Local variation in stress response of juvenile anadromous brown trout, Salmo trutta.

Habitat fragmentation may cut off anadromous salmonids from parts of their potential native habitat and separate previously connected populations. Understanding the consequences of this is vital for fish management and prioritization of restoration activities. Here, we show that there is a significant difference in the body morphology, physiological stress response, and aspects contributing to aerobic capacity between juvenile anadromous brown trout, Salmo trutta, collected at a downstream site and an upstream site, separated by 2 km and several challenging stream sections, in a small unfragmented stream system in western Sweden. Following a standardized stress test, there were significant differences between fish from the upstream and downstream sites (plasma cortisol concentration, plasma osmolality, hematocrit, hemoglobin concentration, and mean corpuscular hemoglobin concentration). Plasma glucose concentration did not significantly differ between fish from the two sites. Fish from the upstream site had larger spleen mass, although there was no evidence of differences in ventricle mass or proportion of compact ventricular myocardium. These physiological differences indicate local variation in stress response and highlight the importance of considering local trait variation in river management. If a section of the river becomes fragmented or degraded, and there are differences in the juveniles in different parts of the river, the consequence for the population might be larger than the proportional loss of habitat.

Variation in phenotypic plasticity across age-at-maturity genotypes in wild Atlantic salmon.

Evolution of phenotypic plasticity requires genotype-environment interaction. The discovery of two large-effect loci in the vgll3 and six6 genomic regions associated with the number of years the Atlantic salmon spend feeding at sea before maturation (sea age), provides a unique opportunity to study evolutionary potential of phenotypic plasticity. Using data on 1246 Atlantic salmon caught in the River Surna in Norway, we show that variation in mean sea age among years (smolt cohorts 2013-2018) is influenced by genotype frequencies as well as interaction effects between genotype and year. Genotype-year interactions suggest that genotypes may differ in their response to environmental variation across years, implying genetic variation in phenotypic plasticity. Our results also imply that plasticity in sea age will evolve as an indirect response to selection on mean sea age due to a shared genetic basis. Furthermore, we demonstrate differences between years in the additive and dominance functional genetic effects of vgll3 and six6 on sea age, suggesting that evolutionary responses will vary across environments. Considering the importance of age at maturity for survival and reproduction, genotype-environment interactions likely play an important role in local adaptation and population demography in Atlantic salmon.

Temporal shifts in the marine feeding of individual Atlantic salmon inferred from scale isotope ratios.

Given the limited information on prey use during the marine residency period for Atlantic salmon, scales were collected from salmon at return to the River Namsen (Norway) for spawning after 1 year at sea, and scale material from the first and second summer marine feeding periods was analysed using stable isotope methods to understand dynamics of their trophic ecology. As the salmon increased in size from the first to second summer, they reduced their feeding niche and specialised more (narrowed the δ13C range) and increased their dependency on higher tropic level (δ15N) prey, likely fish. Changes in δ13C indicated a consistent pattern of movement towards the north and west between summer feeding periods. Hence, salmon during their first year at sea may have a migration route roughly resembling that of previous spawners, as inferred from earlier tagging studies. Feeding conditions and nutrient composition during the last summer at sea, i.e. in the months before returning to the river for spawning, impacted final body size and within-season timing of return. Fish undergoing the largest trophic niche shift (δ13C and δ15N combined) between summer feeding periods, returned earliest. The earliest returning fish had the fastest specific growth rates at sea. Hence, salmon encountering abundant high-quality fish food during the marine migration, particularly during the last months, may reach a size and energetic state whereby it is better to return early to a safer environment in freshwater than risk being eaten by a big predator at sea. Both trophic status (δ15N), resource use (δ13C) and growth rates were significantly correlated between feeding periods. Nutrient composition during the first summer at sea did not impact the fish body length after the following winter, but growth conditions during the first summer evidenced carry-over effects from the first to the second summer of feeding.

Ecological regime shift in the Northeast Atlantic Ocean revealed from the unprecedented reduction in marine growth of Atlantic salmon.

Ecological regime shifts are abrupt changes in the structure and function of ecosystems that persist over time, but evidence of contemporary regime shifts are rare. Historical scale data from 52,384 individual wild Atlantic salmon caught in 180 rivers from 1989 to 2017 reveal that growth of Atlantic salmon across the Northeast Atlantic Ocean abruptly decreased following the year 2004. At the same time, the proportion of early maturing Atlantic salmon decreased. These changes occurred after a marked decrease in the extent of Arctic water in the Norwegian Sea, a subsequent warming of spring water temperature before Atlantic salmon entering the sea, and an approximately 50% reduction of zooplankton across large geographic areas of the Northeast Atlantic Ocean. A sudden decrease in growth was also observed among Atlantic mackerel in the Norwegian Sea. Our results point toward an ecosystem-scale regime shift in the Northeast Atlantic Ocean.

Early-season brown trout (Salmo trutta) migrants grow and survive better at sea.

The timing of seaward migration is a key life-history trait for many anadromous fish species, with growth and survival at sea depending on a match/mismatch scenario between the timing of the sea entry and optimal conditions. Based on a 25-year study with 15,226 individually tagged brown trout (Salmo trutta) in a Norwegian river, we analysed how the within-season timing of sea migration impacted growth and survival. In both first-time and veteran migrants, marine growth was highest for early migrating individuals, large individuals, and those with a low condition factor when entering the sea. Survival was highest for individuals entering the sea early in the season. In first-time migrants, survival increased with body length. Survival also increased with the number of other smolts migrating simultaneously. As the early smolts were the most successful, it may seem strange that many smolts migrate later in the season. We suggest that late-migrating smolts may not be of a size and/or physiological state suitable for early marine conditions, and may make the best of a bad situation.

Global trends in aquatic animal tracking with acoustic telemetry.

Acoustic telemetry (AT) is a rapidly evolving technique used to track the movements of aquatic animals. As the capacity of AT research expands it is important to optimize its relevance to management while still pursuing key ecological questions. A global review of AT literature revealed region-specific research priorities underscoring the breadth of how AT is applied, but collectively demonstrated a lack of management-driven objectives, particularly relating to fisheries, climate change, and protection of species. In addition to the need for more research with direct pertinence to management, AT research should prioritize ongoing efforts to create collaborative opportunities, establish long-term and ecosystem-based monitoring, and utilize technological advancements to bolster aquatic policy and ecological understanding worldwide.

Growth of invasive pink salmon (Oncorhynchus gorbuscha) at sea assessed by scale analysis.

Invasive pink salmon (Oncorhynchus gorbuscha) has been present in variable, but low, numbers in Norwegian waters since c. 1960, but beginning in 2017 their numbers have exploded in rivers in northern Norway, with considerable numbers also recorded in rivers in southern Norway and other countries bordering the North Atlantic. Analysis of pink salmon scales from two rivers draining to the western Barents Sea showed declining growth during the first weeks after entering the sea, and some individuals even showed a pronounced growth arrest, based on detailed scale circulus analyses. This was followed by a period of growth increase and stability during late summer and autumn, which may reflect a transition to better food sources, as the fish migrate from coastal waters to the open ocean, and as they grow larger and can eat larger and more energy efficient food items. Growth declined to a minimum during winter. Fish body size at spawning was positively correlated with the distance from scale focus to the last winter circulus, as well as with the number of circuli. When dividing scale growth into three periods, better growth during the first period at sea was related to increased fish body length at spawning, but this early growth explained only a minor part (6%) of the variation in final body length. The reason for this may be large individual variation in growth combined with large mortality during the first weeks at sea. If mortality is selective, removing fish with poor growth may reduce a correlation between early growth and body size at spawning. Scale growth during late summer and early autumn explained more of the variation in fish length at spawning (27%). Hence, late summer and early autumn was likely an important period for marine growth and survival in the invasive pink salmon.

Redefining the oceanic distribution of Atlantic salmon.

Determining the mechanisms driving range-wide reductions in Atlantic salmon marine survival is hindered by an insufficient understanding of their oceanic ecology and distribution. We attached 204 pop-up satellite archival tags to post-spawned salmon when they migrated to the ocean from seven European areas and maiden North American salmon captured at sea at West Greenland. Individuals migrated further north and east than previously reported and displayed increased diving activity near oceanographic fronts, emphasizing the importance of these regions as feeding areas. The oceanic distribution differed among individuals and populations, but overlapped more between geographically proximate than distant populations. Dissimilarities in distribution likely contribute to variation in growth and survival within and among populations due to spatio-temporal differences in environmental conditions. Climate-induced changes in oceanographic conditions will alter the location of frontal areas and may have stock-specific effects on Atlantic salmon population dynamics, likely having the largest impacts on southern populations.

Keeping close to the river, shore and surface: the first marine migration of brown trout (Salmo trutta) and Arctic charr (Salvelinus alpinus) post-smolts.

Acoustic telemetry was utilized to track 49 brown trout (Salmo trutta) and 37 Arctic charr (Salvelinus alpinus) first-time migrants of wild origin [post-smolts; mean LF (fork length): 169 and 172 mm] in a large fjord in northern Norway. The S. trutta were registered at sea for more than twice the time of the S. alpinus (medians of 54 and 22 days, respectively). Both species were mostly detected near river mouths (>80% of detections) and almost exclusively spent their time (>95%) within the interior 18 km of the fjord. They were surface oriented, with most detections at <1 m depth and S. trutta deeper on average (median mean depths of 0.7 and 0.5 m, respectively). This study concludes that post-smolts of both species stay closer to the surface and to river mouths than larger veteran migrants. This study emphasizes the importance of river mouths and upper water layers for the survival of both species during their first marine migration.

Repeatable individual variation in migration timing in two anadromous salmonids and ecological consequences.

Consistent individual differences in behavior have been demonstrated for many animals, but there are few studies of consequences of such repeated behavior in the wild. We tested consistency in migration timing to and from the sea among anadromous Arctic char (Salvelinus alpinus) and brown trout (Salmo trutta), using data from a study period of about 25 years, including more than 27,000 uniquely Carlin-tagged individuals that migrated to sea for feeding in the spring and returned to the river in late summer for up to 13 successive years. Consistency was found between individuals across time in timing of the seaward migration. Individuals migrating early during their first migration tended to migrate early the following years, and late migrants tended to migrate late. The same pattern was found also at ascent to freshwater. Hence, this study demonstrated that individual fish in nature can differ in behavior related to migration timing and that these differences can be consistent during their lifetime. Early migrants increased their mass more than late migrants and had a higher specific growth rate. Early migrating Arctic char, but not brown trout, experienced a longer life after the first migration to sea than late migrants. In both species, maturity occurred earlier in individuals that migrated early. For brown trout, but not for Arctic char, fecundity was significantly correlated to the timing of smolt migration. Hence, the repeatable individual variation in migration timing seemed to have ecological and fitness consequences in terms of growth, longevity, timing of maturity, and lifetime fecundity.

Age of European silver eels during a period of declining abundance in Norway.

The European eel (Anguilla anguilla) is critically endangered throughout its range. Knowledge about age distribution of future spawners (silver eels) is essential to monitor the status and contribute to the recovery of this species. Determination of age in anguillid eels is challenging, especially in eels from the northern part of the distribution area where growth is slow and age at maturation can be up to 30 years or more. Eels from the river Imsa in Norway have been monitored since 1975, and this reference time series has been used to assess the stock at the European level. Population dynamics in this catchment were analyzed during the late 1980s by estimating ages on whole cleared otoliths. However, techniques for revealing annual increments on otoliths have evolved over the years sometimes yielding significant differences in age estimates. In this study, the historical otolith data were reanalyzed using a grinding and polishing method rather than reading the whole otolith. The new age estimates were considerably higher than the previous ones, sometimes by up to 29 years. Since the 1980s, mean age of silver eels only slightly increased (from 19 to 21 years in the 2010s). This was mainly due to the disappearance of younger silver eels (<15 years) in the 2010s. The new age estimates agreed with the steep decline in recruitment which occurred in the late 1980s in the Imsa catchment. Mean growth (30 mm/year, min-max: 16-64 mm/year) has not changed since the 1980s, although density in the catchment has decreased. Revealing and reading age of slow-growing eels remain a challenge but adding a measure of otolith reading uncertainty may improve age data collection and contribute to recovery measures for this species.

Migration patterns of a potamodromous piscivore, asp (Leuciscus aspius), in a river-lake system.

Potamodromous fishes require safe migration routes between spawning, feeding and wintering habitats to complete their life cycle. As knowledge on asp migrations is restricted, this work investigated the movements of adult asp tagged with acoustic transmitters for 3 years in the large Peipsi-Emajõgi-Võrtsjärv lake-river system, Estonia, which is free of migration barriers. Asp showed complex migration patterns, moving between and within different waterbodies (lakes, river, tributaries) in all seasons, but with a tendency to repeat habitat use patterns between years. Lakes were mainly used for feeding during spring and summer (after spawning 65% of the fish migrated to Lake Peipsi), and more so by large fish. The majority (80-96%) of the fish spent the winter in the rivers, mostly close to their subsequent spawning area. Spawning areas were in swift-flowing waters in tributaries and the main river. The results indicate that asp may benefit from an extensive and diverse complex of habitats, and any migration barrier during any season may restrict the natural habitat use by asp. Maintenance and restoration of habitat heterogeneity and connectivity is critical to protect behaviourally diverse fish populations and increase resilience in rivers negatively impacted by various human activities.

Effects of sound exposure from a seismic airgun on heart rate, acceleration and depth use in free-swimming Atlantic cod and saithe.

Airguns used for offshore seismic exploration by the oil and gas industry contribute to globally increasing anthropogenic noise levels in the marine environment. There is concern that the omnidirectional, high intensity sound pulses created by airguns may alter fish physiology and behaviour. A controlled short-term field experiment was performed to investigate the effects of sound exposure from a seismic airgun on the physiology and behaviour of two socioeconomically and ecologically important marine fishes: the Atlantic cod (Gadus morhua) and saithe (Pollachius virens). Biologgers recording heart rate and body temperature and acoustic transmitters recording locomotory activity (i.e. acceleration) and depth were used to monitor free-swimming individuals during experimental sound exposures (18-60 dB above ambient). Fish were held in a large sea cage (50 m diameter; 25 m depth) and exposed to sound exposure trials over a 3-day period. Concurrently, the behaviour of untagged cod and saithe was monitored using video recording. The cod exhibited reduced heart rate (bradycardia) in response to the particle motion component of the sound from the airgun, indicative of an initial flight response. No behavioural startle response to the airgun was observed; both cod and saithe changed both swimming depth and horizontal position more frequently during sound production. The saithe became more dispersed in response to the elevated sound levels. The fish seemed to habituate both physiologically and behaviourally with repeated exposure. In conclusion, the sound exposures induced over the time frames used in this study appear unlikely to be associated with long-term alterations in physiology or behaviour. However, additional research is needed to fully understand the ecological consequences of airgun use in marine ecosystems.

Timing and pattern of annual silver eel migration in two European watersheds are determined by similar cues.

Many animals perform long-distance migrations in order to maximize lifetime reproductive success. The European eel migrates several thousand kilometers between their feeding habitats in continental waters (fresh-, brackish, and sea water) and their spawning area in the Sargasso Sea. Eels residing in freshwaters usually initiate their spawning migration as silver eels during autumn, triggered by diverse environmental cues. We analyzed the time series of silver eel downstream migration in Burrishoole, Ireland (1971-2015), and Imsa, Norway (1975-2015), to examine factors regulating the silver eel migration from freshwater to the sea. The migration season (90% of the run) generally lasted from 1 August to 30 November. Environmental factors acting in the months before migration impacted timing and duration of migration, likely through influencing the internal processes preparing the fish for migration. Once the migration had started, environmental factors impacted the day-to-day variation in number of migrants, apparently stimulating migration among those eels ready for migration. Both the day-to-day variation in the number of migrants and the onset of migration were described by nearly identical models in the two rivers. Variables explaining day-to-day variation were all associated with conditions that may minimize predation risk; number of migrants was reduced under a strong moon and short nights and increased during high and increasing water levels. Presence of other migrants stimulated migration, which further indicates that silver eel migration has evolved to minimize predation risk. The onset of migration was explained mainly by water levels in August. The models for duration of the migration season were less similar between the sites. Thus, the overall migration season seems governed by the need to reach the spawning areas in a synchronized manner, while during the actual seaward migration, antipredator behavior seems of overriding importance.

Influence of harvest restrictions on angler release behaviour and size selection in a recreational fishery.

Fishing regulations such as harvest restrictions are implemented to limit the exploitation of many fish stocks and ensure the sustainability of fisheries. In Norway, inland recreational fisheries are co-managed by the government and by local riparian rights holders, meaning that Atlantic salmon Salmo salar harvest restrictions differ somewhat among rivers. Data from Norwegian rivers from 2009 to 2013 were used to test for variation in the proportion of salmon released by anglers and the relative size of salmon harvested and released by anglers in rivers that had varying harvest restrictions in terms of quotas, size restrictions, and/or female harvest restrictions. The proportion of the catch released by anglers was higher in rivers where there were harvest restrictions (proportion released = 0.09-0.24) than in rivers with no such restrictions (proportion released = 0.01). On average, salmon released in rivers with size restrictions larger (average mass difference between harvested and released salmon = -1.25 kg) than those released in rivers without harvest restrictions (difference = 0.60 kg). The proportion of the catch released was larger in rivers with seasonal quotas (0.29) than in rivers with daily (0.07) or collective (i.e. total catch for the river; 0.06) quotas. Rivers with low daily (one salmon per angler per day) or seasonal (<5 salmon per angler per year) quotas had a larger proportion of salmon released (0.23, 0.38, respectively) than rivers with moderate (0.10, 0.21) or high (0.07, 0.16) quotas. High seasonal quotas resulted in larger individuals harvested than released (difference = 1.16 kg), on average, compared to moderate (1.22 kg) and high seasonal quotas (-0.30 kg). We conclude that harvest restrictions influenced the extent to which fish were released and thus the stock composition (i.e. size distribution) escaping the recreational fishery with the potential to spawn.

Spatial and temporal genetic structure of a river-resident Atlantic salmon (Salmo salar) after millennia of isolation.

The river-resident Salmo salar ("småblank") has been isolated from other Atlantic salmon populations for 9,500 years in upper River Namsen, Norway. This is the only European Atlantic salmon population accomplishing its entire life cycle in a river. Hydropower development during the last six decades has introduced movement barriers and changed more than 50% of the river habitat to lentic conditions. Based on microsatellites and SNPs, genetic variation within småblank was only about 50% of that in the anadromous Atlantic salmon within the same river. The genetic differentiation (F ST) between småblank and the anadromous population was 0.24. This is similar to the differentiation between anadromous Atlantic salmon in Europe and North America. Microsatellite analyses identified three genetic subpopulations within småblank, each with an effective population size Ne of a few hundred individuals. There was no evidence of reduced heterozygosity and allelic richness in contemporary samples (2005-2008) compared with historical samples (1955-56 and 1978-79). However, there was a reduction in genetic differentiation between sampling localities over time. SNP data supported the differentiation of småblank into subpopulations and revealed downstream asymmetric gene flow between subpopulations. In spite of this, genetic variation was not higher in the lower than in the upper areas. The meta-population structure of småblank probably maintains genetic variation better than one panmictic population would do, as long as gene flow among subpopulations is maintained. Småblank is a unique endemic island population of Atlantic salmon. It is in a precarious situation due to a variety of anthropogenic impacts on its restricted habitat area. Thus, maintaining population size and avoiding further habitat fragmentation are important.

Crossing invisible boundaries: the effectiveness of the Langebaan Lagoon marine protected area as a harvest refuge for a migratory fish species in South Africa.

The application of no-take areas in fisheries remains controversial. Critics argue that many targeted species are too mobile to benefit from area protection and that no-take areas are only appropriate for resident species. The degree of protection does not depend on the size of the no-take area but rather on the time fish reside inside its boundaries during key life-history events (i.e., spawning) and during periods of peak fishing activity. We evaluated the potential of a small no-take marine protected area (MPA) inside a coastal embayment as a harvest refuge for a mobile, possibly migratory, long-lived fish species. We used acoustic telemetry to track movements of 30 transmitter-tagged white stumpnose (Rhabdosargus globiceps) across and on both sides of the boundary of a small (34 km(2)) no-take area over a full year. Being landlocked on 3 sides, the location of the MPA inside the lagoon made it practical to detect all boundary crossings and to calculate the time individual fish used the MPA. We detected frequent movements across the boundary, with strong seasonal and individual variations. There were significant differences in MPA use patterns between fish from different release areas. The time spent in the MPA by individual fish during summer (mean 50%; max 98%) was out of proportion with the size of that area (4% of total habitat). Summer coincided with peak recreational fishing activity and with the spawning season of this species. The small MPA provided a refuge for a part of the spawning stock of white stumpnose. Our findings suggest that if strategically placed, a small no-take area can be effective in protecting mobile species and that models of spillover from no-take areas should account for seasonal and individual variation in area use and the spatiotemporal distribution of fish and fishers.