Autumn outmigrants in brown trout (Salmo trutta) are not a demographic dead-end.

Genetic identity analysis and PIT (passive integrated transponder) tagging were used to examine the freshwater return rates and phenotypic characteristics of n = 1791 downstream migrating juvenile Salmo trutta in the Burrishoole catchment (northwest Ireland) across the period September 2017 to December 2020. In this system, juveniles out-migrate (move from freshwater into brackish or marine habitats) in every month of the year, with distinct seasonal peaks in spring (March through June; mostly silvered smolts) and autumn (September through December; mostly younger, unsilvered fry or parr). Both types exhibited a sex-bias towards females, which was stronger in spring (78% females) than in autumn outmigrants (67%). Sixty-nine returning fish were matched back to previous juvenile outmigrants, and similar return rates were found for spring outmigrants (5.0%), autumn outmigrants (3.3%) and fish that out-migrated outside of spring or autumn (2.8%). Spring and autumn outmigrants returned at similar dates (typically mid to late July), but autumn fish were away for longer periods (median = 612 days; spring outmigrants = 104 days). Autumn outmigrants were 25% smaller than spring outmigrants at outmigration and 6% smaller on their return, and within both groups smaller/younger outmigrants spent longer away than larger/older outmigrants. Autumn outmigrants were more likely to return unsilvered as "slob" trout (84%) than spring outmigrants (31%), suggesting they make greater use of brackish habitats that might be safer, but less productive, than fully marine habitats. Nonetheless, both types also produced silvered "sea trout" (≥1+ sea-age), implying neither is locked into a single life-history strategy. The findings emphasise that autumn outmigrants and the transitional habitats that support their persistence should not be overlooked in salmonid management and conservation.

Long-term changes and effects of significant fishery closures on marine survival and biological characteristics of wild and hatchery-reared Atlantic salmon Salmo salar.

Long-term data, over four decades, were analysed to examine temporal trends in survival indices and phenotypic characteristics of Atlantic salmon Salmo salar returning to the Burrishoole national salmonid monitored river in Ireland. Before 2007, the marine drift net fishery was the major capture method for salmon in Irish home waters, accounting for over 70% of the commercial catch and targeting mixed stocks from multiple rivers. The authors examined size differences in fish captured in marine and freshwater environments and the impact of closure of this fishery on long-term survival indices and fish size. Return rates to Irish home waters for wild one sea-winter (1SW) and a ranching strain of hatchery-reared 1SW Atlantic salmon stocks showed a declining trend up to the time of closure of the fishery (1985-2006). In contrast, closure of the drift net fishery resulted in the anticipated increase in return rate to fresh water in the short term. Nonetheless, the short-term upward trend was not sustained in the following years: the trend for return rate to fresh water (1985-2017) was found to be neither increasing nor decreasing. Mean return rates to fresh water 10 years pre- and post-closure of the drift net fishery increased from 7.4% to 8.5% for wild 1SW and significantly from 2.4% to 3.7% for ranched 1SW suggesting some benefit had accrued as a consequence of drift net closure. For ranched 1SW salmon, entry into fresh water was found to be occurring earlier, which is likely a phenotypical response to changing climatic conditions. A declining trend in fish length was found in the pre-closure period, followed by a more stable trend post-closure. Similar patterns were observed for fish condition and weight parameters. Significantly, a step change in fish size occurred just before the closure of the Irish drift net fishery in both marine and freshwater habitats, when the average length decreased by 3.8 and 4.6 cm, respectively, between 2005 and 2006. This suggests an environmental effect on the population, rather than a fishery closure effect. Similar trends in fish length were observed in wild 1SW salmon kelts and ranched 2SW salmon in fresh water. The stable but not increasing trends post-closure suggest that conditions at sea may not be improving. These findings show that a clear decline occurred in wild and ranched salmon populations' return rates and lengths, while the drift net fishery was still active. Closure of the fishery did not result in a rebound to pre-exploitation levels of these indicators. Nonetheless, the trends went from declining to stable, suggesting the closure helped mitigate the impact of unfavourable environmental and rearing habitat conditions. These findings, based on four decades of data, highlight the urgency of strengthening monitoring of fisheries populations in face of climate change, so as to guide precautionary management measures that, as this study suggests, may be able to mitigate its impacts.

Domestication-induced reduction in eye size revealed in multiple common garden experiments: The case of Atlantic salmon (Salmo salar L.).

Domestication leads to changes in traits that are under directional selection in breeding programmes, though unintentional changes in nonproduction traits can also arise. In offspring of escaping fish and any hybrid progeny, such unintentionally altered traits may reduce fitness in the wild. Atlantic salmon breeding programmes were established in the early 1970s, resulting in genetic changes in multiple traits. However, the impact of domestication on eye size has not been studied. We measured body size corrected eye size in 4000 salmon from six common garden experiments conducted under artificial and natural conditions, in freshwater and saltwater environments, in two countries. Within these common gardens, offspring of domesticated and wild parents were crossed to produce 11 strains, with varying genetic backgrounds (wild, domesticated, F1 hybrids, F2 hybrids and backcrosses). Size-adjusted eye size was influenced by both genetic and environmental factors. Domesticated fish reared under artificial conditions had smaller adjusted eye size when compared to wild fish reared under identical conditions, in both the freshwater and marine environments, and in both Irish and Norwegian experiments. However, in parr that had been introduced into a river environment shortly after hatching and sampled at the end of their first summer, differences in adjusted eye size observed among genetic groups were of a reduced magnitude and were nonsignificant in 2-year-old sea migrating smolts sampled in the river immediately prior to sea entry. Collectively, our findings could suggest that where natural selection is present, individuals with reduced eye size are maladapted and consequently have reduced fitness, building on our understanding of the mechanisms that underlie a well-documented reduction in the fitness of the progeny of domesticated salmon, including hybrid progeny, in the wild.

Captive-bred Atlantic salmon released into the wild have fewer offspring than wild-bred fish and decrease population productivity.

The release of captive-bred animals into the wild is commonly practised to restore or supplement wild populations but comes with a suite of ecological and genetic consequences. Vast numbers of hatchery-reared fish are released annually, ostensibly to restore/enhance wild populations or provide greater angling returns. While previous studies have shown that captive-bred fish perform poorly in the wild relative to wild-bred conspecifics, few have measured individual lifetime reproductive success (LRS) and how this affects population productivity. Here, we analyse data on Atlantic salmon from an intensely studied catchment into which varying numbers of captive-bred fish have escaped/been released and potentially bred over several decades. Using a molecular pedigree, we demonstrate that, on average, the LRS of captive-bred individuals was only 36% that of wild-bred individuals. A significant LRS difference remained after excluding individuals that left no surviving offspring, some of which might have simply failed to spawn, consistent with transgenerational effects on offspring survival. The annual productivity of the mixed population (wild-bred plus captive-bred) was lower in years where captive-bred fish comprised a greater fraction of potential spawners. These results bolster previous empirical and theoretical findings that intentional stocking, or non-intentional escapees, threaten, rather than enhance, recipient natural populations.

Assessment of the quality of European silver eels and tentative approach to trace the origin of contaminants - A European overview.

The European eel is critically endangered. Although the quality of silver eels is essential for their reproduction, little is known about the effects of multiple contaminants on the spawning migration and the European eel management plan does not take this into account. To address this knowledge gap, we sampled 482 silver eels from 12 catchments across Europe and developed methods to assess three aspects of eel quality: muscular lipid content (N = 169 eels), infection with Anguillicola crassus (N = 482), and contamination by persistent organic pollutants (POPs, N = 169) and trace elements (TEs, N = 75). We developed a standardized eel quality risks index (EQR) using these aspects for the subsample of 75 female eels. Among 169 eels, 33% seem to have enough muscular lipids content to reach the Sargasso Sea to reproduce. Among 482 silver eels, 93% were infected by A. crassus at least once during their lifetime. All contaminants were above the limit of quantification, except the 1,2-bis(2,4,6-tribromophenoxy)ethane (BTBPE), Ag and V. The contamination by POPs was heterogeneous between catchments while TEs were relatively homogeneous, suggesting a multi-scale adaptation of management plans. The EQR revealed that eels from Warwickshire were most impacted by brominated flame-retardants and agricultural contaminants, those from Scheldt were most impacted by agricultural and construction activities, PCBs, coal burning, and land use, while Frémur eels were best characterized by lower lipid contents and high parasitic and BTBPE levels. There was a positive correlation between EQR and a human footprint index highlighting the capacity of silver eels for biomonitoring human activities and the potential impact on the suitability of the aquatic environment for eel population health. EQR therefore represents a step forward in the standardization and mapping of eel quality risks, which will help identify priorities and strategies for restocking freshwater ecosystems.

Impacts of a record-breaking storm on physical and biogeochemical regimes along a catchment-to-coast continuum.

The impacts of changes in climate are often most readily observed through the effects of extremes in local weather, effects that often propagate through multiple ecosystem levels. Precise effects of any extreme weather event depend not only on the type of event and its timing, but also on the ecosystem affected. Here the cascade of effects following the arrival of an atmospheric river (directed by record-breaking Storm Desmond) across terrestrial, freshwater and coastal zones is quantified, using the Burrishoole system on the Atlantic coast of Ireland as a natural observatory. We used a network of high-frequency in-situ sensors to capture in detail the effects of an unprecedented period of rainfall, high wind speeds and above-average winter air temperatures on catchment and estuarine dynamics. In the main freshwater lake, water clarity decreased and acidity increased during Storm Desmond. Surface heat input, due to a warm and moist above-lake air mass, was rapidly distributed throughout the water column. River discharge into the downstream coastal basin was estimated to be the highest on record (since 1976), increasing the buoyancy flux by an order of magnitude and doubling the water column stratification stability. Entrainment of salt into the outflowing freshwater plume exported resident salt from the inner estuarine basin, resulting in net salt loss. Here, the increased stratification markedly reinforced isolation of the bottom waters, promoting deoxygenation. Measurements of current between the inner estuarine basin and the adjacent coastal waters indicated a 20-fold increase in the volume of seaward flowing low-salinity water, as a result of storm rainfall over the watershed. Storm impacts spanned the full catchment-to-coast continuum and these results provide a glimpse into a potential future for hydrological systems where these severe hydroclimatic events are expected to occur more frequently.

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.

Telemetry and genetics reveal asymmetric dispersal of a lake-feeding salmonid between inflow and outflow spawning streams at a microgeographic scale.

The degree of natal philopatry relative to natal dispersal in animal populations has important demographic and genetic consequences and often varies substantially within species. In salmonid fishes, lakes have been shown to have a strong influence on dispersal and gene flow within catchments; for example, populations spawning in inflow streams are often reproductively isolated and genetically distinct from those spawning in relatively distant outflow streams. Less is known, however, regarding the level of philopatry and genetic differentiation occurring at microgeographic scales, for example, where inflow and outflow streams are separated by very small expanses of lake habitat. Here, we investigated the interplay between genetic differentiation and fine-scale spawning movements of brown trout between their lake-feeding habitat and two spawning streams (one inflow, one outflow, separated by <100 m of lake habitat). Most (69.2%) of the lake-tagged trout subsequently detected during the spawning period were recorded in just one of the two streams, consistent with natal fidelity, while the remainder were detected in both streams, creating an opportunity for these individuals to spawn in both natal and non-natal streams. The latter behavior was supported by genetic sibship analysis, which revealed several half-sibling dyads containing one individual that was sampled as a fry in the outflow and another that was sampled as fry in the inflow. Genetic clustering analyses in conjunction with telemetry data suggested that asymmetrical dispersal patterns were occurring, with natal fidelity being more common among individuals originating from the outflow than the inflow stream. This was corroborated by Bayesian analysis of gene flow, which indicated significantly higher rates of gene flow from the inflow into the outflow than vice versa. Collectively, these results reveal how a combination of telemetry and genetics can identify distinct reproductive behaviors and associated asymmetries in natal dispersal that produce subtle, but nonetheless biologically relevant, population structuring at microgeographic scales.

Evolutionary stasis of a heritable morphological trait in a wild fish population despite apparent directional selection.

Comparing observed versus theoretically expected evolutionary responses is important for our understanding of the evolutionary process, and for assessing how species may cope with anthropogenic change. Here, we document directional selection for larger female size in Atlantic salmon, using pedigree-derived estimates of lifetime reproductive success as a fitness measure. We show the trait is heritable and, thus, capable of responding to selection. The Breeder's Equation, which predicts microevolution as the product of phenotypic selection and heritability, predicted evolution of larger size. This was at odds, however, with the observed lack of either phenotypic or genetic temporal trends in body size, a so-called "paradox of stasis." To investigate this paradox, we estimated the additive genetic covariance between trait and fitness, which provides a prediction of evolutionary change according to Robertson's secondary theorem of selection (STS) that is unbiased by missing variables. The STS prediction was consistent with the observed stasis. Decomposition of phenotypic selection gradients into genetic and environmental components revealed a potential upward bias, implying unmeasured factors that covary with trait and fitness. These results showcase the power of pedigreed, wild population studies-which have largely been limited to birds and mammals-to study evolutionary processes on contemporary timescales.

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.

Where the lake meets the sea: strong reproductive isolation is associated with adaptive divergence between lake resident and anadromous three-spined sticklebacks.

Contact zones between divergent forms of the same species are often characterised by high levels of phenotypic diversity over small geographic distances. What processes are involved in generating such high phenotypic diversity? One possibility is that introgression and recombination between divergent forms in contact zones results in greater phenotypic and genetic polymorphism. Alternatively, strong reproductive isolation between forms may maintain distinct phenotypes, preventing homogenisation by gene flow. Contact zones between divergent freshwater-resident and anadromous stickleback (Gasterosteus aculeatus L.) forms are numerous and common throughout the species distribution, offering an opportunity to examine these contrasting hypotheses in greater detail. This study reports on an interesting new contact zone located in a tidally influenced lake catchment in western Ireland, characterised by high polymorphism for lateral plate phenotypes. Using neutral and QTL-linked microsatellite markers, we tested whether the high diversity observed in this contact zone arose as a result of introgression or reproductive isolation between divergent forms: we found strong support for the latter hypothesis. Three phenotypic and genetic clusters were identified, consistent with two divergent resident forms and a distinct anadromous completely plated population that migrates in and out of the system. Given the strong neutral differentiation detected between all three morphotypes (mean FST = 0.12), we hypothesised that divergent selection between forms maintains reproductive isolation. We found a correlation between neutral genetic and adaptive genetic differentiation that support this. While strong associations between QTL linked markers and phenotypes were also observed in this wild population, our results support the suggestion that such associations may be more complex in some Atlantic populations compared to those in the Pacific. These findings provide an important foundation for future work investigating the dynamics of gene flow and adaptive divergence in this newly discovered stickleback contact zone.

Identifying the role of environmental drivers in organic carbon export from a forested peat catchment.

Carbon export in streams draining peat catchments represents a potential loss of carbon from long-term stores to downstream aquatic systems and ultimately, through mineralisation, to the atmosphere. There is now a large body of evidence that dissolved organic carbon (DOC) export has increased significantly in recent decades at many sites, although there is still debate about the drivers of this increase. In this study, DOC export and particulate organic carbon (POC) export were quantified from a forested peatland catchment in the west of Ireland over two years at a fine temporal resolution. The principle drivers of change in stream DOC and POC concentrations were investigated using a general additive modelling (GAM) approach. The study period included drought conditions in the early summer of 2010 and clearfelling of some commercial forestry in early 2011. The results indicated that annual loads of 9.5 t DOC km(2) year(-1) and 6.2 t POC km(2) year(-1) were exported from the catchment in 2010. This combined annual load of 15.7 t C km(2) year(-1) would represent between 0.01% and 0.02% of typical estimates for peat soil carbon storage in the region. Soil temperature, river discharge and drought explained 59.7% the deviance in DOC concentrations, while soil temperature, river discharge, and rainfall were the significant drivers of variation in POC concentrations, explaining 58.3% of deviance. Although clearfelling was not a significant factor in either model, large spikes in POC export occurred in 2011 after the first forestry clearance. The results illustrate the complexity of the interactions between climate and land management in driving stream water carbon export. They also highlight the sensitivity of peatland carbon stores to changes in temperature and precipitation, which are projected to be more extreme and variable under future climate scenarios.

Molecular pedigree reconstruction and estimation of evolutionary parameters in a wild Atlantic salmon river system with incomplete sampling: a power analysis.

BACKGROUND: Pedigree reconstruction using genetic analysis provides a useful means to estimate fundamental population biology parameters relating to population demography, trait heritability and individual fitness when combined with other sources of data. However, there remain limitations to pedigree reconstruction in wild populations, particularly in systems where parent-offspring relationships cannot be directly observed, there is incomplete sampling of individuals, or molecular parentage inference relies on low quality DNA from archived material. While much can still be inferred from incomplete or sparse pedigrees, it is crucial to evaluate the quality and power of available genetic information a priori to testing specific biological hypotheses. Here, we used microsatellite markers to reconstruct a multi-generation pedigree of wild Atlantic salmon (Salmo salar L.) using archived scale samples collected with a total trapping system within a river over a 10 year period. Using a simulation-based approach, we determined the optimal microsatellite marker number for accurate parentage assignment, and evaluated the power of the resulting partial pedigree to investigate important evolutionary and quantitative genetic characteristics of salmon in the system. RESULTS: We show that at least 20 microsatellites (ave. 12 alleles/locus) are required to maximise parentage assignment and to improve the power to estimate reproductive success and heritability in this study system. We also show that 1.5 fold differences can be detected between groups simulated to have differing reproductive success, and that it is possible to detect moderate heritability values for continuous traits (h2 ~ 0.40) with more than 80% power when using 28 moderately to highly polymorphic markers. CONCLUSION: The methodologies and work flow described provide a robust approach for evaluating archived samples for pedigree-based research, even where only a proportion of the total population is sampled. The results demonstrate the feasibility of pedigree-based studies to address challenging ecological and evolutionary questions in free-living populations, where genealogies can be traced only using molecular tools, and that significant increases in pedigree assignment power can be achieved by using higher numbers of markers.

Basin-scale phenology and effects of climate variability on global timing of initial seaward migration of Atlantic salmon (Salmo salar).

Migrations between different habitats are key events in the lives of many organisms. Such movements involve annually recurring travel over long distances usually triggered by seasonal changes in the environment. Often, the migration is associated with travel to or from reproduction areas to regions of growth. Young anadromous Atlantic salmon (Salmo salar) emigrate from freshwater nursery areas during spring and early summer to feed and grow in the North Atlantic Ocean. The transition from the freshwater ('parr') stage to the migratory stage where they descend streams and enter salt water ('smolt') is characterized by morphological, physiological and behavioural changes where the timing of this parr-smolt transition is cued by photoperiod and water temperature. Environmental conditions in the freshwater habitat control the downstream migration and contribute to within- and among-river variation in migratory timing. Moreover, the timing of the freshwater emigration has likely evolved to meet environmental conditions in the ocean as these affect growth and survival of the post-smolts. Using generalized additive mixed-effects modelling, we analysed spatio-temporal variations in the dates of downstream smolt migration in 67 rivers throughout the North Atlantic during the last five decades and found that migrations were earlier in populations in the east than the west. After accounting for this spatial effect, the initiation of the downstream migration among rivers was positively associated with freshwater temperatures, up to about 10 °C and levelling off at higher values, and with sea-surface temperatures. Earlier migration occurred when river discharge levels were low but increasing. On average, the initiation of the smolt seaward migration has occurred 2.5 days earlier per decade throughout the basin of the North Atlantic. This shift in phenology matches changes in air, river, and ocean temperatures, suggesting that Atlantic salmon emigration is responding to the current global climate changes.

The European eel quality database: towards a pan-European monitoring of eel quality.

The stocks of the European eel Anguilla anguilla are in decline and there is an increasing awareness that poor health status due to contaminants and/or diseases might be a key element in this decline and might be a hindrance to recovery. Many countries have started compiling data on the health status of eels in their water bodies. Objectives for these monitoring actions are diverse and there is a large amount of information collected by EU member countries. However, this information is widely scattered over Europe in agencies, institutes or universities. As there is a growing need to collect and report on data on the health status of the eel on international level, the Joint EIFAC/ICES Working Group on Eels initiated in September 2007 the set up of an European Eel Quality Database to collect recent data of contaminants and diseases over the distribution area of the eel. This paper describes the aim, the set up and future development of the database in order to give it greater publicity and to call on scientists or managers to submit data on eel health status. The database represents now the first comprehensive pan-European compilation of eel health data, including data from over 10,000 eels from approximately 1,200 sites over 14 countries. Preliminary work has indicated a number of shortcomings and future developments will be needed. Guaranteeing further development of the database, harmonisation of methods, quality assurance, and setting up harmonised eel monitoring strategies over Europe will be a great challenge and will need pan-European cooperative work.

The occurrence of persistent chlorinated and brominated organic contaminants in the European eel (Anguilla anguilla) in Irish waters.

The European eel (Anguilla anguilla) is a relatively high lipid, long lived species capable of living in a variety of brackish, fresh and marine habitats. As such, eels can accumulate organic pollutants and have been incorporated into environmental monitoring programs as a suitable "bioindicator" species for the determination of the levels of organic contaminants within different water bodies. The global eel stock is now in decline and while the cause of the collapse remains unidentified, it is likely to include a combination of anthropogenic mortality in addition to environmental degradation. This study provides valuable data on a range of contaminants (PCDD/Fs, PCBs, OCPs, PBDEs, HBCD, TBBPA and PBBs) and extractable lipid levels in eel muscle tissue collected from five Irish catchments. Extractable lipid levels were lower in the yellow eels compared to those in the silver eels. These levels were similar to those reported elsewhere and it has been posited that a decline in the lipid content in yellow eels may have consequences for the future viability of the stock. With the exception of higher substituted dioxins (especially OCDD), in three samples collected from one catchment (Burrishoole) in the West of Ireland, POP levels in general were determined to be low in eels from Irish waters compared to those in other countries.