Esben Moland Olsen

Are low but statistically significant levels of genetic differentiation in marine fishes ‘biologically meaningful’? A case study of coastal Atlantic cod

A key question in many genetic studies on marine organisms is how to interpret a low but statistically significant level of genetic differentiation. Do such observations reflect a real phenomenon, or are they caused by confounding factors such as unrepresentative sampling or selective forces acting on the marker loci? Further, are low levels of differentiation biologically trivial, or can they represent a meaningful and perhaps important finding? We explored these issues in an empirical study on coastal Atlantic cod, combining temporally replicated genetic samples over a 10‐year period with an extensive capture–mark–recapture study of individual mobility and population size. The genetic analyses revealed a pattern of differentiation between the inner part of the fjord and the open skerries area at the fjord entrance. Overall, genetic differentiation was weak (average FST = 0.0037), but nevertheless highly statistical significant and did not depend on particular loci that could be subject to selection. This spatial component dominated over temporal change, and temporal replicates clustered together throughout the 10‐year period. Consistent with genetic results, the majority of the recaptured fish were found close to the point of release, with <1% of recaptured individuals dispersing between the inner fjord and outer skerries. We conclude that low levels of genetic differentiation in this marine fish can indeed be biologically meaningful, corresponding to separate, temporally persistent, local populations. We estimated the genetically effective sizes (Ne) of the two coastal cod populations to 198 and 542 and found a Ne/N (spawner) ratio of 0.14.

An Evaluation of Visible Implant Elastomer for Marking Age-0 Brown Trout

Abstract Injection of visible implant elastomer (VIE) was evaluated as a way to provide age-0 brown trout Salmo trutta with externally visible internal marks. We first tested this fluorescent elastomer material in a laboratory experiment using a single color and body position to batch-mark the fish (28.9–44.1 mm fork length). The experiment lasted for 77 d, with no mortality or tag loss and no significant (P = 0.44) effect on fish growth. Similar marks were then used to individually tag age-0 brown trout (26–70 mm) in small streams in the wild using four different colors and several body positions. The immediate mortality associated with the marking procedure was low (0.5%), and visual identification of VIE marks on recaptured individuals (169 out of 699 released; 39–83 d between release and recapture) was not difficult. We found no significant differences in mean fork length between tagged and untagged individuals captured on the same date at given sampling locations. This suggests that capture and marki...

Small‐scale biocomplexity in coastal Atlantic cod supporting a Darwinian perspective on fisheries management

Harvesting of marine resources raises concerns about how to identify and preserve biocomplexity, including the diversity of life histories found within and among wild populations of a species. In order to fully accomplish this, there is a need to elucidate the underlying causes of phenotypic variation, and how this variation responds to environmental changes. In general, both evolutionary (genetic) and nonevolutionary (plastic) responses may occur. Plastic responses to environmental change are expected to shift the phenotype along a reaction norm, while an evolutionary response is expected to shift the reaction norm itself. Here, we assess the maturation patterns of coastal Atlantic cod (Gadus morhua) in Skagerrak, where studies using neutral markers have revealed genetically differentiated populations of this harvested fish within tens of kilometres of coastline. Our results suggest that physiological state prior to the spawning season, as well as juvenile growth, both influence the probability of completing sexual maturation at a given age. Furthermore, our results point towards a spatial structuring of this plasticity (i.e. the maturation reaction norms) comparable with population connectivity inferred from neutral markers. We argue that such fine‐scale biocomplexity calls for a Darwinian approach to fisheries management.

Spawning stock and recruitment in North Sea cod shaped by food and climate

In order to provide better fisheries management and conservation decisions, there is a need to discern the underlying relationship between the spawning stock and recruitment of marine fishes, a relationship which is influenced by the environmental conditions. Here, we demonstrate how the environmental conditions (temperature and the food availability for fish larvae) influence the stock–recruitment relationship and indeed what kind of stock–recruitment relationship we might see under different environmental conditions. Using unique zooplankton data from the Continuous Plankton Recorder, we find that food availability (i.e. zooplankton) in essence determines which model applies for the once large North Sea cod (Gadus morhua) stock. Further, we show that recruitment is strengthened during cold years and weakened during warm years. Our combined model explained 45 per cent of the total variance in cod recruitment, while the traditional Ricker and Beverton–Holt models only explained about 10 per cent. Specifically, our approach predicts that a full recovery of the North Sea cod stock might not be expected until the environment becomes more favourable.

Lobster and cod benefit from small-scale northern marine protected areas: inference from an empirical before–after control-impact study

Marine protected areas (MPAs) are increasingly implemented as tools to conserve and manage fisheries and target species. Because there are opportunity costs to conservation, there is a need for science-based assessment of MPAs. Here, we present one of the northernmost documentations of MPA effects to date, demonstrated by a replicated before–after control-impact (BACI) approach. In 2006, MPAs were implemented along the Norwegian Skagerrak coast offering complete protection to shellfish and partial protection to fish. By 2010, European lobster (Homarus gammarus) catch-per-unit-effort (CPUE) had increased by 245 per cent in MPAs, whereas CPUE in control areas had increased by 87 per cent. Mean size of lobsters increased by 13 per cent in MPAs, whereas increase in control areas was negligible. Furthermore, MPA-responses and population development in control areas varied significantly among regions. This illustrates the importance of a replicated BACI design for reaching robust conclusions and management decisions. Partial protection of Atlantic cod (Gadus morhua) was followed by an increase in population density and body size compared with control areas. By 2010, MPA cod were on average 5 cm longer than in any of the control areas. MPAs can be useful management tools in rebuilding and conserving portions of depleted lobster populations in northern temperate waters, and even for a mobile temperate fish species such as the Atlantic cod.

Small-scale genetic structure in a marine population in relation to water circulation and egg characteristics

Until the last decade it was assumed that most marine species have pronounced gene flow over vast areas, largely because of their potential for dispersal during early life stages. However, recent genetic, modeling, and field studies have shown that marine populations may be structured at scales that are inconsistent with extensive dispersal of eggs and larvae. Such findings have stimulated the birth of new studies explaining the mechanisms that promote population structure and isolation in the oceans, in the face of high potential for dispersal. Here we study the vertical and horizontal distribution of cod (Gadus morhua) eggs in relation to small-scale circulation and water column hydrography in a coastal location of southern Norway. Previous studies conducted in this region have shown that cod populations inhabiting fjord locations, which are on average 30 km apart, are genetically differentiated, a remarkable outcome considering that Atlantic cod have pelagic egg stages and long pelagic larval duration. We document that cod eggs are found in greater abundance in shallow water layers, which on average are flowing up the fjord (away from the open ocean), and in the inner portion of the fjord, which is subject to lower current speeds compared to the outer or mouth of the fjord. Eggs were found to be neutrally buoyant at shallow depths, a trait that also favors local retention, given the local circulation. The same patterns held during two environmentally contrasting years. These results strongly suggest that population structure of Atlantic cod is favored and maintained by a balance between water circulation and egg characteristics.

Harvest selection on Atlantic cod behavioral traits: implications for spatial management

Harvesting wild populations may contrast or reinforce natural agents of selection and potentially cause evolutionary changes in life-history traits such as growth and maturation. Harvest selection may also act on behavioral traits, although this field of research has so far received less attention. We used acoustic tags and a network of receivers to monitor the behavior and fate of individual Atlantic cod (Gadus morhua, N = 60) in their natural habitat on the Norwegian Skagerrak coast. Fish with a strong diel vertical migration, alternating between shallow- and deep-water habitats, had a higher risk of being captured in the fishery (traps, gillnet, hand line) as compared to fish that stayed in deeper water. There was also a significant negative correlation between fish size (30–66 cm) and the magnitude of diel vertical migration. Natural selection on behavior was less clear, but tended to favor fish with a large activity space. On a monthly time scale we found significant repeatabilities for cod behavior, meaning that individual characteristics tended to persist and therefore may be termed personality traits. We argue that an evolutionary approach to fisheries management should consider fish behavior. This would be of particular relevance for spatial management actions such as marine reserve design.

Mating patterns and determinants of individual reproductive success in brown trout (Salmo trutta) revealed by parentage analysis of an entire stream living population

Reproductive success and its determinants are difficult to infer for wild populations of species with no parental care where behavioural observations are difficult or impossible. In this study, we characterized the breeding system and provide estimates of individual reproductive success under natural conditions for an exhaustively sampled stream‐resident brown trout (Salmo trutta) population. We inferred parentage using a full probability Bayesian model that combines genetic (microsatellite) with phenotypic data. By augmenting the potential parents file with inferred parental genotypes from sib‐ship analysis in cases where large families had unsampled parents, we could make more precise inference on variance of family size. We observed both polygamous and monogamous matings and large reproductive skew for both sexes, particularly in males. Correspondingly, we found evidence for sexual selection on body size for both sexes. We show that the mating system of brown trout has the potential to be very flexible and we conjecture that environmental uncertainty could be driving the evolution and perhaps select for the maintenance of plasticity of the mating system in this species.

Climate and population density drive changes in cod body size throughout a century on the Norwegian coast

Understanding how populations respond to changes in climate requires long-term, high-quality datasets, which are rare for marine systems. We estimated the effects of climate warming on cod lengths and length variability using a unique 91-y time series of more than 100,000 individual juvenile cod lengths from surveys that began in 1919 along the Norwegian Skagerrak coast. Using linear mixed-effects models, we accounted for spatial population structure and the nested structure of the survey data to reveal opposite effects of spring and summer warming on juvenile cod lengths. Warm summer temperatures in the coastal Skagerrak have limited juvenile growth. In contrast, warmer springs have resulted in larger juvenile cod, with less variation in lengths within a cohort, possibly because of a temperature-driven contraction in the spring spawning period. A density-dependent reduction in length was evident only at the highest population densities in the time series, which have rarely been observed in the last decade. If temperatures rise because of global warming, nonlinearities in the opposing temperature effects suggest that negative effects of warmer summers will increasingly outweigh positive effects of warmer springs, and the coastal Skagerrak will become ill-suited for Atlantic cod.

Nine decades of decreasing phenotypic variability in Atlantic cod

Changes in phenotypic variability in natural populations have received little attention in comparison with changes in mean trait values. This is unfortunate because trait diversity may influence adaptive evolutionary change and population stability. We combine two unique data sets to illuminate complex trait changes in Atlantic cod along the Norwegian Skagerrak coast: (i) an annual beach seine survey starting in 1919, monitoring juvenile body size and abundance and (ii) capture-mark-recapture data from which we estimated selection on juvenile body size and growth. We demonstrate that the variability of juvenile size has been steadily decreasing across nine decades, with no evidence for a similar trend in mean size. We also report that small, slow-growing fish as well as large, fast-growing fish are selected against. Together, these results suggest long-term stabilizing selection acting on Atlantic cod, and emphasize the need for further studies evaluating the full complexity of trait changes in wild populations.