By providing new approaches to the investigation of demographic and evolutionary dynamics of wild populations, molecular genetics has led to fundamental changes in our understanding of marine ecology. In particular, genetic approaches have revolutionized our understanding in three areas: (i) most importantly, they have contributed to the discovery of extensive genetic population structure in many marine species, overturning the notion of large, essentially homogenous marine populations limiting local adaptation and speciation. (ii) Concomitant differences in ecologically important traits now indicate extensive adaptive differentiation and biocomplexity, potentially increasing the resilience to exploitation and disturbance. Evidence for rapid adaptive change in many populations underlies recent concerns about fisheries-induced evolution affecting life-history traits. (iii) A compilation of recent published research shows estimated effective population sizes that are 2‐6 orders of magnitude smaller than census sizes, suggesting more complex recruitment dynamics in marine species than previously assumed. Studies on Atlantic cod are used to illustrate these paradigm shifts. In our synthesis, we emphasize the implications of these discoveries for marine ecology and evolution as well as the management and conservation of exploited marine fish populations. An important implication of genetic structuring and the potential for adaptive divergence is that locally adapted populations are unlikely to be replaced through immigration, with potentially detrimental consequences for the resilience to environmental change ‐ a key consideration for sustainable fisheries management.

Paradigm shifts in marine fisheries genetics: ugly hypotheses slain by beautiful facts

Genetic stochasticity due to small population size contributes to population extinction, especially when population fragmentation disrupts gene flow. Estimates of effective population size (Ne) can therefore be informative about population persistence, but there is a need for an assessment of their consistency and informative relevance. Here we review the body of empirical estimates of Ne for wild populations obtained with the temporal genetic method and published since Frankham's (1995) review. Theoretical considerations have identified important sources of bias for this analytical approach, and we use empirical data to investigate the extent of these biases. We find that particularly model selection and sampling require more attention in future studies.

We report a median unbiased Ne estimate of 260 (among 83 studies) and find that this median estimate tends to be smaller for populations of conservation concern, which may therefore be more sensitive to genetic stochasticity. Furthermore, we report a median Ne/N ratio of 0.14, and find that this ratio may actually be higher for small populations, suggesting changes in biological interactions at low population abundances. We confirm the role of gene flow in countering genetic stochasticity by finding that Ne correlates strongest with neutral genetic metrics when populations can be considered isolated. This underlines the importance of gene flow for the estimation of Ne, and of population connectivity for conservation in general. Reductions in contemporary gene flow due to ongoing habitat fragmentation will likely increase the prevalence of genetic stochasticity, which should therefore remain a focal point in the conservation of biodiversity.

https://onlinelibrary.wiley.com/doi/pdf/10.1111/j.1365-294X.2008.03842.x

Genetic estimates of contemporary effective population size: what can they tell us about the importance of genetic stochasticity for wild population persistence?

An essential prerequisite of a sustainable fisheries management is the matching of biologically relevant processes and management action. In fisheries management and assessment, fish stocks are the fundamental biological unit, but the reasoning for the operational management unit is often indistinct and mismatches between the biology and the management action frequently occur. Despite the plethora of population genetic data on marine fishes, to date little or no use is made of the information, despite the fact that the detection of genetic differentiation may indicate reproductively distinct populations. Here, we discuss key aspects of genetic population differentiation in the context of their importance for fisheries management. Furthermore, we evaluate the population structure of all 32 managed marine fish species in the north-east Atlantic and relate this structure to current management units and practice. Although a large number of studies on genetic population structure have been published in the last decades, data are still rare for most exploited species. The mismatch between genetic population structure and the current management units found for six species (Gadus morhua, Melanogrammus aeglefinus, Merlangius merlangus, Micromesistius poutassou, Merluccius merluccius and Clupea harengus), emphasizes the need for a revision of these units and questions the appropriateness of current management measures. The implementation of complex and dynamic population structures into novel and less static management procedures should be a primary task for future fisheries management approaches.

Genetic population structure of marine fish : mismatch between biological and fisheries management units

Humans have a penchant for unintentionally selecting against that which they desire most. In fishes, unprecedented reductions in abundance have been associated with unprecedented changes in harvesting and aquaculture technologies. Fishing, the predominant cause of fish-population collapses, is increasingly believed to generate evolutionary changes to characters of import to individual fitness, population persistence and levels of sustainable yield. Human-induced genetic change to wild populations can also result from interactions with their domesticated counterparts. Our examination of fisheries- and farming-induced evolution includes factors that may influence the magnitude, rate and reversibility of genetic responses, the potential for shifts in reaction norms and reduced plasticity, loss of genetic variability, outbreeding depression and their demographic consequences to wild fishes. We also suggest management initiatives to mitigate the effects of fisheries- and farming-induced evolution. Ultimately, the question of whether fishing or fish farming can cause evolutionary change is moot. The key issue is whether such change is likely to have negative conservation- or socio-economic consequences. Although the study of human-induced evolution on fishes should continue to include estimates of the magnitude and rate of selection, there is a critical need for research that addresses short- and long-term demographic consequences to population persistence, plasticity, recovery and productivity.

/pdf/the-nature-of-fisheries-and-farming-induced-evolution-1nxs7w7bvd.pdf

The nature of fisheries- and farming-induced evolution.

Knowledge of geographic and temporal scales of adaptive genetic variation is crucial to species conservation, yet understanding of these phenomena, particularly in marine systems, is scant. Until recently, the belief has been that because most marine species have highly dispersive or mobile life stages, local adaptation could occur only on broad geographic scales. This view is supported by comparatively low levels of genetic variation among populations as detected by neutral markers. Similarly, the time scale of adaptive divergence has also been assumed to be very long, requiring thousands of generations. Recent studies of a variety of species have challenged these beliefs. First, there is strong evidence of geographically structured local adaptation in physiological and morphological traits. Second, the proportion of quantitative trait variation at the among-population level (QsT) is much higher than it is for neutral markers (F ST ) and these two metrics of genetic variation are poorly correlated. Third, evidence that selection is a potent evolutionary force capable of sustaining adaptive divergence on contemporary time scales is summarized. The differing spatial and temporal scales of adaptive v. neutral genetic divergence call for a new paradigm in thinking about the relationship between phenogeography (the geography of phenotypic variation) and phylogeography (the geography of lineages) in marine species. The idea that contemporary selective processes can cause fine-scale spatial and temporal divergence underscores the need for a new emphasis on Darwinian fishery science.

Spatial and temporal scales of adaptive divergence in marine fishes and the implications for conservation

The existence of biologically differentiated populations has been credited with a major role in conferring sustainability and in buffering overall productivity of anadromous fish population complexes where evidence for spatial structure is uncontroversial. Here, we describe evidence of correlated genetic and life history (spawning season linked to spawning location) differentiation in an abundant and highly migratory pelagic fish, Atlantic herring, Clupea harengus, in the North Sea (NS) and adjacent areas. The existence of genetically and phenotypically diverse stocks in this region despite intense seasonal mixing strongly implicates natal homing in this species. Based on information from genetic markers and otolith morphology, we estimate the proportional contribution by NS, Skagerrak (SKG) and Kattegat and western Baltic (WBS) fish to mixed aggregations targeted by the NS fishery. We use these estimates to identify spatial and temporal differences in life history (migratory behaviour) and habitat use among genetically differentiated migratory populations that mix seasonally. Our study suggests the existence of more complex patterns of intraspecific diversity than was previously recognized. Sustainability may be compromised if such complex patterns are reduced through generalized management (e.g. area closures) that overlooks population differences in spatial use throughout the life cycle.

E. J. Simmonds

Papers

Biocomplexity in a highly migratory pelagic marine fish, Atlantic herring