Timing matters: species‐specific interactions between spawning time,substrate quality,and recruitment success in three salmonid species

Substratum quality and oxygen supply to the interstitial zone are crucial for the reproductive success of salmonid fishes. At present, degradation of spawning grounds due to fine sediment deposition and colmation are recognized as main factors for reproductive failure. In addition, changes in water temperatures due to climate change, damming, and cooling water inlets are predicted to reduce hatching success. We tested the hypothesis that the biological effects of habitat degradation depend strongly on the species‐specific spawning seasons and life‐history strategies (e.g., fall‐ vs. spring‐spawners, migratory vs. resident species) and assessed temperature as an important species‐specific factor for hatching success within river substratum. We studied the species‐specific differences in their responses to such disturbances using egg‐to‐fry survival of Danube Salmon (Hucho hucho), resident brown trout (Salmo trutta fario), and migratory brown trout (Salmo trutta lacustris) as biological endpoint. The egg incubation and hatching success of the salmonids and their dependence on temperature and stream substratum quality were compared. Hatching rates of Danube salmon were lower than of brown trout, probably due to higher oxygen demands and increased interstitial respiration in spring. Increases in maximum water temperature reduced hatching rates of resident and migratory brown trout (both fall‐spawners) but were positively correlated with hatching rates of Danube salmon (a spring‐spawner). Significantly longer incubation periods of resident and migratory brown trout coincided with relatively low stream substratum quality at the end of the egg incubation. Danube salmon seem to avoid low oxygen concentrations in the hyporheic zone by faster egg development favored by higher water temperatures. Consequently, the prediction of effects of temperature changes and altered stream substratum properties on gravel‐spawning fishes and biological communities should consider the observed species‐specific variances in life‐history strategies to increase conservation success.     

Climate warming causes phenological shift in Pink Salmon, Oncorhynchus gorbuscha, behavior at Auke Creek, Alaska

Thirty‐four years (1972–2005) of water temperature data and extensive biological observations at Auke Creek, Alaska indicate a general warming trend that affected the native pink salmon (Oncorhynchus gorbuscha) population. Serial environmental records at nearby Auke Bay, Alaska over 46 years show trends of increasing air and sea surface temperatures. Trends of increased total precipitation and earlier date of ice out on nearby Auke Lake also occurred, but not at significant rates. Average water temperatures during the incubation of pink salmon in Auke Creek increased at a rate of 0.03 °C yr^?1 over the 34‐year period. For the 1972–2005 broods, midpoints of fry migrations from Auke Creek ranged between April 2 and May 7, and there was a trend of earlier migration of pink salmon fry at a rate of ? 0.5 days yr^?1. The migration timing of adult salmon into Auke Creek also showed a trend toward earlier timing. The earlier adult migration combined with warmer incubation temperatures are related to earlier migration of pink salmon fry. If the observed warming trend continues, Auke Creek may become unsuitable habitat for pink salmon. Given the trend for salmon fry to migrate earlier, a larger portion of the population may become mismatched with optimum environmental conditions during their early marine life history. If salmon adults continue to migrate into the creek earlier when water temperatures are commonly high, it will result in increased prespawning mortality.     

Rearing and migration of juvenile Chinook salmon (<Emphasis Type="Italic">Oncorhynchus tshawytscha</Emphasis>) in a large river floodplain

Off-channel habitat has become increasingly recognized as key for migratory fishes such as juvenile Chinook salmon (Oncorhynchus tshawytscha). Hence, floodplain habitat has been identified as critical for the continued persistence of California’s Central Valley salmon, particularly the Yolo Bypass, the primary floodplain of the Sacramento River. To provide insight into factors supporting juvenile salmon use of this 240 km², partially leveed floodplain, we examined inter- and intra-annual relationships between environmental correlates and residency time, apparent growth, emigration, migratory phenotype, and survival over more than a decade for natural-origin (“wild”) fish and experimentally-released hatchery fish. Flood duration was positively associated with hatchery juveniles residing longer and achieving larger size. Wild juveniles grew larger and emigrated later with cumulative temperature experience (accumulated thermal units) and warmer average annual temperatures during flood years. Within years, both wild and hatchery salmon departed the floodplain as flood waters receded. Parr-sized juveniles dominated outmigrant composition, though fry and smolt-sized juveniles were also consistently observed. Survival to the ocean fishery was not significantly different between hatchery fish that reared in the Yolo Bypass versus those that reared in the main stem Sacramento River. Our study indicates improved frequency and duration of connectivity between the Sacramento River and the Yolo Bypass could increase off-channel rearing opportunities that expand the life history diversity portfolio for Central Valley Chinook salmon.     

Use of radio telemetry and electrofishing to assess spawning by transplanted Atlantic salmon

The river Ingdalselva, which drains to the Trondheims fjord, has no local salmon population due to an impassable waterfall 500 m upstream from the outlet. In the period 1994–97, a total of 31 mature Atlantic salmon (19 females and 12 males) from the rivers Orkla (1994–96) and Vigda (1997) were radio-tagged and released in the river Ingdalselva before spawning. The main goal of the project was to find out whether the fish would stay and spawn in the river, and if the observations during the spawning period could be used to indicate where spawning had taken place. Some fish left the river shortly after release, but 77% of the fish stayed in the river during the spawning period in October. Most of the females (74%) spawned in the river, including multi-sea winter salmon of approximately 10 kg. Some fish remained at the site of the release, while others migrated downstream to hiding places where they stayed until spawning. Long distance upstream migrations were not observed. The observations of the fish during the spawning period were used to identify spawning areas. Electrofishing for salmon fry in the spawning areas the following year in all cases produced fry, and in two of the years salmon fry were also found in areas where no spawning activity had been registered. Radio-tracking was an efficient method for determining whether transplanted salmon would remain in a `new'n river during the spawning season and for locating spawning areas, particularly when the fish were tracked daily.     

The pre‐spawning migratory behaviour of Atlantic salmon Salmo salar in a large lacustrine catchment

The movements of adult Atlantic salmon Salmo salar were determined as they migrated to spawning habitats in a large lacustrine catchment, Lough Neagh, in Northern Ireland. The minimum average ground speed of S. salar through the lake was 2·1 km day^?1 and the mean residence time was 11 days. Tagged S. salar tended to actively migrate through the lake which represented a transitory habitat for adult S. salar. Migration time from the release site, through the lake, to a spawning tributary decreased during the migratory period. During the 4 year study period between 20·5 and 41·6% of tagged S. salar which entered the lake each year, explored at least one other channel before ascending the final spawning tributary. Exploratory behaviour was more likely in S. salar which spawned in the tributaries furthest from the sea. Exploratory behaviour was also more likely to occur during periods of reduced discharge in the natal stream. The fishery management implications of complex pre‐spawning behaviour in a mixed stock lacustrine system, are discussed.     

Juvenile habitat partitioning and relative productivity in allochronically isolated sockeye salmon (Oncorhynchus nerka)

Allochronic divergence, like spatial isolation, may contribute to population diversity and adaptation, however the challenges for tracking habitat utilization in shared environments are far greater. Adult Klukshu River (Yukon, Canada) sockeye salmon, Oncorhynchus nerka, return as genetically distinct "early" and "late" runs. Early and late adult spawning populations (1999 and 2000) and their subsequent fry (sampled at 7 sites in 2000 and at 8 sites in 2001 throughout Klukshu Lake and River) were genotyped at eight microsatellite loci. Bayesian assignment was used to determine the spatial distribution of early versus late fry; although intermixed, the distribution of fry significantly differed in Klukshu Lake and in the Klukshu River in 2001, based on crosstab analyses. Late-run fry predominated in Klukshu Lake at all sites, while early-run fry were most common in the north and south of Klukshu Lake and in Klukshu River. Early-run spawners had significantly higher relative productivity (early life survival) than late-run fish (2.9 times more fry produced per early-run adult in 2000, and 9.2 times more in 2001). This study demonstrates spatial habitat partitioning and differences in the contribution of allochronically isolated populations to fry abundance, and highlights annual variability that likely contributes to recruitment variation.     

Olfactory gene expression in migrating adult sockeye salmon Oncorhynchus nerka

Expression of 12 olfactory genes was analysed in adult sockeye salmon Oncorhynchus nerka nearing spawning grounds and O. nerka that had strayed from their natal migration route. Variation was found in six of these genes, all of which were olfc olfactory receptors and had lower expression levels in salmon nearing spawning grounds. The results may reflect decreased sensitivity to natal water olfactory cues as these fish are no longer seeking the correct migratory route. The expression of olfactory genes during the olfactory‐mediated spawning migration of Pacific salmon Oncorhynchus spp. is largely unexplored and these findings demonstrate a link between migratory behaviours and olfactory plasticity that provides a basis for future molecular research on salmon homing.     

Return migration of Atlantic Salmon in the River Tana: Phases of migratory behaviour

From a total of 174 multi-sea-winter Atlantic salmon radio tagged in the Tanafjord (northern Norway, 70°N) during 1992 and 1993, 48 Atlantic salmon were followed from entering the River Tana until spawning. Three phases were identified: (1) migratory, direct or stepwise migration to, or close to the position held at spawning; (2) search, movements both up and down river at or close to the position held at spawning; (3) holding, a period without movements prior to spawning. During the migratory phase, Atlantic salmon migrated directly to near the spawning area, or stopped between one and nine shorter periods during the upstream migration. Number of stops increased with increasing migratory distance in 1993, but no such correlation was found in 1992. The highest migratory speeds were recorded in the lower parts of the river. A distinct change in migratory pattern was found in 67% of the Atlantic salmon near or at the area held at spawning. Most common was a search phase of erratic movements with more than one down river movement. After the movement terminated, 96% of the Atlantic salmon had a period when no or little movement was recorded until spawning (on average 55 days in 1992 and 51 days in 1993). There was no preference for staying at, up or down river from the spawning area during this holding period. Early ascending Atlantic salmon migrated to spawning areas further from the mouth than the later arriving Atlantic salmon in 1993, but not in 1992. The proportion of time spent on the migratory phase increased, while the proportion of time spent on the holding phase decreased with increasing distance to the spawning area.     

Efficacy of releasing captive reared broodstock into an imperilled wild Atlantic salmon population as a recovery strategy

The strategy of releasing captive reared adult Atlantic salmon Salmo salar into the Magaguadavic River, New Brunswick, Canada, to spawn, was not an effective tool for rebuilding a seriously depressed wild population. The fish were first generation progeny from wild parents, and had spent their entire lives in captivity in either sea or fresh water. No differences in movement or behaviour patterns were observed between freshwater and seawater reared groups. Fish released in the lower river early (35 to 80 days prior to the natural spawning period) moved into a lake low in the system, and most stayed there near the commercial hatchery where they had been reared from egg to smolt. During the spawning season, none moved to the upper river reaches where most spawning habitat exists. Most broodstock released in the upper river reaches near the time of spawning stayed there during the spawning period. The following year few to no Atlantic salmon fry were found, and most appeared not to be offspring of released adults.     

Size-selective mortality of Sea of Okhotsk pink salmon in the ocean in the winter and spring

The mortality of Sea of Okhotsk pink salmon in the winter and spring varies significantly from year to year, which complicates forecasts of its arrival in the following year based on data on the downstream migration of fry and surveys in the fall. The size-selective mortality of pink salmon was studied and the possibility of using the size and weight parameters of juveniles for predicting their return was evaluated through measurements of scale increments in juvenile pink salmon that were caught in the southern Sea of Okhotsk in the fall of 2007 and 2008 and in fish of these year classes that came back to spawn. In the 2007 year class, which had a low overwinter survival level in the ocean, the average scale increments for the first year of life were considerably smaller than those in adult fish that returned to the spawning grounds. In the pink salmon of 2008, which had a very high level of overwinter survival, the values of scale increments in juveniles and adults were similar. This confirms the hypothesis of a critical size and a critical period, according to which slowly growing juveniles that do not accumulate enough energy reserves for summer are eliminated in the winter to a greater extent as compared to fast-growing fish. Correlation analysis revealed a significant negative relationship between the size and weight of juvenile pink salmon and their mortality in the ocean. After conducting further and more extensive studies this will allow using the size parameters of juvenile pink salmon as one of predictors of its return for the year following the fall surveys. These results emphasize how important it is to take the size and growth rate of juvenile salmon into account when forecasting their return.     

Persistent parental effects on the survival and size, but not burst swimming performance of juvenile sockeye salmon Oncorhynchus nerka

Sockeye salmon Oncorhynchus nerka were used as a model in an artificial fertilization experiment to investigate the relationships between individual adult O. nerka and their offspring. Survival, size and burst swimming ability were assessed in fry of known parentage (adult spawners from the Weaver Creek population, British Columbia, Canada). Maternal identity significantly affected the survival rate of eggs at hatch time, though this effect did not extend to fry life stages. The results were also suggestive of a paternal effect on both egg and fry survival, though this could not be separated from the experimental block design. After 4 months of exogenous feeding, fry mass remained under significant maternal influence, though fork length did not, despite having a high correlation with mass. Burst swimming performance was highly variable among individuals, and was not significantly influenced by maternal identity or individual fry size. Collectively, the findings presented here suggest that maternal, and possibly paternal, effects can be integral components of population dynamics in the early life stages of O. nerka . A good understanding of these factors will be essential for scientists and fisheries managers in developing a more holistic view of population-level spawning success and fry survival.     

Survival and growth of salmon, Salmo salar (L.), planted in a Scottish stream

Salmon eggs and unfed fry were planted in reaches (total length 2.8 km, mean width 4 m) of a Scottish stream between 1971 and 1977 and their subsequent progress was studied by sampling 16 sections (areas 38–126 m²) of the stream. Brown trout are the only fish which spawn in the stream, waterfalls and a dam near its mouth preventing adult salmon and sea-trout passing upstream. There were no restraints on the downstream movement of fish except in 1977, when a fry trap was operated. In 1971 and 1974 boxes each containing 300 eggs were buried in groups of 3–6. In other years fry were evenly distributed at 3.6–29.3 m^?2. At the end of the first growing season, salmon occurred at decreasing population densities for a distance of 600 m below the plantings, but after two growing seasons there was little remaining indication of their pattern of dispersion when planted. Rates of survival between planting and the end of the growing season were 9.4–31%. Survival when eggs were planted (11.1–14.8%) was not affected by the numbers planted together at one point (900–1800) or the distance apart of groups of boxes (10–85 m). When fry were planted the instantaneous mortality rate (M) of the 0₊ salmon during their first growing season was related to the initial stocking density (D_p) by the formula M= 0.00637 + 0.00444 log₁₀D_p. Twenty-two to 88% of 0₊ salmon present at the end of the growing season were still surviving in the stream as 1₊ fish one year later. In 1973–1976 only a small number of 2₊ salmon occurred, the majority having migrated between the end of the second growing season and the following spring. There were more 2₊ salmon in 1977 and 1978 resulting from higher stocking densities in 1975 and 1976 and slower growth. Trout of several age classes were present but their population densities were never high (<0.6 m^?2). Salmon reached a greater size than trout by the end of the first growing season. Their mean weight (W^o, g) at this time was inversely related to their population density (D_o No. m^?2) and the biomass (B₁, g m^?2) of 1₊ salmon present, giving the relationship log₁₀w^o= 0.6584–0.0558 D₀-0.0352B₁. The mean weight of 1₊ salmon tended to be highest in sections where the 0₊ salmon had reached a relatively large size the previous year. When a reach of the stream was planted twice (11 and 30 May 1977) with salmon fry (total 13.9 m^?2) at the same stage of development, M during the first growing season was 0.0099 per day. This was less than that of fry in a control (M= 0.0107) where the stocking density was lower (6.8 m^?2) and also less than in previous years when single planting rates of approximately 14 m^?2 were used (M=0.0115). The double planting resulted in a wide range of lengths of 0₊ salmon in September and the highest biomass values encountered during all experiments.     

Genetic versus Rearing-Environment Effects on Phenotype: Hatchery and Natural Rearing Effects on Hatchery- and Wild-Born Coho Salmon

With the current trends in climate and fisheries, well-designed mitigative strategies for conserving fish stocks may become increasingly necessary. The poor post-release survival of hatchery-reared Pacific salmon indicates that salmon enhancement programs require assessment. The objective of this study was to determine the relative roles that genotype and rearing environment play in the phenotypic expression of young salmon, including their survival, growth, physiology, swimming endurance, predator avoidance and migratory behaviour. Wild- and hatchery-born coho salmon adults (Oncorhynchus kisutch) returning to the Chehalis River in British Columbia, Canada, were crossed to create pure hatchery, pure wild, and hybrid offspring. A proportion of the progeny from each cross was reared in a traditional hatchery environment, whereas the remaining fry were reared naturally in a contained side channel. The resulting phenotypic differences between replicates, between rearing environments, and between cross types were compared. While there were few phenotypic differences noted between genetic groups reared in the same habitat, rearing environment played a significant role in smolt size, survival, swimming endurance, predator avoidance and migratory behaviour. The lack of any observed genetic differences between wild- and hatchery-born salmon may be due to the long-term mixing of these genotypes from hatchery introgression into wild populations, or conversely, due to strong selection in nature—capable of maintaining highly fit genotypes whether or not fish have experienced part of their life history under cultured conditions.     

The effect of stocking with 0+ year age‐class Atlantic salmon Salmo salar fry: a case study from the River Bush,Northern Ireland

An enhancement programme based on stocking 0+ year age‐class Atlantic salmon Salmo salar, conducted in the River Bush, Northern Ireland, U.K. over the period 1996–2005, was reviewed with reference to the performance and biological characteristics of wild fish. Wild ova to 0+ year fry (summer) survival was c. 8% with subsequent wild 0+ year fry‐to‐smolt survival c. 9%. Stocked unfed 0+ year juveniles gave c. 1% survival to smolt whilst fed 0+ year S. salar stocked in late summer exhibited survival at c. 5%. Stocking with unfed and fed fry contributed to increased smolt production and helped attain local management objectives between 2001 and 2005. Significant differences in biological characteristics were observed between wild and stocked‐origin fish. Wild‐smolt cohorts were dominated by 2+ year age‐class fish on the River Bush whilst smolts originating from fed fry mostly comprised younger 1+ year individuals. The mean mass of 1+ year smolts derived from stocked fed fry was significantly lower than that of wild 1+ year smolts, although these differences were not evident between older age classes. Differences in run timing between wild smolts and smolts derived from stocked fry were also apparent with the stocked‐origin fish tending to run earlier than wild fish. Although the stocking exercise was useful in terms of maximizing freshwater production, concerns over the quality of stocked‐origin recruits and the long term consequences for productivity are highlighted.     

Variability in Migration Routes Influences Early Marine Survival of Juvenile Salmon Smolts

Variability in animal migratory behavior is expected to influence fitness, but few empirical examples demonstrating this relationship exist. The initial marine phase in the migration of juvenile salmon smolts has been identified as a potentially critical life history stage to overall population productivity, yet how fine-scale migration routes may influence survival are unknown. Large-scale acoustic telemetry studies have estimated survival rates of outmigrant Pacific salmon smolts through the Strait of Georgia (SOG) along the British Columbian coastline to the Pacific Ocean, but these data have not been used to identify and characterize fine-scale movements. Data collected on over 850 sockeye salmon (Oncorhynchus nerka) and steelhead (Oncorhynchus mykiss) smolts detected at an array in the Strait of Georgia in 2004–2008 and 2010–2013 were analyzed to characterize migration routes and link movements to subsequent survival at an array 250 km further along the marine migration pathway. Both species exhibited disproportionate use of the most eastern route in the Strait of Georgia (Malaspina Strait). While many smolts moved across the northern Strait of Georgia acoustic array with no indication of long-term milling or large-scale east-to-west movements, large proportions (20–40% of sockeye and 30–50% of steelhead) exhibited a different behavior, apparently moving in a westward or counterclockwise pattern. Variability in migratory behavior for both species was linked to subsequent survival through the Strait of Georgia. Survival for both species was influenced by initial east-to-west location, and sockeye were further influenced by migration timing and duration of time spent near the northern Strait of Georgia array. Westward movements result in a net transport of smolts from Malaspina Strait to the Strait of Georgia, particularly for steelhead. Counterclockwise movements may be due to the currents in this area during the time of outmigration, and the higher proportion of steelhead smolts exhibiting this counterclockwise behavior may reflect a greater exposure to wind-altered currents for the more surface-oriented steelhead. Our results provide an empirical example of how movements can affect migration survival, for which examples remain rare in movement ecology, confirming that variability in movements themselves are an important part of the migratory process.     

Temperature seasonality during fry out‐migration influences the survival of hatchery‐reared chum salmon Oncorhynchus keta

Among years, fry‐to‐adult survival of hatchery‐reared chum salmon Oncorhynchus keta was positively correlated with the length (in days) of the fry out‐migration period with temperatures suitable for migration. Furthermore, survival decreased with increasing difference in mean temperature between May and June. Thus, prolonged out‐migration periods increased the probability of survival from fry to adult, lending support to the hypothesis that long migration periods decrease the risk of mortality (bet‐hedging), and increase the probability of migration when environmental conditions in fresh water and the ocean are suitable (match–mismatch).     

Swimming performance and morphology of juvenile sockeye salmon, <Emphasis Type="Italic">Oncorhynchus nerka</Emphasis>: comparison of inlet and outlet fry populations

We raised two populations of sockeye salmon fry from fertilized eggs in the laboratory and tested the hypothesis that outlet fry populations, fish which must migrate upstream to reach rearing lakes after yolk-sac absorption, have better swimming ability and morphological characteristics conducive to enhanced swimming performance than inlet fry populations, fish which migrate downstream to rearing lakes. Despite being of identical age, fry from the outlet population were larger (approx. 6.7% longer, ~5 mm on average) and more laterally compressed than inlet fry at the time of our initial experiments. Using an open-top box flume, we found that the burst-swimming performance (in cm s⁻¹) of the outlet population was 31% better. We found no differences between populations in prolonged-swimming performance. We were unable to find any direct relationships between measures of swimming performance and size or shape variables, suggesting that the larger, more robust morphology of outlet fry was not responsible for the superior burst ability. Recent biochemical studies indicate outlet fry may be metabolically better provisioned for burst swimming than inlet fry. It is possible that the morphological differences between the populations of fry reflect adaptations needed by adults during their migration and spawning.     

Carcass analogues provide marine subsidies for macroinvertebrates and juvenile Atlantic salmon in temperate oligotrophic streams

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Estimation of migratory history in female masu salmon Oncorhynchus masou (Brevoort, 1856) with the growth hormone pseudogene

To estimate the migration timing, length, and location of females with the growth hormone pseudogene (GHp), trace element analyses were performed on the otoliths of 40 wild adult masu salmon, Oncorhynchus masou (12 males, 19 normal females, and nine females with GHp [all fish were 3‐ or 4‐years‐old]) caught during the reproductive season (September) from 2004 to 2010 in the Atsuta River in Japan. Sr concentrations at the end of the otolith translucent area formed during the winter suddenly increased in all individuals, suggesting that they had migrated in the spring. In addition, the estimated migration length and location of females with GHp indicated that they had spent a fixed period of 1 year in the ocean, suggesting that females with GHp have a similar migratory history as that of normal females.     

Environmental signature in the otolith elemental fingerprint of the tapertail anchovy, Coilia mystus, from the Changjiang estuary, China

The ontogenetic patterns of habitat use and the migratory history of the tapertail anchovy, Coilia mystus, collected in the Changjiang estuary around Chongming Island, China, were studied by examining the environmental signature in the otolith strontium (Sr) and calcium (Ca) fingerprints using electron probe microanalysis (EPMA). Our results suggest that the migration strategy of C. mystus is much more flexible than supposed in the literature to date. The spring spawning population of C. mystus from the studied area was found to consist of individuals with different migration histories. Although the tapertail anchovy seems to be an anadromous fish that spawns and hatches in a freshwater habitat, it can also use a freshwater environment in non‐spawning seasons. The otolith EPMA of the elemental fingerprint (Sr x‐ray maps and Sr : Ca ratios) is an environmental indicator that can be applied to the migratory ecology of other important diadromous species in China.