A survey of the distribution of the PKD‐parasite Tetracapsuloides bryosalmonae (Cnidaria: Myxozoa: Malacosporea) in salmonids in Norwegian rivers – additional information gleaned from formerly collected fish

The malacosporean Tetracapsuloides bryosalmonae was detected in kidneys from Atlantic salmon parr in 64 of 91 sampled Norwegian rivers. Using real‐time PCR, this parasite was found to be present in Atlantic salmon parr in rivers along the whole coast, from the northernmost and southernmost areas of the country. In addition, T. bryosalmonae was found in kidneys from brown trout parr in 17 of 19 sampled rivers in south‐east Norway, and in Arctic charr sampled in the River Risfjordelva, located at the northernmost edge of the European mainland. In conclusion, T. bryosalmonae has a widespread distribution in salmonids in Norwegian watercourses. Proliferative kidney disease (PKD) caused by T. bryosalmonae and PKD‐induced mortality has been observed in salmonids in several Norwegian rivers and it can be speculated that more PKD outbreaks will occur as a result of climate change.     

A genetic tool for evaluating male‐mediated stock introgression in Atlantic salmon

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Inter-stage survival of wild juvenile Atlantic salmon, Salmo salar L.

A biological model was developed to calculate annual survival between life stages of juvenile Atlantic salmon, Salmo salar L., in Catamaran Brook, a small stream basin (52 km²) in the Miramichi River catchment in New Brunswick, Canada. Seven years’ data (1990–1996) were used in the model. Input variables included: daily fish counts and measurements of parr (3–4 age classes), smolts, and adult salmon at a fish-counting fence near the stream mouth; biennial quantification of all habitat types along the watercourse; fish density estimated by electric fishing at 30 sites; and estimates of young-of-the-year emigration via stream drift. Continuous recording of stream discharge provided data to assist in interpretation of survival estimates. Annual survival for juvenile salmon in their first 3 years of life in the stream averaged between 31% and 34%. The greatest annual variation (CV = 0.699) occurred at the egg to 0+ (summer) stage with a low of 9.2% survival recorded for a winter with an atypical midwinter flood event; parr and pre-smolt survival were similarly affected. Survival from egg deposition (after correction for losses caused by predation and retention/non-fertilization) to smolt emigration was between 0.16% and 0.52%, which is low relative to estimates from many other studies. Survival of smolts to returning 1-sea-winter adults (grilse) averaged 8.5%. Potential errors in the computation of the model are discussed, e.g. inaccurate counts of spawning adults during high autumn stream flow. A possible explanation for the low egg to smolt survival was the environmental conditions experienced during various winters. Mean egg survival was 1.3 times higher (39.3%) and egg to smolt survival increased to 1.03% when the two winters characterized by extremely low discharge or midwinter freshets were excluded from the calculation. Density-dependent factors related to a beaver dam, which limited spawning distribution, may also have contributed to poor survival and increased fry emigration in one year. Environmental factors, particularly winter conditions, in streams such as Catamaran Brook may act as bottlenecks to natural production of Atlantic salmon.     

Reproductive strategies in small populations: using Atlantic salmon as a case study

Abstract –  Wild salmonid populations with only a few breeding adults may not exhibit a significant reduction in genetic variability compared with larger populations. Such an observation suggests that effective population sizes are larger than population size estimates based on direct adult counts and/or the mating strategy maximises outbreeding, contributing to increased heterozygosity. In the case of wild Atlantic salmon Salmo salar populations, stratification by age classes and sexes on the spawning grounds avoids inbreeding and increases genetic variability. We studied the breeding composition of four Spanish salmon populations. Over a 7-year period we concluded that the probability of within-cohort mating is very low: females generally reproduce after two sea-winters whereas males reproduce mostly as one sea-winter ( grilse ) and/or mature parr. Considering different levels of contribution of mature parr to spawning derived from field surveys, we developed a simple model for estimating effective population sizes and found that they doubled with 65% parr contribution expected for rivers at this latitude (43°N), and ranged from 100–800 individuals. The effect of between-cohort mating was modelled considering different ranges of differences in allele frequencies between cohorts and resulted in 28–50% increases in heterozygosity when considering a 65% parr contribution. The complex mating strategy of Atlantic salmon contributes to explain the high levels of genetic variability found for small populations of this species. This model can probably be extended to other animal species with mating strategies involving different cohorts.     

Reproductive success of Atlantic salmon (Salmo salar) mature male parr in a small river,the Nivelle: influence of shelters

Grimardias D, Merchermek N, Manicki A, Garnier J, Gaudin P, Jarry M, Beall E. Reproductive success of Atlantic salmon (Salmo salar) mature male parr in a small river, the Nivelle: influence of shelters.
Ecology of Freshwater Fish 2010: 19: 510–519. © 2010 John Wiley & Sons A/S Abstract – The breeding activity of a small population of Atlantic salmon (Salmo salar L.) was monitored on the Nivelle River in southwest France to estimate the overall contribution of mature male parr and the numbers involved in egg fertilisation. In the twelve redds sampled, 563 eggs were collected for parental assignment and the physical habitat was characterised to investigate the possible relationship between parr reproductive success and habitat complexity. The overall contribution of mature male parr was particularly high (87%, one of the highest estimates ever reported). Concerning habitat complexity, granulometry and shelter diversity were correlated with the number of parr breeders contributing to egg fertilisation in each redd. A complex habitat increased multiple paternity, which could affect the effective size and genetic variability of small salmon populations.     

Transmission dynamics of the monogenean Gyrodactylus salaris under seminatural conditions

Tracking individual variation in the dynamics of parasite infections in wild populations is often complicated by lack of knowledge of the epidemiological history of hosts. Whereas the dynamics and development of Gyrodactylus salaris Malmberg, 1957, on Atlantic salmon, Salmo salar L., are known from laboratory studies, knowledge about infection development on individual wild fishes is currently sparse. In this study, the dynamics of an infection of G. salaris on individually marked Atlantic salmon parr was followed in a section of a natural stream. During the 6‐week experiment, the prevalence increased from 3.3 to 60.0%, with an average increase in intensity of 4.1% day^?1. Survival analyses showed an initially high probability (93.6%) of staying uninfected by G. salaris, decreasing significantly to 37% after 6 weeks. The results showed that even at subarctic water temperatures and with an initially low risk of infection, the parasite spread rapidly in the Atlantic salmon population, with the capacity to reach 100% prevalence within a short summer season. The study thus track individual infection trajectories of Atlantic salmon living under near‐natural conditions, providing an integration of key population parameters from controlled experiments with the dynamics of the epizootic observed in free‐living living populations.     

Influence of long term ammonia exposure on Atlantic salmon (Salmo salar L.) parr growth and welfare

The objective of this study was to determine the long‐term effects of ambient unionized ammonia nitrogen (NH₃‐N) combined with different feeding regimes on Atlantic salmon Salmo salar L parr growth, welfare and smoltification. Previous studies on the parr stage of Atlantic salmon have mostly focused on acute exposure, or at low temperatures. Atlantic salmon parr were exposed for 105 days (at 12°C, pH 6.8) to four sublethal ammonia concentrations ranging from 0.1 to 35 μg L^?1 NH₃‐N (0.1–25 mg L^?1 TAN) at two feeding levels: full feed strength (+20% overfeeding) and 1/3 of full feed strength. After 21 days, it was observed that 32 μg L^?1 NH₃‐N reduced growth rate of parr fed full ration, but this effect was not evident at the end of the exposure. Feed utilization was not affected by ammonia exposure at any sampling point. Increasing ammonia levels were associated with a higher prevalence and severity of gill damage at 22 days but not at the end of the exposure. The examination of welfare indicators revealed only a few pathologies, not related to ammonia exposure. In addition, higher ammonia concentrations did not appear to influence the development of hypo‐osmoregulatory ability during parr‐smolt transformation.     

Regional variability in Atlantic salmon (Salmo salar) riverscapes: a simple landscape ecology model explaining the large variability in size of salmon runs across Gaspé watersheds,Canada

Kim M, Lapointe M. Regional variability in Atlantic salmon (Salmo salar) riverscapes: a simple landscape ecology model explaining the large variability in size of salmon runs across Gaspé watersheds, Canada.
Ecology of Freshwater Fish 2011: 20: 144–156. © 2010 John Wiley & Sons A/S Abstract – Atlantic salmon (Salmo salar) rivers in the Gaspé Peninsula, Quebec, present a 20 to 1 variability in the average numbers of returning adult salmon per km² of watershed area (the ‘specific run size’). These variations are very poorly explained by interbasin differences in total stream length (R² = 0.033, P = 0.533) or in estimates of total area of salmon habitat that fail to take into account complementarity and interconnections across life stage habitats (R² = 0.065, P = 0.448). The relative spatial distribution of three complementary habitat types (adult holding pools, spawning beds, parr habitats) is hypothesised to be an important factor controlling the production of salmon at any given watershed area or given total stream length, and by extension the numbers of adults returning each year to spawn. We developed a simple riverine landscape ecology (or riverscape) model that uses easily accessible topographic map sources to identify optimally productive river segments. These segments were identified based on large‐scale river and valley features and the associated spatial organisation of complementary salmon habitats. We tested the ability of this model to predict salmon run sizes for 14 watersheds in the Gaspé Peninsula. The aggregate length of optimally productive segments, as defined in our model, is a strong predictor of the average size of the annual salmon runs for these watersheds (R² = 0.913, P < 0.0005). Furthermore, specific salmon run sizes were accurately predicted (R² = 0.771, P < 0.0005) after removal of the obvious scale effects of watershed size.     

Assessing the diet of North American Atlantic salmon (Salmo salar L.) off the West Greenland coast using gut content and stable isotope analyses

Investigations on the marine feeding of Atlantic salmon (Salmo salar L.) in the Northwest Atlantic are limited compared with the Northeast Atlantic. Climate‐induced changes to food webs in Atlantic salmon feeding areas have been noted, alongside increased mortality despite a cessation of most marine fisheries. As forage efficiency may be hampering survival, it was important to address this knowledge gap. Atlantic salmon were sampled at three sites on the West Greenland coast (Sisimiut, Nuuk and Qaqortoq) between 2009 and 2011. Gut content and stable isotope analyses were combined to assess spatial and temporal differences in feeding. Capelin (Mallotus villosus) dominated the diet at Nuuk and Qaqortoq, whereas boreoatlantic armhook squid (Gonatus fabricii) was the dominant prey at Sisimiut. Hyperiid amphipods (Themisto spp.) and sand lance (Ammodytes spp.) were also important. Significant differences were found among sites for both gut contents and stable isotope analyses, with fewer differences evident temporally. Dietary differences were also evident across larger scales, with little overlap demonstrated with Northeast Atlantic diets and the emergence of boreoatlantic armhook squid as an important prey item over time. Atlantic salmon diets are frequently anchored on one or two prey items, on which they appear to specialize, but they will diversify to consume other available pelagic prey. Thus, Atlantic salmon are an opportunistic, generalist predator within the pelagic food web. The variability evident in diet suggests that the limited data available are insufficient to appropriately understand potential vulnerabilities that the species may have to ecosystem changes, and suggest further research is needed.     

Interactions between naturalised exotic salmonids and reintroduced Atlantic salmon in a Lake Ontario tributary

Abstract – Atlantic salmon (Salmo salar) was once native to Lake Ontario, however, its numbers rapidly declined following colonisation by Europeans and the species was extirpated by 1896. Government agencies surrounding Lake Ontario are currently undertaking a variety of studies to assess the feasibility of reintroducing Atlantic salmon. We released hatchery‐reared adult Atlantic salmon into a Lake Ontario tributary to examine spawning interactions between this species and fall‐spawning exotic salmonids found in the same stream. Chinook salmon, coho salmon and brown trout were observed interacting with spawning Atlantic salmon in nearly one‐quarter of our observation bouts, with chinook salmon interacting most frequently. Whereas a previous investigation found that chinook salmon caused elevated agonistic behaviour and general activity by spawning Atlantic salmon, the present study found that interspecific courtship was the most common form of exotic interaction with spawning Atlantic salmon. In particular, we observed precocial male Chinook salmon courting female Atlantic salmon and defending the female against approach by male Atlantic salmon. We discuss the potential implications of these interactions on the Lake Ontario Atlantic salmon reintroduction programme.     

Do habitat measurements in the vicinity of Atlantic salmon (Salmo salar) parr matter?

Atlantic salmon, Salmo salar L., parr habitat characterisation is usually performed by in situ measures of key environmental variables taken at the exact fish location if the fishing gear allows precise pinpointing of this location, or in large sampling sections covering a river reach or mesohabitat, often ignoring variability in the immediate vicinity around individual fish. These data may be critically important in the development and validation of habitat preference models. The influences of seven increasing distances of measures, the variation of the number of considered measures and the depth of velocity measurement (bottom or 0.6 of the depth) in the calculations of HSI (Habitat Suitability Index) from a multiple‐experts fuzzy model of Atlantic salmon parr habitat were tested. When a parr was present, six measures collected in a 50‐cm radius around the fish to provide an average measure as input data and velocity measured at 60% of the depth gave the highest HSI values. These results show some potential for the use of an intermediate study scale, between micro‐ and mesohabitat, and questions how fish habitat conditions are currently measured.     

Low concentrations of sodium hypochlorite affect population dynamics in Gyrodactylus salaris (Malmberg, 1957): practical guidelines for the treatment of the Atlantic salmon,Salmo salar L. parasite

Atlantic salmon, Salmo salar L. parr (age 1+), infected by the monogenean ectoparasite Gyrodactylus salaris (Malmberg, 1957), were exposed to chlorine (Cl)‐enriched water at three different concentrations: Cl_low (0–5 μg Cl L^?1), Cl_medium (18 μg Cl L^?1) and Cl_high (50 μg Cl L^?1). There was a negative correlation between G. salaris infections and the hypochlorite concentrations added. The parasite infection was eliminated by day 6–8 and day 2–4 in the groups Cl_medium and Cl_high, respectively, while inhibition of G. salaris population growth was observed in the Cl_low group. An important note to this matter, however, is that the G. salaris specimens observed at day 6 in Cl_medium and at day 2 in Cl_high were all considered dead by subjective judgement. No mortality in the salmon parr was observed during the first 8 days of the experiment, demonstrating that Cl has a stronger effect on G. salaris than on the salmonid host. The differences in sensitivity between the parasite and the Atlantic salmon indicate that hypochlorite has a potential use as a parasiticide with a therapeutic margin. The low‐dose sensitivity may imply that Cl pollution in urban areas may pose a greater risk towards biodiversity than previously assumed.     

An additional life-history tactic of masu salmon: Migration of parr to coastal habitats

Alternative life-history tactics of masu salmon Oncorhynchus masou are well documented. Subsequent to the freshwater parr stage (age ≥ 1 + years), an anadromous form migrates to the sea after smolting, while a resident form matures without seaward migration. In addition to this typical migratory dimorphism, anecdotal reports based on field observations have indicated that some underyearling masu salmon use estuarine waters. However, no empirical evidence indicates saltwater utilisation and subsequent survival in the early parr stage. Here, we used otolith microchemistry to examine whether a portion of masu salmon parr in northern Japan enters coastal habitats. The otolith Sr:Ca ratios of most juveniles collected from six rivers had consistently low values, indicating that masu salmon parr inhabiting these rivers stay only in freshwater. In contrast, in individuals from a steep-gradient river the Sr:Ca ratios increased at about a 200-µm distance from the otolith core. These results suggest that some masu salmon parr might use brackish water or sea water temporarily. In addition, three masu salmon parr were found in another steep river where a culvert located only ten metres from the river mouth completely blocked upstream migration for spawning. The Sr:Ca ratios in these fish increased at about >200 µm from the otolith core, indicating the parr had immigrated to the non-natal river from the sea. Such flexible behaviour at an early life stage may contribute to the spatial expansion of masu salmon, and the movement could moreover help to stabilise its population dynamics.     

Metabolic adaptations of oxidative muscle during spawning migration in the Atlantic salmon <Emphasis Type="Italic">Salmo salar</Emphasis> L.

The adaptability/plasticity of the highly oxidative red muscle in Atlantic salmon was demonstrated during spawning migration. Substrate concentrations and the enzymatic pathways of ATP production were examined in red muscle obtained from Atlantic salmon at different sites along their migratory route in the Exploits River, Newfoundland, Canada. Individuals were chronologically sampled from a seawater site, two sites upstream, and at spawning. The 20% decrease in salmon body weight during the later stages of migration was accompanied by large decreases (mg dry weight⁻¹) in both glycogen (P < 0.01) and total muscle lipid (P < 0.01). In contrast, water content and protein concentration (mg dry weight⁻¹) of the red muscle increased by 25 and 34%, respectively, at spawning. Enzymes of the glycolytic pathways demonstrated a significant (P < 0.001) decrease in maximal activity as migration proceeded whereas enzymes of the oxidative phosphorylation pathways, specifically the citric acid cycle enzymes, exhibited an increase (P < 0.001) in maximal activity at spawning. The antioxidant enzyme superoxide dismutase also demonstrated an increase (P < 0.001) in maximal activity during the latter stages of migration. These adaptations imply that the red epaxial muscle of Atlantic salmon has a more efficient means of oxidizing lipids, while minimizing free radical damage, during the later stages of migration and spawning, thereby potentially increasing post spawning survival.     

Wild origin and enriched environment promote foraging rate and learning to forage on natural prey of captive reared Atlantic salmon parr

Abstract – After release to the wild, captive reared salmon have shown lower foraging rates on natural prey and impaired ability to avoid natural predators and thus lower survival compared with wild‐born conspecifics. Here, we examine whether captive breeding influences learning of foraging on natural prey and how enriched rearing methods may improve foraging on natural prey by Atlantic salmon (Salmo salar) parr. We reared offspring of hatchery or wild salmon of the same population in either a standard or enriched environment at production‐scale densities. The enriched environment featured submerged overhead shelter, varying water current, depth and direction and consequently alterations in food dispersion. Parr reared in the enriched environment expressed higher feeding rates, and parr of wild origin started to forage earlier on natural prey. The enriched method promoted foraging of hatchery reared parr on natural prey and is easily applicable to commercial production of salmonids for reintroduction or stock enhancement purposes.     

Migratory behaviour of adult wild and escaped farmed Atlantic salmon, Salmo salar L., before, during and after spawning in a Norwegian river

The migratory behaviour of adult wild and escaped farmed Atlantic salmon, Salmo salar L., before, during after spawning in the River Namsen, Norway, was analysed using radio telemetry. The fish were caught, radio tagged and released into the fjord between 7 and 25 km from the river mouth. A significantly higher proportion of wild (74%) than farmed (43%) salmon was subsequently recorded in the river. Wild salmon (33%) were more frequently captured in the sea and in rivers than farmed salmon (14%). The migration speed from release to passing a data logger 11 km upstream from the river mouth was not significantly different between wild (20.6 km day^?1) and farmed (19.8 km day^?1) salmon. Wild salmon tagged when water flow in the river was increasing had a significantly higher migration speed than wild salmon tagged when water flow was decreasing. This was not true for farmed salmon. Farmed salmon were distributed significantly higher up the river than wild salmon during spawning, although both types of fish were found together in spawning areas. Thus, there was no geographical isolation to prevent spawning between wild and escaped farmed salmon. Farmed salmon had significantly more and longer up- and downstream movements than wild salmon during the spawning period. Unlike farmed salmon, the number of riverine movements by wild salmon increased significantly when variation in water flow increased. A smaller proportion of wild (9%) than farmed (77%) salmon survived through the winter after spawning.     

Some behavioural and ecological factors affecting distribution,biomass and production of juvenile Atlantic salmon

Observations were made in an experimental stream tank (total area 14.7 m²) on juvenile Atlantic salmon, Salmo salar L (parr), relating experimental observations to field observations, including the reported diurnal fasting behaviour of juvenile salmon at water temperatures <10 °C. Densities in the tank ranged from five to twenty parr, at water temperatures ranging from 4.6 °C to 15.8 °C. The wide channel of the stream tank, with mean water velocity of 18.8 cm·s^?1, was the preferred section, where territorial behaviour was observed. Biomass was regulated in the wide channel by territorial mosaics or by dominance hierarchies. Dominance hierarchies were reflected in coloration of the fish. Dominant salmon were generally in the wide channel. Densities of salmon parr (of mean fork length 10.2 cm) in the channel ranged from 0.84 m^?2 to 1.73 m^?2, with an average biomass of 14.2 g·m^?2. Growth was least at the 5.9 °C temperature treatment. In experiments at temperatures below 10 °C, feeding, dominance hierarchy and territorial behaviour were observed in daylight hours, contrary to the published literature. Interactions with other species may affect behaviour. Some observations were made on a closely related species, brown trout (Salmo trutta L.), a commonly cohabiting species in many systems. Trout displaced salmon from their preferred locations in the tank and were more aggressive than the salmon, reducing agonistic behaviour by the salmon. The commonest agonistic act shown by salmon was ‘charge’ and that by the trout was ‘approach’. Some field observations affecting behaviour and production are discussed.     

Allozyme heterozygosity and development in Atlantic salmon, Salmo salar

The development of Atlantic salmon (Salmo salar) parr was monitored during the first year in a fresh-water hatchery environment to test whether multilocus heterozygosity (computed for six allozyme loci: _s AAT-4^*, IDDH-1^*, IDDH-2^*, IDPH-3^*, _s MDH-3,4^* and_m MEP-2^*) is associated with growth (body mass, fork length) and maturity status (mature male parr). The length frequency distribution of population was unimodal in September, but was clearly bimodal by December. UMG fish tended to be more heterozygous and had higher growth rates than LMG fish. The heterozygosity of UMG fish and mature male parr (MMP) did not differ significantly but MMP showed lower growth rates than UMG. There were positive correlations between the number of heterozygotic loci and the fork length, weight or growth rates of individuals.     

Lake-use by juvenile Atlantic salmon (Salmo salar L.) and other salmonids in northern Norway

Abstract– The utilization of lakes, and inlet and outlet streams by juvenile Atlantic salmon ( Salmo salar L.), brown trout ( Salmo trutta L.) and Arctic charr ( Salvelinus alpinus (L.), were investigated in 16 watercourses northern Norway, all known to inhabit salmon stocks. In lakes, fish were caught by small mesh size gill nets, while in rivers fish were caught electrofishing. In the shallow littoral (0-3 m depth) there were juvenile salmon in 15 of 19 investigated lakes, juvenile trout in 17 and juvenile charr in seven. Trout dominated significantly in numbers in the shallow littoral of seven lakes, while salmon and charr dominated in three lakes each. When trout and salmon were frequent in the shallow littoral, charr was usually not present in this habitat, but were found in the profundal zone in most of the lakes. Atlantic salmon parr utilized both shallow and deep lakes, and used both stones and macrophytic vegetation as shelter. The utilization of lakes by salmon parr seemed to be closely related to utilization of small inlet streams for spawning. In most inlet and outlet streams salmon dominated over trout in numbers, while charr were absent. This is the first documentation of lake-use by naturally occurring salmon parr in Scandinavia.     

Prespawning migratory behaviour of wild and farmed Atlantic salmon, Salmo salar L, in a north Norwegian river

The migration patterns of wild and released farmed Atlantic salmon, Salmo salar L., were studied by radio-telemetry during migration from entering a river to after spawning. The wild salmon were caught during return migration in bagnets and the farmed salmon were produced in a near by fish farm. Both groups were tagged and released at sea near the river mouth of the River Alta, North Norway. A pronounced individual variation in migration pattern was observed. No significant differences was found between wild and farmed salmon in the distance from entering the river mouth to the place of the first stop (recorded in the same pool for 5 days or longer) and the days from entering the river mouth to arrival at the first stop. For both wild and farmed salmon, mean migration speed was 2.6 km day¹, varying from 0.4 to 11.0 km day^-1. A larger proportion of farmed salmon distributed to the upper part of the river at spawning; mean distances from the river mouth were 30.1 and 19.1 km. respectively. Farmed salmon spent a significantly longer time from entering the river to reaching the area occupied during spawning.