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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.

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.

That sexually mature fish skip reproduction, especially in response to poor condition, has been documented in many species. We present results from an energy-allocation life history model that shed light on the underlying logic of skipped spawning, based on the Northeast Arctic stock of Atlantic cod (Gadus morhua). The model predicts that skipped spawning is a regular phenomenon, with up to 30% of the sexually mature biomass skipping spawning. Spawning should be skipped if the expected future gain in reproductive output, discounted by survival, more than balances the expected reproductive success the current year. Skipped spawning was most common (i) among potential second-time spawners and (ii) early in life, (iii) when fishing mortality at the spawning grounds was high, (iv) when fishing mortality at the feeding grounds was low, (v) when natural mortality was low, and (vi) when the energetic and mortality costs associated with migration and spawning were high. Cod skipped spawning more often when food availability was both increased (opportunities for better growth) and decreased (too little energy for gonad development), and this pattern interacted with mortality rate. We conclude that skipped spawning may be more widespread than appreciated and highlight potential consequences for the understanding of stock–recruitment relationships.

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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.

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