Associations between metabolic traits and growth rate in brown trout (Salmo trutta) depend on thermal regime.

Metabolism defines the energetic cost of life, yet we still know relatively little about why intraspecific variation in metabolic rate arises and persists. Spatio-temporal variation in selection potentially maintains differences, but relationships between metabolic traits (standard metabolic rate (SMR), maximum metabolic rate (MMR), and aerobic scope) and fitness across contexts are unresolved. We show that associations between SMR, MMR, and growth rate (a key fitness-related trait) vary depending on the thermal regime (a potential selective agent) in offspring of wild-sampled brown trout from two populations reared for approximately 15 months in either a cool or warm (+1.8°C) regime. SMR was positively related to growth in the cool, but negatively related in the warm regime. The opposite patterns were found for MMR and growth associations (positive in warm, negative in the cool regime). Mean SMR, but not MMR, was lower in warm regimes within both populations (i.e. basal metabolic costs were reduced at higher temperatures), consistent with an adaptive acclimation response that optimizes growth. Metabolic phenotypes thus exhibited a thermally sensitive metabolic 'floor' and a less flexible metabolic 'ceiling'. Our findings suggest a role for growth-related fluctuating selection in shaping patterns of metabolic variation that is likely important in adapting to climate change.

The spawning location of vulnerable ferox trout (Salmo trutta L.) in the Lough Corrib and Lough Mask catchments, Western Ireland.

Ferox trout are large, long-lived, piscivorous trout normally found in deep lakes; they are highly prized by trophy anglers. Lough Corrib and Lough Mask, Western Ireland, have recorded the majority of Irish specimen ferox trout since angling records began. Little was known regarding the spawning location of ferox trout relative to sympatric brown trout, and a radio telemetry study was initiated in both catchments in 2005. Over the period 2005-2009, 79 ferox were captured by angling and radio tagged in Lough Corrib, while 55 ferox were tagged in Lough Mask. Manual and helicopter tracking were carried out on all spawning streams entering both lakes over the autumn/winter period to detect tagged fish. Overall, 37 radio-tagged trout (46.8%) were detected in Lough Corrib streams and 21 tagged trout (38.2%) were recorded from Lough Mask streams. Results from radio tracking indicate that the majority (92%) of ferox trout tagged in Lough Corrib spawned in a single spawning stream, the Cong river, while the majority (76%) of ferox trout tagged in Lough Mask spawned in the Cong canal and Cong river. These results suggest that these streams are most likely the principle spawning locations of ferox trout in both lakes. The occurrence of ferox trout predominantly in single spawning rivers in both catchments highlights the vulnerability of the study ferox populations. As a result of these findings, conservation measures were introduced for ferox trout in both catchments.

Body condition of returning Atlantic salmon Salmo salar L. correlates with scale δ13 C and δ15 N content deposited at the last marine foraging location.

Patterns of feeding and growth of Atlantic salmon Salmo salar L. in the marine environment are critical to understanding how observed declines in recruitment may reflect warming or other oceanic drivers. The isotopic composition of scales can provide insight into differences in marine feeding location and possibly temperature regime. The authors used archived scale samples to measure δ13 C and δ15 N deposited in the scales of one sea-winter (1SW) salmon during their last season of growth at sea before they returned to five Irish rivers. δ13 C values were related statistically to observed salmon body condition (Fulton's K), and fish with higher δ13 C values tended to show significantly better condition. In contrast, δ15 N values were negatively related to body condition. There was no important effect on condition of length at smolt migration, and the effect of duration of marine residence varied among rivers. It is likely that δ13 C values partly reflected ambient ocean temperature and recent marine feeding environment before return migration, such that the observed relationship between higher δ13 C values and increased body condition may express an advantage for adult fish feeding in warmer, potentially closer, waters. If greater body condition influences fitness, then a changing temperature regime in the Northeast Atlantic may drive shifts in salmon survival and reproduction. This study provides evidence that there is spatial and trophic variation at sea between salmon from rivers of origin that are located relatively close to each other, with potential consequences for body condition and, consequently, fitness and life history; this suggests that salmon populations from geographically proximate rivers within regions may exhibit differential responses to ocean-scale climatic changes across the Northeast Atlantic.

Food and temperature stressors have opposing effects in determining flexible migration decisions in brown trout (Salmo trutta).

With rapid global change, organisms in natural systems are exposed to a multitude of stressors that likely co-occur, with uncertain impacts. We explored individual and cumulative effects of co-occurring environmental stressors on the striking, yet poorly understood, phenomenon of facultative migration. We reared offspring of a brown trout population that naturally demonstrates facultative anadromy (sea migration), under different environmental stressor treatments and measured life history responses in terms of migratory tactics and freshwater maturation rates. Juvenile fish were exposed to reduced food availability, temperatures elevated to 1.8°C above natural conditions or both treatments in combination over 18 months of experimental tank rearing. When considered in isolation, reduced food had negative effects on the size, mass and condition of fish across the experiment. We detected variable effects of warm temperatures (negative effects on size and mass, but positive effect on lipids). When combined with food restriction, temperature effects on these traits were less pronounced, implying antagonistic stressor effects on morphological traits. Stressors combined additively, but had opposing effects on life history tactics: migration increased and maturation rates decreased under low food conditions, whereas the opposite occurred in the warm temperature treatment. Not all fish had expressed maturation or migration tactics by the end of the study, and the frequency of these 'unassigned' fish was higher in food deprivation treatments, but lower in warm treatments. Fish showing migration tactics were smaller and in poorer condition than fish showing maturation tactics, but were similar in size to unassigned fish. We further detected effects of food restriction on hypo-osmoregulatory function of migrants that may influence the fitness benefits of the migratory tactic at sea. We also highlight that responses to multiple stressors may vary depending on the response considered. Collectively, our results indicate contrasting effects of environmental stressors on life history trajectories in a facultatively migratory species.

Unprecedented widespread occurrence of Pink Salmon Oncorhynchus gorbuscha in Ireland in 2017.

The occurrence of non-native pink salmon Oncorhynchus gorbuscha in multiple river systems in Ireland in 2017 was unprecedented and reflected similar concurrent events observed in other countries in the North Atlantic region. Thirty-six fish were captured in a total of 11 river systems located in the south-west, west and north-west of Ireland in the period June to September 2017. The biological characteristics of sampled specimens are described and the potential for establishment in Ireland are considered.

Impact of parasites on salmon recruitment in the Northeast Atlantic Ocean.

Parasites may have large effects on host population dynamics, marine fisheries and conservation, but a clear elucidation of their impact is limited by a lack of ecosystem-scale experimental data. We conducted a meta-analysis of replicated manipulative field experiments concerning the influence of parasitism by crustaceans on the marine survival of Atlantic salmon (Salmo salar L.). The data include 24 trials in which tagged smolts (totalling 283 347 fish; 1996-2008) were released as paired control and parasiticide-treated groups into 10 areas of Ireland and Norway. All experimental fish were infection-free when released into freshwater, and a proportion of each group was recovered as adult recruits returning to coastal waters 1 or more years later. Treatment had a significant positive effect on survival to recruitment, with an overall effect size (odds ratio) of 1.29 that corresponds to an estimated loss of 39 per cent (95% CI: 18-55%) of adult salmon recruitment. The parasitic crustaceans were probably acquired during early marine migration in areas that host large aquaculture populations of domesticated salmon, which elevate local abundances of ectoparasitic copepods-particularly Lepeophtheirus salmonis. These results provide experimental evidence from a large marine ecosystem that parasites can have large impacts on fish recruitment, fisheries and conservation.

Alternative migratory tactics in brown trout (Salmo trutta) are underpinned by divergent regulation of metabolic but not neurological genes.

The occurrence of alternative morphs within populations is common, but the underlying molecular mechanisms remain poorly understood. Many animals, for example, exhibit facultative migration, where two or more alternative migratory tactics (AMTs) coexist within populations. In certain salmonid species, some individuals remain in natal rivers all their lives, while others (in particular, females) migrate to sea for a period of marine growth. Here, we performed transcriptional profiling ("RNA-seq") of the brain and liver of male and female brown trout to understand the genes and processes that differentiate between migratory and residency morphs (AMT-associated genes) and how they may differ in expression between the sexes. We found tissue-specific differences with a greater number of genes expressed differentially in the liver (n = 867 genes) compared with the brain (n = 10) between the morphs. Genes with increased expression in resident livers were enriched for Gene Ontology terms associated with metabolic processes, highlighting key molecular-genetic pathways underlying the energetic requirements associated with divergent migratory tactics. In contrast, smolt-biased genes were enriched for biological processes such as response to cytokines, suggestive of possible immune function differences between smolts and residents. Finally, we identified evidence of sex-biased gene expression for AMT-associated genes in the liver (n = 12) but not the brain. Collectively, our results provide insights into tissue-specific gene expression underlying the production of alternative life histories within and between the sexes, and point toward a key role for metabolic processes in the liver in mediating divergent physiological trajectories of migrants versus residents.

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.

Metabolic traits in brown trout (Salmo trutta) vary in response to food restriction and intrinsic factors.

Metabolic rates vary hugely within and between populations, yet we know relatively little about factors causing intraspecific variation. Since metabolic rate determines the energetic cost of life, uncovering these sources of variation is important to understand and forecast responses to environmental change. Moreover, few studies have examined factors causing intraspecific variation in metabolic flexibility. We explore how extrinsic environmental conditions and intrinsic factors contribute to variation in metabolic traits in brown trout, an iconic and polymorphic species that is threatened across much of its native range. We measured metabolic traits in offspring from two wild populations that naturally show life-history variation in migratory tactics (one anadromous, i.e. sea-migratory, one non-anadromous) that we reared under either optimal food or experimental conditions of long-term food restriction (lasting between 7 and 17 months). Both populations showed decreased standard metabolic rates (SMR-baseline energy requirements) under low food conditions. The anadromous population had higher maximum metabolic rate (MMR) than the non-anadromous population, and marginally higher SMR. The MMR difference was greater than SMR and consequently aerobic scope (AS) was higher in the anadromous population. MMR and AS were both higher in males than females. The anadromous population also had higher AS under low food compared to optimal food conditions, consistent with population-specific effects of food restriction on AS. Our results suggest different components of metabolic rate can vary in their response to environmental conditions, and according to intrinsic (population-background/sex) effects. Populations might further differ in their flexibility of metabolic traits, potentially due to intrinsic factors related to life history (e.g. migratory tactics). More comparisons of populations/individuals with divergent life histories will help to reveal this. Overall, our study suggests that incorporating an understanding of metabolic trait variation and flexibility and linking this to life history and demography will improve our ability to conserve populations experiencing global change.

Ocean predation and mortality of adult Atlantic salmon.

Predation and mortality are often difficult to estimate in the ocean, which hampers the management and conservation of marine fishes. We used data from pop-up satellite archival tags to investigate the ocean predation and mortality of adult Atlantic salmon (Salmo salar) released from 12 rivers flowing into the North Atlantic Ocean. Data from 156 tagged fish revealed 22 definite predation events (14%) and 38 undetermined mortalities (24%). Endothermic fish were the most common predators (n = 13), with most of these predation events occurring in the Gulf of St. Lawrence and from the Bay of Biscay to the Irish Shelf. Predation by marine mammals, most likely large deep-diving toothed whales (n = 5), and large ectothermic fish (n = 4) were less frequent. Both the estimated predation rates (ZP) and total mortality rates (ZM) where higher for Atlantic salmon from Canada, Ireland, and Spain (ZP = 0.60-1.32 y-1, ZM = 1.73-3.08 y-1) than from Denmark and Norway (ZP = 0-0.13 y-1, ZM = 0.19-1.03 y-1). This geographical variation in ocean mortality correlates with ongoing population declines, which are more profound for southern populations, indicating that low ocean survival of adults may act as an additional stressor to already vulnerable populations.

Empirical observations of the spawning migration of European eels: The long and dangerous road to the Sargasso Sea.

The spawning migration of the European eel (Anguilla anguilla L.) to the Sargasso Sea is one of the greatest animal migrations. However, the duration and route of the migration remain uncertain. Using fishery data from 20 rivers across Europe, we show that most eels begin their oceanic migration between August and December. We used electronic tagging techniques to map the oceanic migration from eels released from four regions in Europe. Of 707 eels tagged, we received 206 data sets. Many migrations ended soon after release because of predation events, but we were able to reconstruct in detail the migration routes of >80 eels. The route extended from western mainland Europe to the Azores region, more than 5000 km toward the Sargasso Sea. All eels exhibited diel vertical migrations, moving from deeper water during the day into shallower water at night. The range of migration speeds was 3 to 47 km day-1. Using data from larval surveys in the Sargasso Sea, we show that spawning likely begins in December and peaks in February. Synthesizing these results, we show that the timing of autumn escapement and the rate of migration are inconsistent with the century-long held assumption that eels spawn as a single reproductive cohort in the springtime following their escapement. Instead, we suggest that European eels adopt a mixed migratory strategy, with some individuals able to achieve a rapid migration, whereas others arrive only in time for the following spawning season. Our results have consequences for eel management.