The effects of environmental factors on the migratory movement patterns of Sacramento River yearling late-fall run Chinook salmon (Oncorhynchus tshawytscha)

Understanding smolt migration dynamics is a critical step in the preservation and conservation of imperiled salmonids in California’s Sacramento River system. Late-fall run Chinook salmon yearling smolts were acoustically tagged and tracked during their outmigration through California’s Sacramento River and San Francisco Estuary during 2007–2009. Migration rates were 14.3 km·day^-1 (± 1.3 S.E.) to 23.5 km?day^-1 (± 3.6 S.E.), similar to rates published for other West Coast yearling Chinook salmon smolt emigrations. Region-specific movement rates were fastest through the upper river regions, and slowest in the Sacramento/San Joaquin River Delta. River travel times were recorded for smolts travelling through a series of ten monitor-delimited reaches. Using these, a smolt travel time model determined by two parameters (movement rate and rate of population spreading) was then used to determine the influence of different factors on the model’s fit, using model selection with Akaike’s Information Criterion. The model that allowed for both year and reach to be expressed additively for both travel time and population spreading rate estimates, while accounting for a “release” effect, was the best supported model. Finally, several models incorporated environmental data as a linear predictor of movement rates. The addition of the environmental variables, in order of importance, river width to depth ratio, river flow, water turbidity, river flow to mean river flow ratio, and water velocity all resulted in improved model fit. Water temperature did not improve model fit. These environmental associations are discussed and potential improvements on the travel time model are suggested.     

Holding behavior of Chinook salmon (Oncorhynchus tshawytscha) and steelhead (O. mykiss) smolts, as influenced by habitat features of levee banks, in the highly modified lower Sacramento River, California

Using acoustic telemetry methods on large numbers of tagged fish, we studied how the holding behavior of Chinook salmon and steelhead smolts could be related to habitat features and spatial and temporal variables on a highly altered section of the Sacramento River. We viewed downstream migration as a process in which fish transition between moving and holding states, and used a binomial and negative binomial Generalized Linear Model to analyze two aspects of holding: 1) probability of holding, and 2) holding time. For Chinook salmon, the probability of holding increased as wood size and fine substrates increased; holding time increased as overhead shade increased. For steelhead, holding behavior was only weakly related to habitat variables, in contrast to the strong relationships with spatial and temporal variables. For both species, the probability of holding increased when distance from the release location decreased and instream flows decreased. We found support for three main findings: 1) spatial and temporal factors have considerably greater influence on Chinook salmon and steelhead smolt holding behavior than nearshore habitat features; 2) holding behaviors of Chinook salmon smolts are influenced more strongly by habitat features than steelhead smolts; and 3) incorporation of habitat features such as large woody material and overhead shade should be considered when conducting nearshore bank rehabilitation projects to increase cover from predators and provide velocity refuge, improving holding habitat during downstream migration.     

Interannual variation of reach specific migratory success for Sacramento River hatchery yearling late-fall run Chinook salmon (Oncorhynchus tshawytscha) and steelhead trout (Oncorhynchus mykiss)

The release of hatchery reared salmonid smolts is a common management tool aimed at enhancing depleted wild stocks and maintaining fisheries throughout Northern California and the Pacific Northwest. In the Sacramento River watershed, smolts must migrate through the river, delta and estuary in order to successfully reach the Pacific Ocean. Migration success (success defined as apparent survival from one monitor location to another) may vary between species, year and habitat. We released 500 late-fall run Chinook salmon and 500 steelhead smolts in 2009 and 2010 in the Sacramento River (river kilometer 207). Each smolt was implanted with a coded ultrasonic tag, which was detected by an array of over 300 underwater receiver stations deployed throughout the system. Less than 25 % of fish migrated successfully to the Pacific Ocean in both years. We found that reach specific success was greater in the Delta in 2009 (>60 %) than in 2010 (<33 %), whereas this pattern was reversed in the Bay (<57 % in 2009, >75 % in 2010). Identifying the location, timing and causes of smolt mortality can lead to improved management of the resource.     

Rhythmic aspects of estuarine migration of hatchery-reared Atlantic salmon (Salmo salar) smolts

Seasonal, diel and tidal rhythmic activity of hatchery-reared Atlantic salmon ( Salmo salar ) smolts migrating through a large estuary was studied by ultrasonic tracking of 46 individuals during two seasons. Prior to 10 May each year most smolts were inactive and remained near shore in shallow water. After 10 May nearly all smolts moved away from the release point into swift water and made rapid seaward progress. Initiation of migration each year occurred when river and hatchery water temperatures rose above 9°C. Migration in the estuary was largely passive drift, and as a result there were distinct tidal rhythms of ground ('swimming') speed and net seaward travel. There were no diel rhythms in ground speed or net seaward travel; smolts drifted seaward on the tides during both day and night. Smolts may be slightly deeper during day than night.     

The effect of hatchery release strategy on marine migratory behaviour and apparent survival of Seymour River steelhead smolts (Oncorhynchus mykiss)

Early marine migratory behaviour and apparent survival of hatchery-reared Seymour River steelhead (Oncorhynchus mykiss) smolts was examined over a four year period (2006-2009) to assess the impact of various management strategies on improving early marine survival. Acoustically tagged smolts were released to measure their survival using estuary and coastal marine receivers forming components of the Pacific Ocean Shelf Tracking (POST) array. Early marine survival was statistically indistinguishable between releases of summer run and winter run steelhead races, night and day releases, and groups released 10 days apart. In 2009, the survival of summer run steelhead released into the river was again trialed against groups released directly into the ocean at a distance from the river mouth. Apparent survival was improved significantly for the ocean released groups. The health and physiological status of the various release groups were monitored in years 2007-2009, and results indicate that the fish were in good health, with no clinical signs of disease at the time of release. The possibility of a disease event contributing to early marine mortality was further examined in 2009 by vaccinating half of the released fish against common fish diseases (vibriosis, furunculosis). The results suggest that marine survival may be enhanced using this approach, although not to the extent observed when the smolts were transported away from the river mouth before release. In summary, direct experimental testing of different release strategies using the POST array to measure ocean survival accelerated the scientific process by allowing rapid collection of data which enabled the rejection of several existing theories and allowed tentative identification of several new alternative approaches that might improve early marine survival of Seymour River steelhead.     

Archival and acoustic tags reveal the post-spawning migrations, diving behavior, and thermal habitat of hatchery-origin Sacramento River steelhead kelts (Oncorhynchus mykiss)

Acoustic and geolocating archival tags were deployed on steelhead kelts to study their post-spawning migrations, diving behavior and thermal habitat. Fourteen reconditioned steelhead kelts were surgically implanted with LAT2510 archival and V7-2 L acoustic tags, and released from Coleman National Fish Hatchery. An array of acoustic receivers in the Sacramento River and Delta, and the San Francisco Bay estuary detected the downstream movement of the steelhead. Two steelhead kelts (J and M) with archival tags were recovered at the hatchery after 219 and 285 days at liberty respectively. Based on changes in geolocations, vertical movements, and water temperatures, the migrations of steelhead J and M were divided into five and six phases respectively. Steelhead J moved into coastal California waters while steelhead M remained in freshwater for the majority of its time at liberty. Large increases in temperature and opacity were recorded before and after the ocean phases, likely as the steelhead moved through the Sacramento Delta. Both steelhead kelts remained relatively close to the surface throughout their migration but there were diurnal differences in the vertical movements. In freshwater, the steelhead tended to be deeper during the day (e.g., steelhead J: 3.08?±?1.50 vs 1.65?±?1.15 m, day vs night) but in the ocean, the steelhead were typically deeper during the night (1.32?±?1.61 vs 5.63?±?6.11 m). Both steelhead kelts appeared to be less oceanic than a previous study in Scott Creek, a small coastal stream approximately 100 km south of the mouth of San Francisco Bay. Although this study has a small sample size, the results suggest that steelhead kelts from a large river and estuary system, like the Sacramento River, may be relatively less oceanic than steelhead kelts from small coastal streams.     

Migratory Patterns of Wild Chinook Salmon Oncorhynchus tshawytscha Returning to a Large,Free-Flowing River Basin

Upriver movements were determined for Chinook salmon Oncorhynchus tshawytscha returning to the Yukon River, a large, virtually pristine river basin. These returns have declined dramatically since the late 1990s, and information is needed to better manage the run and facilitate conservation efforts. A total of 2,860 fish were radio tagged during 2002–2004. Most (97.5%) of the fish tracked upriver to spawning areas displayed continual upriver movements and strong fidelity to the terminal tributaries entered. Movement rates were substantially slower for fish spawning in lower river tributaries (28–40 km d^-1) compared to upper basin stocks (52–62 km d^-1). Three distinct migratory patterns were observed, including a gradual decline, pronounced decline, and substantial increase in movement rate as the fish moved upriver. Stocks destined for the same region exhibited similar migratory patterns. Individual fish within a stock showed substantial variation, but tended to reflect the regional pattern. Differences between consistently faster and slower fish explained 74% of the within-stock variation, whereas relative shifts in sequential movement rates between “hares” (faster fish becoming slower) and “tortoises” (slow but steady fish) explained 22% of the variation. Pulses of fish moving upriver were not cohesive. Fish tagged over a 4-day period took 16 days to pass a site 872 km upriver. Movement rates were substantially faster and the percentage of atypical movements considerably less than reported in more southerly drainages, but may reflect the pristine conditions within the Yukon River, wild origins of the fish, and discrete run timing of the returns. Movement data can provide numerous insights into the status and management of salmon returns, particularly in large river drainages with widely scattered fisheries where management actions in the lower river potentially impact harvests and escapement farther upstream. However, the substantial variation exhibited among individual fish within a stock can complicate these efforts.     

Migration of Atlantic salmon smolts and post-smolts in the Rivière Saint-Jean, QC north shore from riverine to marine ecosystems

The characteristics of the Atlantic salmon (Salmo salar) smolt run of the Rivière Saint-Jean, Quebec, Canada, in 2009 and 2010 were determined using acoustic telemetry. Tagged smolts were tracked from freshwater release sites for 17 km, then through the estuary and for their entry into the Gulf of St. Lawrence. The smolt migration began in both years at water temperatures of 10 °C, but lasted twice as long in the cooler year. The smolts in 2009 crossed the river to ocean boundary faster than the smolts of 2010, despite being similar in size. Smolt speed over ground increased from the river to the estuary to the marine environment. Smolts migrated both during the day and the night, but most movements began near or just after sunset, with increasingly nocturnal movements in the ocean. Smolts crossed the estuary during ebbing tides and moved faster during the night than the day. The tide cycle also influenced estuarine smolt travelling rates, but only secondarily to the diel cycle.     

Thermal biology and bioenergetics of different upriver migration strategies in a stock of summer-run Chinook salmon

By combining biotelemetry with animal-borne thermal loggers, we re-created the thermal histories of 21 summer-run Chinook salmon (Oncorhynchus tshawytscha) migrating in the Puntledge River, a hydropower impacted river system on Vancouver Island, British Columbia, Canada. Daily maximum water temperatures in the Puntledge River during the summer-run adult Chinook salmon migration and residency period frequently exceeded 21 °C, a value that has been observed to elicit behavioral thermoregulation in other Chinook salmon populations. We therefore compared river temperatures to body temperatures of 16 fish that migrated through the river to understand if cool-water refuge was available and being used by migrants. In addition, we used thermal histories from fish and thermal loggers distributed in the river to model the effect of thermal habitat on energy density using a bioenergetics model. In general, we found no evidence that cool-water refuge existed in the river, suggesting that there is no opportunity for fish to behaviorally thermoregulate during upriver migration through the regulated portion of the river. Of the thermal histories used in the bioenergetics model, fish that reached an upstream lake were able to access cooler, deeper waters, which would have reduced energy consumption compared to fish that only spent time in the warmer river. Consequently, the Puntledge River water temperatures are likely approaching and in some cases exceeding the thermal limits of the summer-run Chinook salmon during the spawning migration. Further warming may cause more declines in the stock.     

Riverine, estuarine and marine migratory behaviour and physiology of wild and hatchery-reared coho salmon Oncorhynchus kisutch (Walbaum) smolts descending the Campbell River, BC, Canada

Eighty coho salmon Oncorhynchus kisutch smolts (40 wild and 40 hatchery-reared) were surgically implanted with acoustic transmitters and released into the Quinsam River over 2 days. Differences in physiology, travel time and migratory behaviour were examined between wild and hatchery-reared fish. In addition, tagged and control fish of both wild and hatchery-reared stock were raised for 3 months following surgery to compare survival and tag retention. Detection ranges of the acoustic receivers were tested in the river, estuary and ocean in a variety of flow conditions and tide levels. Receivers were placed in the river, estuary and up to 50 km north and south from the river mouth in the marine environment. Wild smolts were significantly smaller by mass, fork length and condition factor than hatchery-reared smolts and exhibited significantly higher levels of sodium, potassium and chloride in their blood plasma than hatchery-reared smolts. The gill Na⁺K⁺-ATPase activity was also significantly higher in the wild coho smolts at the time of release. Ninety-eight per cent of wild and 80% of hatchery-reared fish survived to the estuary, 8 km downstream of the release site. No difference was found in migration speed, timing or survival between smolts released during daylight and those released after dark. Wild smolts, however, spent less time in the river and estuary, and as a result entered the ocean earlier than hatchery-reared smolts. Average marine swimming speeds for wild smolts were double those of their hatchery-reared counterparts. While hatchery smolts dispersed in both a northward and southward direction upon entering the marine environment, the majority of wild smolts travelled north from the Campbell River estuary. The wild coho salmon smolts were more physiologically fit and ready to enter sea water than the hatchery-reared smolts, and as a result had higher early survival rates and swimming speeds.     

Migration of hatchery reared juvenile Atlantic salmon, Salmo salar L., smolts down a release ladder. 1. Environmental effects on migratory activity

The effects of light intensity, water temperature and river spate conditions on the rate of migration of hatchery-reared Atlantic salmon smolts down a release ladder were examined. Low light intensity and high day time water temperatures raised smolt migration rates: water temperature had littleor no effect at night. Thediel patternofdown-ladder movement wasdetermined by these two environmental stimuli. Smolts demonstrated a threshold response to both light and water temperature: a fall in light intensity, or increase in water temperature, below or above their respective thresholds elicited no further response. As fish progressed through the ladder their movement became increasingly nocturnal, and most entry into the estuary occurred at night. Heavy rainhll caused rapid migration of smolts through the ladder. As ladder discharge remained constant, smolts must have been responding to some other change in the water conditions. Smolts responded to spate conditions irrespective of water temperature and light intensity, and they responded to light intensity irrespective of water temperature. A hierarchy of environmental cues, responsible for triggering migratory behaviour on a day to day basis during the smolt run, of spate> light intensity > water temperature is therefore suggested.     

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.     

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.     

Characteristics of the spawning stock of chinook salmon <Emphasis Type="Italic">Oncorhynchus tshawytscha</Emphasis> from the Apuka River (northeastern Kamchatka)

Biological features of chinook salmon Oncorhynchus tshawytscha from the Apuka River, the largest river of the northeastern Kamchatka inflowing Olyutorskii Bay of the Bering Sea, are studied. Chinook salmon from the Apuka River spend mainly a year in the river before downstream migration to the sea. The fish live in the sea for 1–4 years. The spawning migration of chinook salmon into the Apuka River begins in late May just after ice melting, and it continues until early August. The main part of the spawners enters the river during June. A hypothesis on the occurrence of two seasonal races in the Apuka River is proposed.     

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.     

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.     

Growth, survival, and tag retention of steelhead trout (Oncorhynchus mykiss) and its application to survival estimates

Steelhead trout, Oncorhynchus mykiss, are known to expel acoustic tags which can negatively bias population survival estimates. Tag burden may also affect the development and behavior of smolts, thereby impacting the results of population and behavioral studies. We monitored the growth, condition, and tag expulsion rate of steelhead trout in similar-sized individuals and used these data to adjust survival rates from an acoustic telemetry study conducted in the Sacramento River. Eighty fish were surgically implanted with tags—40 with cylindrical tags of 9 mm diameter and 21 or 24 mm length (V9, Vemco Ltd) and 40 with a 7 mm diameter and 20 mm length tag (V7, Vemco Ltd)—to examine the impact of tag size on peritoneal retention and survival rate of juvenile steelhead trout. A total of 20 % (16/80) of all tags were expelled by smolts during the 143-day study. Ten V9 tags were expelled between day 18 and day 66. Six V7 tags were expelled between day 21 and day 143. A statistical difference was found for retention rate by surgeon even though the surgeons were of equal experience and received the same training. There were no significant differences in the tag retention rate in relation to the tag/body weight ratio, or in growth (weight or fork length) among the control, V7 or V9 treatment groups over the duration of the study. All individuals survived throughout the experiment. Two methods were used to adjust the survival estimates of an acoustic telemetry data set from the Sacramento River based on the tag retention study. First, a simple individual censorship approach in Program MARK was utilized and next ATLAS, a software program designed to compensate for bias in survival estimates caused by tag failures was used. The results of the adjusted survival estimates were not significantly different from the unadjusted rates suggesting that it may be more important to focus on improving surgical techniques to reduce tag expulsion rather than adjusting survival estimates dependent on the study. The surgical techniques utilized in this study did not have significant impacts on the growth rates of either of the tag treatment groups compared to the control. However, tag retention was an issue regardless of the size and weight of the implanted tag and the size of the steelhead.     

Comprehensive passage (COMPASS) model: a model of downstream migration and survival of juvenile salmonids through a hydropower system

Migratory fish populations are impacted worldwide by river impoundments. Efforts to restore populations will benefit from a clear understanding of survival and migration process over a wide-range of river conditions. We developed a model that estimates travel time and survival of migrating juvenile salmonids (Oncorhynchus spp.) through the impounded Snake and Columbia rivers in the northwestern United States. The model allows users to examine the effects of river management scenarios, such as manipulations of river flow and spill, on salmonid survival. It has four major components: dam passage and survival, reservoir survival, fish travel time, and hydrological processes. The probability that fish pass through specific routes at a dam and route-specific survival probabilities were based on hydroacoustic, radio telemetry, PIT tag, and acoustic tag data. We related reservoir mortality rate (per day and per km) to river flow, water temperature, and percentage of fish passing through spillways and then fit the relationships to PIT-tag survival data. We related fish migration rate to water velocity, percentage of fish passing through spillways, and date in the season. We applied the model to two threatened “Evolutionarily Significant Units” (as defined under the US Endangered Species Act): Snake River spring/summer Chinook salmon (O. tshawytscha Walbaum) and Snake River steelhead (O. mykiss Walbaum). A sensitivity analysis demonstrated that for both species survival through the hydropower system was responsive to water temperature, river flow, and spill proportion. The two species, however, exhibited different patterns in their response. Such information is crucial for managers to effectively restore migratory fish populations in regulated rivers. Guest editors: R. L. Welcomme & G. Marmulla Hydropower, Flood Control and Water Abstraction: Implications for Fish and Fisheries     

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.