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The loss of longitudinal connectivity in regulated rivers, both up- and downstream, has been detrimental for biodiversity worldwide. While progress has been made regarding upstream fish passage solutions, many questions remain unanswered regarding downstream passage alternatives. To address these knowledge gaps, we used Atlantic salmon (S. salar) smolts to experimentally assess the guidance efficiency and passage rates produced by several common screen-and-bypass fish guidance systems. Vertical screens with horizontally oriented bars extending across a turbine intake channel at a shallow angle (angled guidance screens), combined with a single, full-depth bypass entrance at their downstream end, were on average 20% more effective and produced passage rates that were 10 times higher than screens which extended perpendicularly across a turbine intake channel with vertically oriented bars that rose gradually towards the surface (inclined guidance screens) and with a bypass at the surface, on either side of the screen. Among inclined screens, gap width was negatively associated with guidance efficiencies and the smallest gap width (15 mm) exhibited a 41% greater guidance efficiency than the largest (30 mm). Among angled screens, performance was more closely linked to construction material as metal racks produced passage rates over three times faster than flexible Kevlar netting. Overall, passage through the guidance screens, and therefore into a tentative turbine intake area, was positively associated with gap width and was twice as prevalent among the inclined relative to angled guidance screens. Ultimately, an angled guidance screen with a 30 mm gap width produced the highest guidance efficiency and passage rates (a 30% improvement over the next best screen), while an inclined screen with a 30 mm gap width produced the lowest guidance efficiencies and passage rates. These results have implications for the suitability and performance of downstream fish passage solutions at both large- and small-scale hydropower plants where passage solutions are currently lacking or inadequate.

Known as the "king of fishes," the Atlantic salmon (Salmo salar, Salmonidae) is an iconic freshwater species whose contribution to human well-being has long been recognized, as have widespread declines in its abundance, partly due to river regulation. To understand how salmon conservation has been addressed within the ecosystem services (ES) framework, we synthesized the peer-reviewed literature on ES provided by salmon in regulated rivers. We developed a search string to capture allusions to provisioning, regulating, supporting and cultural ES and assessed the results to identify knowledge gaps. The effects of hydropower on fisheries catches and on modelled populations were shown in several publications. Overall, few studies focused explicitly on ES from salmon and hydropower; this is surprising given the considerable body of literature on salmon in regulated rivers. Wild salmon as a food source and other provisioning services are less important today than historically. Because predators such as salmon are important for facilitating biodiversity by cycling nutrients and controlling food webs, there is a scope of work for future assessments of these regulating and supporting services. Few papers explicitly addressed cultural ES, despite the salmon's longstanding iconic status; this is a knowledge gap for future ES assessments in relation to hydropower. The influence of ES assessments for policy makers is growing through the Intergovernmental Panel for Biodiversity and Ecosystem Services (IPBES) and the post-2020 biodiversity strategy. Explicitly addressing ES poses an opportunity for river managers to raise awareness of aquatic conservation efforts and well-informed decision-making for sustaining ES.

Selection induced by human harvest can lead to different patterns of phenotypic change than selection induced by natural predation and could be a major driving force of evolution of wild populations. The vulnerability of individuals to angling depends on the individual decision to ingest the bait, possibly mediated by their neuroendocrine response towards the associated stimulus. To investigate the mechanisms behind individual vulnerability to angling, we conducted angling experiments in replicated ponds and quantified individual behavioral traits and neuroendocrine stress responsiveness in two salmonid species, rainbow trout (Oncorhynchus mykiss) and brown trout (Salmo trutta). We discovered a phenotypic syndrome in rainbow trout, but not in brown trout, where lower serotonergic and dopaminergic brain activity and cortisol levels (i.e., lower stress responsiveness) in response to a standardized experimental stressor were associated with higher activity, forming a proactive phenotype that showed increased vulnerability to angling. Our results show that angling targets the most stress-resilient and active phenotypes of rainbow trout, supporting the suggestion that fishing-induced phenotypic selection may lead to an increased representation of stress-responsive and low-activity phenotypes in harvested populations.

The salmonid fish Brown trout is iconic as a model for the application of conservation genetics to understand and manage local interspecific variation. However, there is still scant information about relationships between local and large-scale population structure, and to what extent geographical and environmental variables are associated with barriers to gene flow. We used information from 3,782 mapped SNPs developed for the present study and conducted outlier tests and gene–environment association (GEA) analyses in order to examine drivers of population structure. Analyses comprised >2,600 fish from 72 riverine populations spanning a central part of the species' distribution in northern Europe. We report hitherto unidentified genetic breaks in population structure, indicating strong barriers to gene flow. GEA loci were widely spread across genomic regions and showed correlations with climatic, abiotic and geographical parameters. In some cases, individual loci showed consistent GEA across the geographical regions Britain, Europe and Scandinavia. In other cases, correlations were observed only within a sub-set of regions, suggesting that locus-specific variation was associated with local processes. A paired-population sampling design allowed us to evaluate sampling effects on detection of outlier loci and GEA. Two widely applied methods for outlier detection (pcadapt and bayescan) showed low overlap in loci identified as statistical outliers across sub-sets of data. Two GEA analytical approaches (LFMM and RDA) showed good correspondence concerning loci associated with specific variables, but LFMM identified five times more statistically significant associations than RDA. Our results emphasize the importance of carefully considering the statistical methods applied for the hypotheses being tested in outlier analysis. Sampling design may have lower impact on results if the objective is to identify GEA loci and their population distribution. Our study provides new insights into trout populations, and results have direct management implications in serving as a tool for identification of conservation units.

Cost‐effective implementation of fish‐friendly hydropower flow operation and habitat restoration measures require an understanding of their effects on fitness‐related behaviours of stream fish. Here, we investigated how changes in flow and bottom structure influence the social behaviour of European grayling, using large experimental flumes (700 L s−1), with and without added boulders (i.e., restored and unrestored habitat). Grayling increased their distance to nearest neighbour at the start of flow ramping up and after a flow peak compared to stable base flow. At the start of ramping up the flow, grayling made less position changes (movements >1 m) than at stable base flow and after a flow peak. In the unrestored habitat, the proportion of time grayling spent actively swimming was lower before a flow peak than it was both at the start of ramping up the flow and after the peak, an effect not found in the restored habitat. In addition, we compared two static flows, and habitat restoration mediated their effect on distance to nearest neighbour. Grayling in the restored habitat were positioned closer to each other in the low (~10 cm s−1) than in the intermediate static flow (~40 cm s−1), whereas in the unrestored habitat, grayling showed the opposite pattern. Moreover, grayling reduced their number of position changes in the intermediate static flow, which was reflected by a reduction in active swimming. Stomach analysis after the trials revealed that foraging success was higher in variable than in the stable flow treatment. These results show that flow magnitude, flow changes and instream structure play important roles in the behaviour of stream fishes.

The salmonid fish Brown trout is iconic as a model for the application of conservation genetics to understand and manage local interspecific variation. However, there is still scant information about relationships between local and large‐scale population structure, and to what extent geographical and environmental variables are associated with barriers to gene flow. We used information from 3,782 mapped SNPs developed for the present study and conducted outlier tests and gene–environment association (GEA) analyses in order to examine drivers of population structure. Analyses comprised >2,600 fish from 72 riverine populations spanning a central part of the species' distribution in northern Europe. We report hitherto unidentified genetic breaks in population structure, indicating strong barriers to gene flow. GEA loci were widely spread across genomic regions and showed correlations with climatic, abiotic and geographical parameters. In some cases, individual loci showed consistent GEA across the geographical regions Britain, Europe and Scandinavia. In other cases, correlations were observed only within a sub‐set of regions, suggesting that locus‐specific variation was associated with local processes. A paired‐population sampling design allowed us to evaluate sampling effects on detection of outlier loci and GEA. Two widely applied methods for outlier detection (pcadapt and bayescan) showed low overlap in loci identified as statistical outliers across sub‐sets of data. Two GEA analytical approaches (LFMM and RDA) showed good correspondence concerning loci associated with specific variables, but LFMM identified five times more statistically significant associations than RDA. Our results emphasize the importance of carefully considering the statistical methods applied for the hypotheses being tested in outlier analysis. Sampling design may have lower impact on results if the objective is to identify GEA loci and their population distribution. Our study provides new insights into trout populations, and results have direct management implications in serving as a tool for identification of conservation units.

Phenotypic scoring of wild animals under standardized laboratory conditions is important as it allows field ecologists and evolutionary biologists to understand the development and maintenance of interindividual differences in plastic traits (e.g. behaviour and physiology). However, captivity is associated with a shift from a natural familiar environment to an unfamiliar and artificial environment, which may affect estimates of plastic phenotypic traits. In this study, we tested how previous experience with laboratory environments and time spent in captivity affects behavioural (i.e. activity) and metabolic (i.e. standard and maximum metabolic rates) scoring of our model species, wild brown trout Salmo trutta. We found that individuals with previous experience of laboratory captivity (10.5 months earlier) showed higher activity in an open field test than individuals with no prior experience of laboratory captivity. Previous experience with captivity had no significant effect on metabolic rates. However, metabolic rates seemed to increase with increasing time spent in captivity prior to the collection of measurements. Although there are benefits of keeping wild animals in captivity prior to scoring, our results suggest that while allowing for sufficient acclimatization researchers should aim at minimizing time in captivity of wild animals to increase accuracy and ecological relevance of the scoring of plastic phenotypic traits.

The hypothesis that interindividual differences in the activity of brown trout alter the exposure to parasitic freshwater pearl mussel glochidia was tested in a Swedish stream. Wild yearling brown trout (N = 103) were caught, individually tagged for identification and scored for open-field activity during standardized laboratory tests in June. Fifty gravid freshwater pearl mussels were relocated to the stream, where after the trout were released back into the stream. The fish were recaptured in October (N = 35), checked for glochidia encystment (infested individuals: n = 6) and re-scored for open-field activity traits. Swimming velocity during the test was higher in fish infected with glochidia, suggesting that high activity could increase their exposure to glochidia. Potentially, as metabolism rate and ventilation rate typically increase with activity, elevated activity may lead to an increased likelihood of glochidia passing over the gills. This novel finding suggests that glochidia infestation is non-random and that the behaviour of the host fish can influence the likelihood of glochidia infestation.

A new, potentially cost efficient, concept for improving the attraction flow to a fish ladder has been investigated in a case study. For the upstream migrating Atlantic salmon to reach the fish ladder and by-pass the case study hydropower plant, it must be able to localize the attraction flow where it enters the main flow from the tailrace of the hydropower plant in the so-called confluence area. Here the comparatively small and limited attraction flow from the old river channel must be improved in order to be able compete with the substantially larger main flow. The objective of the present study is to investigate the feasibility of a new concept for further improvement of the attraction flow using guiding walls forming a contraction channel. Field measurements were performed tracing tagged fish in the confluence area downstream of the case study hydropower plant in order to understand the movement pattern of the fish. Based on the results, and results from bathymetry measurements in the same area, a physical scale model was constructed where it was experimentally demonstrated that it is hydraulically feasible to construct guiding walls, forming a contraction, which accelerate the attraction flow and generate a concentrated turbulent jet with a higher velocity, while keeping the flow rate unchanged. The attraction flow penetrates about half-way (70 m) into the main flow and reaches the position where most fish are positioned according to fish position measurements and therefore potentially has a good ability to attract upstream migrating fish. There is no negative impact on the water level in the confluence area and thereby not on electricity production. It was shown that the results can be scaled up to prototype conditions and the strategy can presumably be generalized to similar flow situations, existing at other hydropower plants, allowing for improved upstream fish migration in coexistence with a sound hydropower production.

Movement activity levels of wild animals often differ consistently among individuals, reflecting different behavioral types. Previous studies have shown that laboratory-scored activity can predict several ecologically relevant characteristics. In an experiment on wild brown trout Salmo trutta, spanning from June to October, we investigated how spring swimming activity, measured in a standardized laboratory test, related to relative recapture probability in autumn. Based on laboratory activity scores, individuals clustered into 2 groups, which showed contrasting patterns in the size-dependency of their recapture probability. Size had a slightly positive effect on recapture probability for passive fish but a clear negative effect on active fish. Our results show that the population structure in a cohort, in terms of relative proportions of behavioral types in different size classes, can vary over time. The results of this study could depend on either selective mortality or migration. However, selective disappearance of individuals with specific phenotypes, regardless of the mechanism, will have implications for trout population management, such as stocking efficiency of hatchery fish with high growth rates or maintenance of fishways past migration barriers.

The deleterious effect of competition for space and food in animals increases with increasing population density. In contrast, familiarity towards conspecifics can relax the intensity of interference competition. Here, we hypothesized that familiarity towards conspecifics mitigates the effect of density-dependent growth and dispersal behaviour in territorial animals. To test this, wild-captured juvenile brown trout were subjected to two consecutive laboratory experiments. First, growth and fin erosion were measured for 40 d in a 2 × 2 factorial design manipulating density and familiarity. The density was manipulated via size of experimental tanks, while per capita food abundance and fish number was constant. All fish were subsequently exposed to an emergence test, giving them the option to leave their group and disperse to a novel unoccupied environment. The results show that familiarity increases growth and decreases the level of fin erosion (i.e. proxy of intensity of aggressive interactions). We found no significant effect of population density on growth rate. However, there was a tendency towards higher fin erosion in fish kept under high density. The growth of individuals was also affected by their size rank within the group, with the largest individuals in each group growing disproportionally faster than the rest of the group, probably due to their high social rank. However, the second and third fish in the size rank did not grow significantly faster and tended to suffer higher mortality than the rest of the group. During the emergence test, the largest individuals in the familiar groups left the shelter either as the first (six of 12 groups) or last (five of 12 groups) individual in the group, while no such pattern was observed in unfamiliar groups. Our results suggest that individuals in familiar groups receive less aggression and stress (i.e. fin damage) and grow faster than fish in unfamiliar groups. The mechanisms indicated in this laboratory study may be especially important in highly fecund organisms like fish which undergo density-dependent bottlenecks during early life.

In this paper we demonstrate that eggs from anadromous salmonines can constitute a large proportion of the dietary intake of individual juvenile Atlantic salmon Salmo salar and brown trout Salmo trutta during the spawning period in late autumn. The majority of fish older than 1 year residing on a spawning ground were found to have ingested eggs from spawning salmonines (Salmo sp.). Young-of-the-year brown trout were also able to consume eggs, at least down to a minimum fork length of 68 mm. The smallest Atlantic salmon found to have consumed eggs had a fork length of 79 mm. Larger fish were more likely to consume eggs, and consumed more eggs, than smaller individuals. The eggs of anadromous salmonines provide a rich source of marine-derived nutrients to the freshwater residing juveniles at the onset of winter. Our study highlights that these eggs may constitute an important energy input for juvenile salmonines in European anadromous populations, and could possibly influence winter survival rates and subsequent smoltification in the following spring.

Theory suggests that consistent individual differences in activity are linked to life history where high activity is associated with rapid growth, high dispersal tendency, and low survival (the pace-of-life syndrome hypothesis). We addressed this influential hypothesis by combining behavioral studies with fine-scale positional scoring in nature, estimating how individual movement strategies in brown trout (Salmo trutta) associate with fitness correlates (growth and survival) in the wild. Initial dispersal in the wild was positively related to the laboratory activity. Moreover, the growth of individuals with high laboratory activity decreased with increasing home range size, whereas the growth of individuals with lower laboratory activity increased slightly with increasing home range size. Survival in the wild was not associated with laboratory activity. Our results do not support the original pace-of-life syndrome hypothesis. As an alternative explanation, we suggest that the growth of individuals adopting a high-activity strategy is more sensitive to variation in resource abundance (indicated by home range size) than the fitness individuals adopting a more passive strategy.

Theory suggests that high activity levels in animals increase growth at the cost of increased mortality. This growth-mortality tradeoff has recently been incorporated into the wider framework of the pace-of-life syndrome (POLS) hypothesis. However, activity is often quantified only in the laboratory and on a diurnal basis, leaving open the possibility that animals manage predation risk and feeding efficiency in the wild by modulating their circadian activity rhythms. Here we investigate how laboratory activity in wild brown trout parr (Salmo trutta L.) associates with circadian activity, growth, and mortality in their natal stream. We found that individuals with high activity in the laboratory displayed high dispersal and cathemeral activity in their natal stream. In contrast, trout with low laboratory activity showed variation of activity in the wild, which was negatively related to the light intensity. Our results do not support the growth-mortality trade-off of the POLS hypothesis as highly active, fast-growing individuals showed higher survival than inactive conspecifics. These novel results show for the first time that active and inactive individuals, as scored in the lab, can show different circadian patterns of behavior in the wild driven by light intensity. This implies that studies conducted under a narrow range of light conditions can bias our understanding of individual behavioral variation and its fitness consequences in the wild.