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It is generally assumed that fish populations are regulated primarily in the juvenile (pre-recruit) phase of the life cycle, although density dependence in growth and reproductive parameters within the recruited phase has been widely reported. Here we present evidence to suggest that density-dependent growth in the recruited phase is a key process in the regulation of many fish populations. We analyse 16 fish populations with long-term records of size-at-age and biomass data, and detect significant density-dependent growth in nine. Among-population comparisons show a close, inverse relationship between the estimated decline in asymptotic length per unit biomass density, and the long-term average biomass density of populations. A simple population model demonstrates that regulation by density-dependent growth alone is sufficient to generate the observed relationship. Density-dependent growth should be accounted for in fisheries' assessments, and the empirical relationship established here can provide indicative estimates of the density-dependent growth parameter where population-specific data are lacking.

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.

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.

Dickey-Collas, M., Nash, R. D. M., Brunel, T., van Damme, C. J. G., Marshall, C. T., Payne, M. R., Corten, A., Geffen, A. J., Peck, M. A., Hatfield, E. M. C., Hintzen, N. T., Enberg, K., Kell, L. T., and Simmonds, E. J. 2010. Lessons learned from stock collapse and recovery of North Sea herring: a review. – ICES Journal of Marine Science, 67: 1875–1886. The collapse and recovery of North Sea herring in the latter half of the 20th century had both ecological and economic consequences. We review the effect of the collapse and investigate whether the increased understanding about the biology, ecology, and stock dynamics gained in the past three decades can aid management to prevent further collapses and improve projections of recovery. Recruitment adds the most uncertainty to estimates of future yield and the potential to reach biomass reference points within a specified time-frame. Stock–recruitment relationships must be viewed as being fluid and dependent on ecosystem change. Likewise, predation mortality changes over time. Management aimed at maximum sustainable yield (MSY) fishing mortality targets implies interannual variation in TACs, and variability in supply is therefore unavoidable. Harvest control rules, when adhered to, aid management greatly. We advocate that well-founded science can substantially contribute to management through improved confidence and increased transparency. At present, we cannot predict the effects of collapse or recovery of a single stock on the ecosystem as a whole. Moreover, as managers try to reconcile commitments to single-species MSY targets with the ecosystem-based approach, they must consider the appropriate management objectives for the North Sea ecosystem as a whole.

Heino, M., Baulier, L., Boukal, D. S., Ernande, B., Johnston, F. D., Mollet, F. M., Pardoe, H., Therkildsen, N. O., Uusi-Heikkilä, S., Vainikka, A., Arlinghaus, R., Dankel, D. J., Dunlop, E. S., Eikeset, A. M., Enberg, K., Engelhard G. H., Jørgensen, C., Laugen, A. T., Matsumura, S., Nusslé, S., Urbach, D., Whitlock, R., Rijnsdorp, A. D., and Dieckmann, U. 2013. Can fisheries-induced evolution shift reference points for fisheries management? – ICES Journal of Marine Science, 70: 707–721. 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, Blim and Bpa, and the target reference point for fishing mortality, F0.1. 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.

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