheries-induced evolution for stock rebuilding and recovery

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 results 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.

Eco-genetic modeling of contemporary life-history evolution.

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 populations 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 a harvest moratorium. This illustrates how predictions offered by eco-genetic models can enable and guide evolutionarily sustainable resource management.

THE EVOLUTION OF SPAWNING MIGRATIONS: STATE DEPENDENCE AND FISHING‐INDUCED CHANGES

Individuals migrate to exploit heterogeneities between spatially separated environments to modulate growth, survival, or reproduction. We devised a bioenergetics model to investigate the evolution of migration distance and its dependence on individual states. Atlantic cod Gadus morhua ranges from sedentary populations to stocks that migrate several thousand kilometers annually. We focused on the Northeast Arctic cod stock, which migrates south to spawn. A linear relationship between migration distance and the expected survival of offspring was assumed, here understood as the prospects for future survival and development that a fertilized egg faces at a particular spawning location. Reasons for why it may increase southward include warmer water that increases development rates, and thereby survival, along the pelagic drift trajectory. In the model, ingested energy can either be allocated to growth or stored for migration and reproduction. When migrating, individuals forgo foraging opportunities and expend energy. Optimal energy allocation and migration strategies were found using state-dependent optimization, with body length, age, condition, and current food availability as individual states. For both a historical and contemporary fishing regime we modeled two behaviors: (1) homing cod returning to the same spawning site each year and (2) roaming cod with no such constraints. The model predicted distinct regions of locally high spawning stock biomass. Large individuals in good condition migrated farthest, and these also tended to mature later in life. The roaming cod spread farther south as they grew larger and older. Homing cod did not have this freedom, and spawning was generally concentrated along a narrower stretch of the coastline. Under contemporary fishing, individuals matured earlier at a smaller size, had shorter migrations, spawned over a contracted geographical range, and tended to be in poorer condition. The effects were most pronounced for the homing behavior.

The impact of fishing-induced mortality on the evolution of alternative life-history tactics in brook charr

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 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.

Evolutionary impact assessment: accounting for evolutionary consequences of fishing in an ecosystem approach to fisheries management

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.

Demographic and Evolutionary Consequences of Selective Mortality: Predictions from an Eco-Genetic Model for Smallmouth Bass

Abstract We use an individual-based eco-genetic model to examine the demographic and evolutionary consequences of selective mortality on a species with parental care, the smallmouth bass Micropterus dolomieu. Our analyses are grounded in a long-term (1936-2003) empirical study of the dynamics of two populations that differ widely in both density and life history. The model we construct extends previous approaches by including phenotypic plasticity in age and size at maturation, by permitting density-dependent somatic growth, and by analyzing how costs associated with parental care alter model predictions. We show that, first, additional mortality on age-0 individuals applied for 100 years causes reduced population abundance and biomass, faster somatic growth rates, and phenotypic plasticity toward slightly larger sizes at maturation. Second, mortality on individuals above a minimum size limit, also applied for 100 years, has a small influence on population abundance and somatic growth, causes a reduction ...

Propensity of marine reserves to reduce the evolutionary effects of fishing in a migratory species.

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 a 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.

Toward Darwinian fisheries management

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Economic repercussions of fisheries-induced evolution

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.

Evolution: Unnatural selection

Fishing and hunting by humans are the main causes of mortality in many populations of wild animals. The consequence is that large and rapid changes occur in certain characteristics that far exceed changes due to other agents.