Does size matter? A test of size‐specific mortality in Atlantic salmon Salmo salar smolts tagged with acoustic transmitters

Mortality rates of wild Atlantic salmon Salmo salar smolts implanted with acoustic transmitters were assessed to determine if mortality was size dependent. The routinely accepted, but widely debated, ‘2% transmitter mass: body mass’ rule in biotelemetry was tested by extending the transmitter burden up to 12·7% of body mass in small [mean fork length (L_F) 138·3 mm, range 115–168 mm] downstream migrating S. salar smolts. Over the short timescale of emigration (range 11·9–44·5 days) through the lower river and estuary, mortality was not related to S. salar size, nor was a relationship found between mortality probability and transmitter mass: body mass or transmitter length: L_F ratios. This study provides further evidence that smolt migration studies can deviate from the ‘2% rule’ of thumb, to more appropriate study‐specific measures, which enables the use of fishes representative of the body size in natural populations without undue effects.     

Precision and accuracy of Dahl-Lea back-calculated smolt lengths from adult scales of Atlantic salmon Salmo salar

Using tagged and recaptured Atlantic salmon Salmo salar (n = 106) the present analysis shows that the most commonly applied linear back-calculation method for estimating past length, the Dahl-Lea method, resulted in overestimation of the length of large smolts and underestimation of small smolts. A correction equation (y = 0.53x + 6.23) for estimating true smolt length (y) from lengths back-calculated from adult scale measures (x) to account for these systematic discrepancies is proposed.     

Polymorphism of smolts of pink salmon <Emphasis Type="Italic">Oncorhynchs gorbuscha</Emphasis> in the Indera River (Kola Peninsula)

Pink salmon introduced into the White Sea started to exploit as spawning grounds middle and upper reaches of the river 20 years after its appearance in the Indera River. As a result of this, the migration pathway of smolts and late smolts appeared in addition to early smolts. The intraspecies polymorphism of smolts is confirmed by differences of early and late smolts by body length and weight, migration dates, food spectrum, and indices of stomach fullness. The food spectra of late juveniles of pink salmon coincide with those of parr of Atlantic salmon Salmo salar and of brown trout S. trutta. Greater abundance of late migrants of pink salmon may cause competition of these species for food.     

Net ground speed of downstream migrating radio-tagged Atlantic salmon (Salmo salar L.) and brown trout (Salmo trutta L.) smolts in relation to environmental factors

The downstream migration of Atlantic salmon (Salmo salarL.) and sea trout smolt (S. trutta L.) was investigated using radio telemetry in the spring of 1999 and 2000. Forty wild sea trout smolts, 20 F1 sea trout smolts, 20 hatchery salmon smolts and 20 salmon smolts from river stockings were radio tagged and released in the Danish River Lilleaa. The downstream migration of the different groups of fish was monitored by manual tracking and by three automatic listening stations. The downstream migration of radio tagged smolts of both species occurred concurrently with their untagged counterparts. The diel migration pattern of the radio tagged smolts was predominantly nocturnal in both species. Wild sea trout smolt migrated significantly faster than both the F1 trout and the introduced salmon. There was no correlation between net ground speed, gill Na⁺,K⁺-ATPase activity or fish length in any of the different groups. The migration speed of wild sea trout smolts was positively correlated with water discharge in both years. In F1 sea trout smolts, migration speed was positively correlated with temperature in 1999. The migration speed of salmon smolts did not correlate to any of the investigated parameters.     

Body and scale growth of wild Atlantic salmon smolts during seaward emigration

The relationship between scale and body growth for emigrating Atlantic salmon, Salmo salar, smolts was previously not understood and therefore was examined in this study using mark-recapture techniques. The size of smolts at time of recapture was significantly greater than when marked (P = 0.0002). The growth in length of smolts emigrating 5 km over an average of 20 days was 7.7 ± 6.1 mm per day. Instantaneous somatic growth (G _body) ranged from 7.0 × 10⁻⁴ to 5.1 × 10⁻³ (mean = 2.7 × 10⁻³ ± 1.3 × 10⁻³). The mean number of plus growth circuli present per scale was significantly greater for smolts when recaptured compared to when marked (P = 0.0014). The instantaneous growth rate of scales (G _scale) ranged from 1.4 × 10⁻³ to 11.5 × 10⁻³ (mean = 6.6 × 10⁻³ ± 4.3 × 10⁻³). The relationship between body size and scale radius showed positive allometry rather than isometry. The relationship of G _scale with G _body showed positive allometry indicating that scales grew at a slightly faster rate than the body during the emigratory period.     

Size-independent growth in fishes: patterns, models and metrics

A combination of a dynamic energy budget (DEB) model, field data on Atlantic salmon Salmo salar and brown trout Salmo trutta and laboratory data on Atlantic salmon was used to assess the underlying assumptions of three different metrics of growth including specific growth rate (G), standardized mass‐specific growth rate (G_S) and absolute growth rate in length (G_L) in salmonids. Close agreement was found between predictions of the DEB model and the assumptions of linear growth in length and parabolic growth in mass. Field data comparing spring growth rates of age 1+ year and 2+ year Atlantic salmon demonstrated that in all years the larger age 2+ year fish exhibited a significantly lower G, but differences in growth in terms of G_S and G_L depended on the year examined. For brown trout, larger age 2+ year fish also consistently exhibited slower growth rates in terms of G but grew at similar rates as age 1+ year fish in terms of G_S and G_L. Laboratory results revealed that during the age 0+ year (autumn) the divergence in growth between future Atlantic salmon smolts and non‐smolts was similar in terms of all three metrics with smolts displaying higher growth than non‐smolts, however, both G_S and G_L indicated that smolts maintain relatively fast growth into the late autumn where G suggested that both smolts and non‐smolts exhibit a sharp decrease in growth from October to November. During the spring, patterns of growth in length were significantly decoupled from patterns in growth in mass. Smolts maintained relatively fast growth though April in length but not in mass. These results suggest G_S can be a useful alternative to G as a size‐independent measure of growth rate in immature salmonids. In addition, during certain growth stanzas, G_S may be highly correlated with G_L. The decoupling of growth in mass from growth in length over ontogeny, however, may necessitate a combination of metrics to adequately describe variation in growth depending on ontogenetic stage particularly if life histories differ.     

A size-dependent migration strategy in Atlantic salmon smolts: Small smolts favour nocturnal migration

Migration theory states that migration behavioural strategies should be optimised to maximise fitness. Many studies have shown that in downstream migrating Atlantic salmon Salmo salar L. smolts, mortality from predation is high and negatively size dependent. The most common predators are birds and piscivorous fish that are mainly daylight feeders. Given the high mortality during this stage we should expect to observe smolts to follow predator avoidance strategies that may be affected by body size. We tested the hypothesis that small smolts have a higher tendency to exhibit predator avoidance strategies (i.e. nocturnal versus diurnal migration) than larger smolts. The number and size of out-migrating/downstream-migrating wild Atlantic salmon smolts was recorded as they passed through a glass-sided channel during April-May, 1996–1999. In all years, the mean size of nocturnal migrating smolts was significantly lower than the mean size of diurnal migrating smolts. Analysis of the size of smolts, during early and late stages of the migration period showed size-dependent nocturnal migration behaviour up to the end of April. After this, no such size dependent migration pattern was observed. However, small smolts (<100 mm) were absent during this period. We suggest that nocturnal migration is an adaptive behaviour that small Atlantic salmon smolts have to avoid predation by large daylight feeding visual piscivorous predators (e.g. pike Esox Lucius L. and fish eating birds).     

Migration and survival of Atlantic salmon Salmo salar smolts in a large natural lake

An investigation with acoustic telemetry of the passage of Salmo salar smolts through a large natural lake found heavy mortality occurred at the river‐to‐lake confluences (mean 31.2% km^?1), but was lower in the main body of the lake (mean 2.4% km^?1). Predation was a significant pressure on emigrating smolts as tagged pike Esox lucius aggregated at river‐to‐lake confluences during the peak of the smolt run. Tagged smolts mainly emmigrated into the lake in the late evening after dusk, possibly as a predator‐avoidance behaviour.     

Individual growth and phase differentiation of lacustrine masu salmon, Oncorhynchus masou, under artificial rearing conditions

 In lacustrine masu salmon, Oncorhynchus masou, originated from anadromous fish and inhabiting an artificial lake (Shumarinai Lake), we examined the relationship between individual growth during the juvenile stage and phase differentiation under artificial rearing conditions. In females, the mean fork length of potential (subsequent) 1+ smolts and potential 1+ parr (1+ parr show fish that neither smolted nor matured until 1+ autumn) always differed after their first summer (0+ summer), the former being larger. In males, the juveniles that grew faster during their first spring became 0+ mature male parr. After their first summer, potential 1+ smolts were always larger in body length than potential 1+ mature parr and potential 1+ parr. These results were similar to previous reports of anadromous masu salmon, suggesting that the lacustrine masu salmon studied seems to have maintained the phase differentiation as in the original fish, likely because only 60 years have passed since the formation of the lacustrine population.     

The effect of domestication on some life history traits of sea trout and Atlantic salmon

During the period 1968–1991, certain morphological traits of Atlantic salmon Salmo salar and sea trout S. trutta have been recorded regularly at the hatchery at Älvkarleby, central Sweden. Total body length, weight (for females both before and after stripping), number of eggs, egg size and date of ovulation. A smaller data set for fish marked and released as smolts, providing information about total body length of released smolts, time spent at sea and body size of the recovered adults, was also available for analysis. According to theory and empirical data, the process of artificial breeding results in an evolutionary divergence of the cultured strain from the wild phenotypic norm. The reason for such a divergence is that both natural and sexual selection pressures are altered or relaxed during the process of artificial breeding, as well as random genetic processes, such as founder effects and in- and outbreeding. A path analysis of both species and sexes revealed that the size of the released smolts had increased during the study period. Time spent in sea has decreased for both female and male sea trout, but not for Atlantic salmon. Adult body size for female and male trout have increased as well as female trout condition factor. The increase found in egg size of both species was greater for the salmon than for the trout, indicating that female salmon invest more in egg size with increasing body size. These results support the view that domestication probably has a significant and selective impact on the life history traits of the two salmonid species studied.     

Development of seawater tolerance and subsequent downstream migration in wild and stocked young‐of‐the‐year derived Atlantic salmon Salmo salar smolts

This study investigated the development of hypo‐osmoregulatory capacity and timing of downstream migration in wild Atlantic salmon Salmo salar smolts from the River Stjørdalselva and stocked young‐of‐the‐year (YOY), derived S. salar smolts from the tributary River Dalåa. Both wild and stocked S. salar smolts developed seawater (SW) tolerance in early May, persisting through June, measured as their ability to regulate plasma osmolality and chloride following 24 h SW (salinity = 35) exposure. Although the majority of downstream migration among the stocked S. salar smolts occurred later than observed in their wild counterparts, the development of SW tolerance occurred concurrently. The wild S. salar from Stjørdalselva and stocked YOY smolts from the River Dalåa started to migrate on the same cumulative day‐degrees (D°). The study revealed no downstream migration before development of SW tolerance. This emphasizes the importance of incorporating physiological status when studying environmental triggers for downstream migration of S. salar smolts. Overall, these findings suggest that the onset of smolt migration in stocked S. salar smolts was within the smolt window from an osmoregulatory point of view.     

Individual growth and smoltification of juvenile masu salmon, Oncorhynchus masou Brevoort, under rearing conditions

In a reared population of individually marked juvenile masu salmon, individual growth was monitored from the first autumn in 1983 to the following spring. The potential smolts were not significantly greater in mean fork length and body weight than the potential parr in late August of the first year, but they then grew faster until March of the second year. As a result, the potential smolts formed the upper mode of the bimodal length distribution after February. Especially in autumn (October and November) the specific growth rates of potential smolts were significantly greater than those of parr, and the bimodality in growth rate distribution was more distinct for males than females. These suggest that there are two groups having different growth rates in autumn of the first year and that sufficient growth in this period may play an important role in smoltification in the following spring.     

Riverine and near coastal migration performance of hatchery brown trout Salmo trutta

To study migration performance and return rates of hatchery brown trout Salmo trutta smolts the first 5 months after release, 50 fish in each year (fork length, L_F, 158–288 mm) were in two subsequent years tagged with acoustic transmitters and recorded by automatic listening stations in the River Nidelva (central Norway), its estuary and in the marine environment. More than half of the smolts became anadromous migrants (52% in 2011 and 70% in 2012). The fish spent longer time in the estuary than in the marine environment and the results suggest that migratory behaviour of S. trutta smolts is not only restricted to be resident or anadrome–lacustrine, but that there is also an intermediary strategy of estuarine feeding. There were no differences in L_F or mass between groups of smolts with different migration patterns. Return rates from the sea within the first 5 months after release were in both years 16%. Median progression rate in the river was 0·090 L_F s^?1 but decreased significantly as the smolts entered the estuary (0·015 L_F s^?1). The long residential time in the estuary may increase the risk of negative effects of anthropogenic activities in estuaries, such as harbours and industrial development, and special attention should be given to evaluate effects of such activities.     

Time and size at seaward migration influence the sea survival of Salmo salar

Whether time of seaward migration of young Atlantic salmon Salmo salar influences their subsequent survival and growth was investigated in the River Imsa, south‐western Norway. Salmo salar were tagged when moving downstream through a trap near the outlet between 1976 and 2010 and recaptured on their adult return. Most descended as smolts in April and May, but some descended during the other months of the year. Annual variation in timing of the smolt migration was significantly correlated with variation in water temperature during spring. Mean total body length of the descending S. salar varied with month of seaward migration. The sea survival of S. salar emigrating from the River Imsa between January and May was 2·8 times higher than for those descending between June and December. The sea survival of the various cohorts decreased with increasing river temperature in April to May, prior to the smolt migration, and decreasing day number when the smolts moved to sea. The size of smolts descending the river between April and May did not affect the survival at sea as much as it affected the survival of migrants descending in any other month of the year. The majority of the downstream migrating S. salar were 2 years old, but proportionally, more 1 year olds moved downstream in the autumn than in the rest of the year. Mean duration between downstream migration of the young and the return migration of the grilse was shortest (12·7 months) for those descending in July and August and longest for those descending in October (21 months). Mean monthly specific growth rate was highest for those migrating downstream between May and July and lowest for those emigrating in September. Based on the present results, it was hypothesized that S. salar emigrating between April and August migrated directly out into the ocean, while those that emigrated between October and March stayed in the estuary until the subsequent spring.     

Migration of Atlantic Salmon, Salmo salar, Smolt through the Estuary Area of River Ellidaar in Iceland

Synopsis We tagged both wild and hatchery Atlantic salmon, Salmo salar, smolts from River Ellidaar (64 ° 08′ N, 21 ° 50′ W) with ultrasonic tags. We caught the wild smolts in a smolt trap and selected the largest individuals from the run. We implanted the transmitters in the abdominal cavity of the fish and then released them in River Ellidaar close to the estuary. We used four ultrasonic receivers; one in the river’s estuary, one outside the estuary and two further away on both sides of an island in the estuary zone. The receivers recorded all transmitters within a 600 m radius. The hatchery smolts were larger than the wild smolts. Some of the smolts were lost on the way through the estuary. Only 4 out of 9 wild smolts and 14 of 17 hatchery smolts were recorded all the way through. The tags and the tagging likely affected the survival of the smolts especially the smaller fish. There were no differences in the smolt migration between the 2 years of study and no differences in the migration behavior between the wild and the hatchery smolts. After being released the smolts stayed on average for 10 h in the river then migrated into the estuary were they stayed for 54 h on average. Then they migrated straight through the area to the sea at approximately 0.2 fish lengths per second. We recorded large differences in the migration.     

Evidence of changing migratory patterns of wild Atlantic salmon Salmo salar smolts in the River Bush,Northern Ireland,and possible associations with climate change

The migration patterns, timing and biological characteristics of wild Atlantic salmon Salmo salar smolts in the River Bush, Northern Ireland, were examined over the period 1978–2008. A distinct change in the timing of the smolt run was detected with progressively earlier emigration periods evident across the time series. The shift in run timing ranged from 3·6 to 4·8 days 10 years^?1 for a range of standard migratory audit points. The timing of smolt emigration has been linked to ambient river temperature patterns. Distinct seasonal patterns were evident for biological characteristics of River Bush smolts with mean age and fork length decreasing throughout the emigration period. Marine survival patterns in 1 sea winter River Bush S. salar were strongly influenced by the run timing of the preceding smolt year such that later emigrating cohorts demonstrated increased survival. Possible mechanisms for this relationship based on local climatic variation have been explored, including the effect of potential thermal mismatch between freshwater and marine environments.     

Changes in smolt traits of Atlantic salmon (Salmo salar Linnaeus, 1758) and linkages to parr density and water temperature

Smolt traits (length, age) and timing of smolt migration of wild Atlantic salmon, Salmo salar L., were investigated in the Simojoki River, northern Baltic Sea. The aim was to determine whether they responded to changes in parr length, parr density and temperature from 2000 to 2014. Annual electrofishing surveys and smolt numbers determined parr densities by springtime trapping in the river mouth. During the smolt trapping period captured parr and smolts were aged from scales. Water temperature was measured daily. Mean length decreased from 137 mm (TL) to 129 mm among 2‐year‐old smolts, and from 150 mm to 139 mm among 3‐year‐olds. Median date of the smolt migration was 10 days earlier, from early June to late May during the study period, linked to the rise in air temperature in May at the nearby Kemi‐Tornio airport. However, the median day temperature and the mean daily water temperatures during the second (Q₂) and third (Q₃) migration quartiles did not change. This implied that migration began when a suitable water temperature was reached, independent of the date.     

Passing a seawater challenge test is not indicative of hatchery‐reared Atlantic salmon Salmo salar smolts performing as well at sea as their naturally produced conspecifics

Despite satisfactory reactions to seawater challenge tests indicative of appropriate physiological state, hatchery‐reared Atlantic salmon Salmo salar smolts stocked in the Eira River in Norway between 2001 and 2011 performed less well at sea in terms of growth, age at maturity and survival than smolts of natural origin. The mean rates of return to the river for hatchery‐reared and naturally produced S. salar were 0·98 and 2·35%. In the Eira River, c. 50 000 hatchery‐reared S. salar smolts of local origin were stocked annually to compensate for reduced natural smolt production following regulation for hydroelectric purposes, while a mean of 17 262 smolts were produced naturally in the river. This study demonstrates that, although captive S. salar perform well in seawater challenge tests, hatchery‐reared smolts are not necessarily as adaptable to marine life as their naturally produced counterparts. These findings suggest that production of hatchery‐reared smolts more similar to naturally produced individuals in morphology, physiology and behaviour will be necessary to improve success of hatchery releases. Where possible, supplementary or alternative measures, including habitat restoration, could be implemented to ensure the long‐term viability of wild stocks.     

Survival and migration patterns of naturally and hatchery-reared Atlantic salmon (Salmo salar) smolts in a Lake Ontario tributary using acoustic telemetry

1. Atlantic salmon (Salmo salar) smolts are often stocked into rivers to supplement natural reproduction, yet hatchery-reared fish have lower survival compared to wild conspecifics. However, few studies have assessed riverine migratory performance and survival differences in hatchery and wild smolts, or more specifically naturally reared smolts (hatchery fish released earlier as parr), particularly in rivers with weirs which may further reduce survival.
2. Using acoustic telemetry, including a subset of fish with novel transmitters that identify predation events, we assessed survival and migration patterns of hatchery- (2017: n = 32; 2018: n = 30) and naturally reared Atlantic salmon smolts (2017: n = 8; 2018: n = 30) in a Lake Ontario tributary with two weirs to better understand their ecology and assess the influence of environmental parameters on migration.
3. Naturally reared smolts were 13.9 times more likely to survive than hatchery-reared smolts and mark–recapture models indicated that weirs did not reduce survival for either group. Survival per km was lowest at the release site, indicating pre-migration mortality, and specifically high stocking-related mortality of hatchery-reared smolts. Speed and times of day fish migrated (i.e. migratory performance) did not vary by rearing group, suggesting that the high mortality of hatchery-reared smolts may be due to other factors related to hatchery and stocking operations. Overall mean (± SD) migration speed for smolts was 0.70 ± 0.39 km/hr and movements occurred significantly more frequently at night (18:00–06:00).
4. Smolts were detected in Lake Ontario after they left the river; however, the array in Lake Ontario was not conducive to providing much detail regarding movement patterns. There was no predation of the two predation tags detected in Lake Ontario, indicating that movements were made by smolts and not predators.
5. With ongoing restoration efforts of Atlantic salmon in Lake Ontario, it was important to understand the smolt migration patterns and success of the stocked fish. Our findings of similar migratory performance yet different relative survival of hatchery- and naturally reared smolts help inform management with regards to stocking strategies that could improve Atlantic salmon reintroduction success.

     

Evidence for non-random spatial positioning of migrating smolts (Salmonidae) in a small lowland stream

1. The ontogenetic development of anadromous salmonids includes downstream emigration of immature individuals from freshwater towards the marine environment. Although this migration of juvenile salmonids (smolts) may be associated with severe mortalities, only limited attention has been paid to the spatial positioning of smolts in small streams. 2. Using a novel approach, this study examined the vertical and horizontal positioning of brown trout and Atlantic salmon smolts while performing downstream migration in a small lowland stream. 3. Pre‐smolts of indigenous and hatchery‐reared (F1) brown trout (Salmo trutta), and two different populations of Atlantic salmon (S. salar), were tagged with passive integrated transponder (PIT) tags and subsequently released upstream of an antenna array consisting of five circular swim‐through PIT antennas. Antennas were positioned in order to determine whether the migrating smolts were bottom or surface oriented, and if they were oriented towards the mid‐channel or the stream bank. 4. During the smolt emigration period, data describing both the detection of the migrating fish and the amount of water passing through the antennas were collected. This was accomplished in order to determine if the fish were performing active positioning behaviour independently of the vertical and horizontal discharge distributions in the stream. 5. The results showed that the smolts migrated in a non‐random spatial pattern independently of the stream discharge distributions. Vertically, the indigenous brown trout and the Atlantic salmon demonstrated a preference for the bottom orientated positions. In contrast, the distribution of the F1 brown trout was not different from the discharge distribution. The latter observation suggests random vertical positioning, which may be indicative of inferior migratory performance. Horizontally, all tested smolt populations strongly preferred the mid‐channel positions. 6. The discharge‐corrected preferences for certain spatial positions suggest that smolt emigration is not entirely a matter of passive displacement in lowland streams. 7. Anthropogenically altered channels may inhibit or delay downstream emigration of smolts resulting in increased mortalities. Given that the smolts in this study actively selected spatial positions in the mid‐channel and near the bottom, it is suggested that deep, mid‐channel furrows may be used to help guide migrating smolts past adverse habitats in lowland streams.