Is there a threshold size regulating seaward migration of brown trout and Atlantic salmon?

We investigated the influence of variation in body size and growth rate on age of smolting in Atlantic salmon and brown trout in four different Norwegian rivers. In Atlantic salmon smolt ages varied between 2 and 6 years, and in brown trout between 2 and 7 years. Smolt age was negatively correlated with parr growth, and positively correlated with smolt size. Age at smolting was more variable in the two northern than the two southern rivers. Smolt sizes and ages were also more variable in brown trout than in Atlantic salmon. Based on the observed variation in smolt size and age, we reject the hypothesis that a threshold size alone regulates age at smolting. Within populations smolt age depends on growth rate so that fast-growing parr smolted younger and smaller than slow-growing parr. We hypothesize that smolt size and age is a trade-off between expected benefits and costs imposed by differences in individual growth rate.     

Latitudinal clines in life-history variables of anadromous brown trout in Europe

Based on published data, we reviewed clinal variations in life-history characteristics of anadromous brown trout Salmo trutta from 102 European rivers at latitudes between 54 and 70° N. Growth rate in fresh water, mean smolt age, mean sea age at first sexual maturity, proportion of repeal spawners among adults, longevity, and length of adult life span exhibited latitudinal clines. Brown trout grew faster in fresh water, smolted and matured younger, lived fewer years but spawned more times in the south than in the north. The life-history traits studied were often correlated. Longevity, smolt age and sea age at maturity were negatively and smolt length and proportion of repeat spawners among adults were positively correlated with growth rate in fresh water. Longevity was positively correlated with smolt age and sea age at maturity. The latter also increased with increasing smolt age. None of these significant correlations among life history variables, except for those between smolt age and parr growth and proportion of repeat spawners and parr growth, are latitudinal effects. We do not know to what extent the latitudinal variation in life–history traits reflects phenotypic plasticity and to what extent it is caused by inherited differences among populations.     

Pre‐migratory differentiation of wild brown trout into migrant and resident individuals

In February to March, wild brown trout Salmo trutta were captured by electrofishing in a natural watercourse (tributaries of the River Lille Aa, Denmark), individually tagged (Passive Integrated Transponders), and released. Representatives of the tagged brown trout were recaptured on the release sites in April by electrofishing and eventually caught in downstream smolt traps ('migrants') placed in the main river or by electrofishing ('residents') on the initial sites in June. Upon each capture, smolt appearance and body size were evaluated, and a non‐lethal gill biopsy was taken and used for Na⁺,K⁺‐ATPase analysis. Based on repetitive gill enzyme analysis in individual fish, a retrospective analysis of the rate of development in individual brown trout ultimately classified as migrants or residents was performed. Two months prior to migration, a bimodal morphological and physiological (gill Na⁺,K⁺‐ATPase) development concurred and was related to the subsequent differentiation into resident and migratory fractions of each population. This differentiation was unrelated to growth rate and body size of individual fish but skewed in favour of migratory females. Individuals destined to become migrants developed a smolt‐like appearance before the onset of migration and had higher rate of change of gill Na⁺,K⁺‐ATPase activity than fish remaining residents. The rate of change of gill Na⁺,K⁺‐ATPase activity was independent of the distance migrated to the trap (3–28 km). Thus in bimodal wild brown trout populations a major increase in enzyme activity takes place before migration is initiated and is a characteristic of migratory individuals only.     

Risk of inter-river transmission of Gyrodactylus salaris by migrating Atlantic salmon smolts, estimated by Monte Carlo simulation

The possibility of Gyrodactylus salaris infection of wild North Atlantic salmon Salmo salar spreading to new rivers poses a major threat in Norway. This freshwater parasite can survive for some time in brackish water, and it has been suggested that smolts leaving infected rivers could transport vital parasites to new rivers. A Monte Carlo simulation model was used to estimate the risk that infected smolts would ascend a new river. Data from an infected watercourse in Norway, where the salmon population is maintained constant by cultivation, were used. The model included information on prevalence of infection, hydrographical conditions, survival of G. salaris in brackish water, fish population characteristics, and smolt behaviour during seaward migration. The annual risk was estimated for 3 neighbouring rivers situated at different distances from the index river. For the nearest river, which shares the same brackish water zone with the index river, the model estimated an annual risk of 31% that at least 1 infected smolt would ascend this river. The results of the simulation were highly sensitive to the water salinity along the migration route. For the other rivers, the annual risk was lower than 0.5%. Risk was positively correlated with the number of fish leaving the index river, indicating control of this number as a possible tool in risk management.     

Reconstructing the Migratory Behavior and Long-Term Survivorship of Juvenile Chinook Salmon under Contrasting Hydrologic Regimes

The loss of genetic and life history diversity has been documented across many taxonomic groups, and is considered a leading cause of increased extinction risk. Juvenile salmon leave their natal rivers at different sizes, ages and times of the year, and it is thought that this life history variation contributes to their population sustainability, and is thus central to many recovery efforts. However, in order to preserve and restore diversity in life history traits, it is necessary to first understand how environmental factors affect their expression and success. We used otolith ⁸⁷Sr/⁸⁶Sr in adult Chinook salmon (Oncorhynchus tshawytcha) returning to the Stanislaus River in the California Central Valley (USA) to reconstruct the sizes at which they outmigrated as juveniles in a wetter (2000) and drier (2003) year. We compared rotary screw trap-derived estimates of outmigrant timing, abundance and size with those reconstructed in the adults from the same cohort. This allowed us to estimate the relative survival and contribution of migratory phenotypes (fry, parr, smolts) to the adult spawning population under different flow regimes. Juvenile abundance and outmigration behavior varied with hydroclimatic regime, while downstream survival appeared to be driven by size- and time-selective mortality. Although fry survival is generally assumed to be negligible in this system, >20% of the adult spawners from outmigration year 2000 had outmigrated as fry. In both years, all three phenotypes contributed to the spawning population, however their relative proportions differed, reflecting greater fry contributions in the wetter year (23% vs. 10%) and greater smolt contributions in the drier year (13% vs. 44%). These data demonstrate that the expression and success of migratory phenotypes vary with hydrologic regime, emphasizing the importance of maintaining diversity in a changing climate.     

Impact of habitat fragmentation on the genetics of populations in dendritic landscapes

1. The effect of habitat fragmentation on freshwater species has been addressed using brown trout Salmo trutta L. as a model species with a dendritic population structure. 2. Microsatellite loci were employed as molecular markers. Levels of gene flow and population subdivision were determined in more than 1200 brown trout individuals inhabiting four south European rivers with contrasting patterns of fragmentation, defined by the presence of barriers. 3. The genetic units in the four rivers were restricted by artificial barriers, and gene flow among samples within each river was associated with the level of fragmentation of the river. 4. Loss of genetic diversity and dislocation of the dendritic model have been detected in fragmented rivers. These results emphasise the importance of mitigating the impact of dams by constructing passages to restore gene flow along the river, for fish and other migratory species, as well as the need for caution in relation to stocking in isolated areas to avoid problems of inbreeding.     

Migratory timing, marine survival and growth of anadromous brown trout Salmo trutta in the River Imsa, Norway

The aim of the paper was to study sea migration, growth and survival of brown trout Salmo trutta of the River Imsa, 1976–2005. The migratory S. trutta were individually tagged and fish leaving or entering the river were monitored daily in traps located near the river mouth. The mean annual duration of the sea sojourn was 6–9 months for first-time migrants moving to sea between January and June. It was 8–18 months for those migrating to sea between July and December. Veteran migrants stayed 12 months or less at sea and most returned to the river in August. Early ascending fish stayed the longest in fresh water because most returned to sea in April to May. The day number of 50% cumulative smolt descent correlated negatively with mean water temperature in February to March and the February North Atlantic Oscillation index (NAOI). Mean annual sea growth during the first 2 years after smolting was higher for S. trutta spending the winter at sea than those wintering in the River Imsa. First year's sea growth was lower for S. trutta descending in spring than autumn. For first-time migrants, it correlated negatively with the February NAOI of the smolt year. Sea survival was higher for spring than autumn descending first-time migratory S. trutta with a maximum in May (14·9%). Number of anadromous S. trutta returning to the river increased linearly with the size of the cohort moving to sea, with no evidence of density-dependent sea mortality. Sea survival of S. trutta smolts moving to sea between January and June correlated positively both with the annual number of Atlantic Salmo salar smolts, the specific growth rate at sea, and time of seaward migration in spring. This is the first study indicating how environmental factors at the time of seaward migration influence the sea survival of S. trutta .     

Sea trout in North Argyll Sea lochs, population, distribution and movements

Sea trout were sampled by shore seining in the sea lochs of the west coast of Scotland between 1970 and 1974, This area is of special interest because of the indentation of the coastline and the varying degree of freshwater and marine influence in the sea lochs. The majority of fish caught were in their first year after smolt migration. These post-smolts were caught mainly in May and June, after which the number of trout present was low until fish at the 'whirling' stage appeared in the catches at the end of August, continuing through to the following spring. An additional recruitment of mainly unsilvered young trout from the rivers to the sea lochs was found in the autumn. Age and sex composition of both spring and autumn recruits, and of mature fish, were investigated and compared.
A total of 3228 sea trout were tagged, with 311 recaptures. These recaptures supported the evidence from smolt trapping and beach seine catches that, in the first post-migration year, the smolts migrate from the rivers from late March to early May, they then migrate from the sea lochs in May and June and return in late summer and autumn.     

Survival and behavior of juvenile steelhead trout (Oncorhynchus mykiss) in two estuaries in Oregon, USA

Anadromous salmonids are viewed as a prized commodity and cultural symbol throughout the Pacific coast of North America. Unfortunately, several native salmonid populations are threatened or at risk of extinction. Despite this, little is known about the behavior and survival of these fish as the juveniles transition from freshwater to the ocean. Our primary objectives were to estimate survival of juvenile steelhead migrating between trapping sites and the ocean and evaluate whether survival in the estuary varies temporally (within a year) or spatially (within and between estuaries) within the same distinct population segment. We also evaluated whether flow or fork length were correlated with survival and collected information on variables that have been demonstrated to affect smolt survival in other studies to lend insight regarding differences in survival estimates between basins. We compared run timing, migration rate, survival, condition factor, age composition and time of residence in the estuary for steelhead outmigrants from each basin and measured parasite loads in outmigrating steelhead to evaluate potential differences in parasite density and parasite community between basins. In 2009, we implanted acoustic transmitters in 139 wild steelhead smolts in two small rivers on the Oregon Coast. In general, only 40–50 % of the wild steelhead smolts tagged at upstream smolt traps were detected entering the ocean. The majority of mortality occurred in the lower estuary near the ocean. Wild steelhead smolts typically spent less than 1 day in the estuary in both basins. Using similar data from previous studies in the Nehalem and Alsea basins, we showed that survival appears to be negatively correlated with flow in most releases, and in 2009 fork length was not correlated with survival. Our observations provide baseline information on factors that could influence smolt survival through the estuary as well as smolt to adult survival in these basins, and emphasize the importance of monitoring smolt survival in the estuary.     

Morphological and physiological changes of female tropical eels, Anguilla celebesensis and Anguilla marmorata, in relation to downstream migration

Acoustic telemetry and microchemical analysis of otolith strontium-calcium ratios were used to evaluate how exotic brown trout Salmo trutta have responded to Japanese riverine environments of south-western Hokkaido by observing their migratory patterns. The existence of anadromous S. trutta was also verified. Most S. trutta caught in rivers for otolith analysis were freshwater residents (95·6%), whereas those caught in the sea were mainly smolts (91·3%), which had just migrated from rivers during spring. Anadromous S. trutta (n = 6) were captured in rivers and in the sea, confirming the existence of mature pre- and post-spawning fish. According to telemetry results, both mature and immature S. trutta used the river in winter, and their estimated sea-run timings showed individual differences. Through the combination of these two methods, migratory patterns on various spatio-temporal scales were observed. This first documentation of the presence of both male and female anadromous S. trutta in the same region within Japan indicated the risk of further colonization of exotic S. trutta via oceanic migration.     

Variation in some biological characteristics of British sea trout, Salmo trutta L.

Almost 50 assessments of British sea trout are available in the literature and the objective in this work is to examine variation in the parameters by which the fish are usually described and to discover biological criteria on which stocks may be classified. Eight statistics are common to the majority of stock evaluations: mean smolt age, proportion of finnock to spawn, mean age of a stock at first maturation, size achieved by the smolt at migration, rate of growth at sea, survival at sea, diversity of age categories in a stock and condition factor of sea-run fish. When the influence of each factor on the others is tested two characteristics emerge as being of key importance in the biology of adult sea-trout; the life expectancy and the weight: percentage previous spawners in a sample. The distribution of stocks is discussed in this context and two main groupings of British sea trout, corresponding geographically with the Irish sea and the Atlantic sea-board, are proposed.     

Effects of an Introduced Piscivore on Native Trout: Insights from a Demographic Model

Introductions of exotic species pose a significant threat to the persistence of many native populations, including many inland fishes. In 1994, piscivorous lake trout (Salvelinus namaycush) were discovered in Yellowstone Lake, Yellowstone National Park, Wyoming, USA, one of the last strongholds of the native Yellowstone cutthroat trout (Oncorhynchus clarki bouvieri). Predation by lake trout is expected to lead to a substantial decline in the native cutthroat trout population, which may have significant negative consequences for terrestrial predators that depend on cutthroat trout for prey and for the recreational fishery of the Park. We developed a matrix demographic model for the cutthroat trout population in Yellowstone Lake to identify the life stages that are most critical for understanding population dynamics. Parameter estimates (vital rates) were manipulated to explore the possible consequences of lake trout invasion. Comparisons of our results with current estimates of population trend and age structure suggested that our model reflected current conditions of the system. Elasticity analysis of the model revealed that population growth was most sensitive to annual survival of young trout, the group that is expected to be most vulnerable to lake trout predation. Projection of our deterministic model suggested that, in addition to a decline in abundance of cutthroat trout, the effects of lake trout may be manifest as changes in age and breeding structure of the population. Simulations of a stochastic version of the model indicated that a 60% or greater decline in the cutthroat trout population could be expected within 100 years if the lake trout population were permitted to grow uncontrolled. However, an effective control strategy that prevented the establishment of a large population of lake trout substantially reduced population decline, although the reduction in the availability of adult trout to terrestrial predators and anglers may be still be substantial (20–40%). In addition to current control activities in place in the Park, we recommend a renewed emphasis on understanding and monitoring juvenile life stages of cutthroat trout. Our results demonstrate the value of existing data sets for developing models to estimate the potential impact of biological invasions on the management and conservation of native populations, especially when opportunities and resources for additional empirical studies are limited.     

Habitat use and dispersal of post-smolt sea trout Salmo trutta in a Scottish sea loch system

Post-smolt anadromous brown trout Salmo trutta , sea trout, from two Scottish west coast rivers, the Balgy and Shieldaig, flowing into adjacent sea lochs were tracked simultaneously using arrays of moored acoustic receivers to determine dispersal patterns and loss rates. Fish tended to stay close to their natal rivers for the first 14 day after entering the sea, during which time about half the fish were lost to the study. Although initially the overall pattern of dispersal was similar for individual fish from both rivers, towards the end of the study the groups had converged into one of the loch basins. There were also pronounced individual differences in habitat use with all those fish detected for >42 days exhibiting different patterns of habitat use. Loss rates were similar between the two rivers despite differences in the range of air-breathing predators to which the fish were initially exposed. These findings suggest that any management of predators or other mortality agents should be targeted towards mouths of rivers during and immediately following smolt emigration.     

Comparing competitive ability and associated metabolic traits between a resident and migratory population of bull trout against a non-native species

Juvenile bull trout Salvelinus confluentus from two geographically and ecologically distinct populations were compared with regard to their ability to compete with non-native brook trout Salvelinus fontinalis in an artificial stream, and with respect to their rates of oxygen consumption. Bull trout collected from a migratory population foraged more successfully against brook trout competitors than those from a resident population, capturing more of a limited amount of food items presented. The migratory population was also more aggressive (measured by the number of nips, chases and lateral threat displays) against brook trout competitors than the resident population. Bull trout from the migratory population had a higher oxygen consumption rate (203 mg O₂ kg · hr^-1) in the field than similar sized fish from the resident population (183 mg O₂ kg · hr^-1). These results suggest native bull trout have population-level variation in competitive ability against a non-native species and such competitive ability is positively associated with metabolism and migratory life history.     

Migration route selection of juvenile Chinook salmon at the Delta Cross Channel, and the role of water velocity and individual movement patterns

We examined movement tracks of ultrasonic-tagged juvenile Chinook salmon (Oncorhynchus tshawyscha) smolts at the juncture of two migratory pathways. This migratory juncture occurs where the Delta Cross Channel splits from the Sacramento River in California’s Sacramento–San Joaquin Delta. Smolt tracks were analyzed to compare the importance of river flow and individual parameters in migratory route selection. The two routes differ significantly in smolt survival probabilities (Perry et al. N Am J Fish Manag 30:142–156, 2010), thus a clearer understanding of the variables contributing to route selection will be valuable for management of this declining species. A comparison of the two migratory groups showed that fish remaining within the Sacramento River: 1) Encountered the migratory juncture when river water velocities were much higher than those in the Delta Cross Channel (p?<?0.0001), 2) showed more direct swimming paths (p?=?0.03) and 3) migrated at higher speeds (p?=?0.04). Logistic regression models showed that the ratio of mean water velocity between the two routes was a much stronger predictor of ultimate route selection than any other variable tested. However, parameters for both the lateral position of smolts within the river and smolt size added predictive power to the final model. Our results suggest that river flow remains the most important variable for predicting smolt migration route, but note that knowledge of individual smolt attributes and movement patterns can increase our predictive ability.     

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.     

Riverine and early ocean migration and mortality patterns of juvenile steelhead trout (<Emphasis Type="Italic">Oncorhynchus mykiss</Emphasis>) from the Cheakamus River,British Columbia

High mortality (65–73%) occurred in the first month of the smolt migration in a population of wild steelhead trout. We used acoustic telemetry to monitor the downstream, estuarine, and early ocean migration of tagged smolts and estimate their mortality rates. After entering the Strait of Georgia most smolts migrated north through Johnstone and Queen Charlotte Straits rather than south through the Strait of Juan de Fuca. Of 51 smolts tagged in 2004 (49 in 2005), 36–38 (41–42) survived to leave freshwater and 14–19 (13–14) survived to leave the Strait of Georgia system. Mortality rates in separate segments of the migration were correlated with segment distances. An additional component of mobile sampling showed that few smolts died during the migration through Howe Sound. Migration rates averaged 0.7–0.9 body lengths per second (BL s⁻¹) downstream and 1.0–2.6 BL s⁻¹ in ocean waters. Aggregated detection probabilities of 92–96% on lines of ocean receivers suggest that migration routes of small fishes can be quantified over several hundred kilometres, and survival rates can be estimated for even a modest number of tagged fish. Quantifying mortality patterns during the smolt migration could help to determine causes of low marine survival rates observed in recent years.     

Strontium content of scales as a marker for distinguishing between sea trout and brown trout

Strontium was determined in trout scales from a river where it is often difficult to distinguish between sea trout and resident brown trout by coloration or other visual marks. Sr values were compared with values in scales from brown trout caught above the anadromous stretch of the same river and in scales from a river where sea trout coloration is typical. In the first river, the Sr concentration was generally low, and as a mean only 50 ppm higher in scales from individuals classified as sea trout from the anadromous stretch than in brown trout scales from the upper stretch. There was no consistency between fish coloration and Sr concentration in scales from presumed sea trout on the anadromous stretch. Individuals with a typical sea trout coloration could have a lower concentration of Sr than individuals that were classified as uncertain sea trout by coloration. Fish weight did not seem to influence Sr levels. The mean Sr concentration in scales from the typical sea trout colored population in the second river was 2.8 times higher than that of the anadromous part of the first river. The high variability of Sr concentration in sea trout scales may be explained by differences in individual and population life history. The Sr levels reflect differences in saltwater exposure, either expressed by length of stay or concentration of salt in marine habitats. The study has shown that fish coloration is an inadequate mean to distinguish between resident and migratory trout. Nor is Sr determination of scales alone sufficient, because of low inter-group and high intra-group variability in some rivers. However, Sr values can give valuable information on individual and population migration on a large scale.     

The relationship between smolt density and fry density in salmonids

If the Ricker stock–recruitment model describes the relationship between egg density and survivor density at different stages of the life cycle, then the relationship between smolt density and fry density is not simple. Using data from a long-term study (1966–1990) of a sea trout population Salmo trutta , the relationship between density of potential smolts and fry densities in late M ay/early June or late August/early September is shown to be a reflexed curve with zero origin, so that there are two smolt densities for each fry density. A linear relationship is obtained only when the exponential parameter in the Ricker model is constant for the different life Stages.     

Influence of hydrologic attributes on brown trout recruitment in low-latitude range margins

Factors controlling brown trout Salmo trutta recruitment in Mediterranean areas are largely unknown, despite the relevance this may have for fisheries management. The effect of hydrological variability on survival of young brown trout was studied during seven consecutive years in five resident populations from the southern range of the species distribution. Recruit density at the end of summer varied markedly among year-classes and rivers during the study period. Previous work showed that egg density the previous fall did not account for more than 50% of the observed variation in recruitment density. Thus, we expected that climatic patterns, as determinants of discharge and water temperature, would play a role in the control of young trout abundance. We tested this by analyzing the effects of flow variation and predictability on young trout survival during the spawning to emergence and the summer drought periods. Both hatching and emergence times and length of hatching and emergence periods were similar between years within each river but varied considerably among populations, due to differences in water temperature. Interannual variation in flow attributes during spawning to emergence and summer drought affected juvenile survival in all populations, once the effect of endogenous factors was removed. Survival rate was significantly related to the timing, magnitude and duration of extreme water conditions, and to the rate of change in discharge during hatching and emergence times in most rivers. The magnitude and duration of low flows during summer drought appeared to be a critical factor for survival of young trout. Our findings suggest that density-independent factors, i.e., hydrological variability, play a central role in the population dynamics of brown trout in populations from low-latitude range margins. Reported effects of hydrologic attributes on trout survival are likely to be increasingly important if, as predicted, climate change leads to greater extremes and variability of flow regimes.