Patterns of gametic investment by female stream-and ocean-type chinook salmon

Investment in fecundity and egg size is compared among two types of chinook salmon Oncorhynchus tshawytscha : stream-type, which undertake long, arduous oceanic and upstream migrations and often return to their natal rivers (where they do not feed)months before spawning, and ocean-type, which undertake relatively short oceanic and upstream migrations and remain at sea feeding and accumulating energy until a few days or weeks before spawning. The coefficient of variation in egg size for both life-history types was significantly less than variation in fecundity, reflecting the expected strong selection on egg size relative to egg number. Total investment in the gonad varied significantly among years in both life-history types, apparently reflecting variation in oceanic feeding conditions. Stream-type chinook were smaller in total body size and more than 1 year older than ocean-type. Stream-type also invested relatively less of their total body energy in eggs and produced smaller eggs than ocean-type, reflecting the greater energetic cost of their longer migration and freshwater maintenance. These differences suggest that stream-type chinook pay a considerable fitness penalty in reproductive output. This may be offset by the fact that stream-type go to sea at larger size and migrate far offshore, thereby avoiding some of the mortality costs of oceanic migration.     

Divergence in juvenile growth and life history in two recently colonized and partially isolated chinook salmon populations

In 47 families of New Zealand chinook salmon, Oncorhynchus tshawytscha derived from two populations, differing in mean spawning date by 11–17 days and in juvenile life history, neither mean ovum weight nor mean time to hatch differed between the two populations, but substantial differences in mean family weight were apparent from 6 months after fertilization. Differences in growth rates from 12 to 24 months were relatively small, suggesting that most of the divergence in size and growth occurred during the first 6 months of life. There were differences in mean weight for families spawned 11–17 days apart, the approximate interval by which peak spawning date differs between the two populations, but these did not persist beyond 7 months. Differences between the two populations are consistent with their natural life history. The slower growing population experiences cooler temperatures during stream residence and is dominated by fish which spend a year in fresh water before seaward migration, whereas the faster growing population normally experiences warmer temperatures and migrates to sea in the first year of life. These results provide further evidence of life history evolution in partially isolated chinook salmon populations within 90 years ( c. 30 generations) of becoming established.     

Evolution of temporal isolation in the wild: genetic divergence in timing of migration and breeding by introduced chinook salmon populations

Abstract. The timing of migration and breeding are key life-history traits; they are not only adaptations of populations to their environments, but can serve to increase reproductive isolation, facilitating further divergence among populations. As part of a study of divergence of chinook salmon, Oncorhynchus tshawytscha , populations, established in New Zealand from a common source in the early 1900s, we tested the hypotheses that the timing of migration and breeding are under genetic control and that the populations genetically differ in these traits despite phenotypic overlap in timing in the wild. Representatives of families from two populations were collected within a day or two of each other, reared in a common environment, and then released to sea from each of two different rivers, while other family representatives were retained in fresh water to maturity. The date of maturation of fish held in fresh water and the dates of return from the ocean and maturation of fish released to sea all showed significant differences between the two populations and among families within populations. The very high heritabilities and genetic correlations estimated for migration and maturation date indicated that these traits would respond rapidly to selection. Combined with the results of related studies on these chinook salmon populations, it appears that spawning time may not only evolve during the initial phases of divergence, but it may play an important role in accelerating divergence in other traits.     

Sea lice and salmon population dynamics: effects of exposure time for migratory fish

The ecological impact of parasite transmission from fish farms is probably mediated by the migration of wild fishes, which determines the period of exposure to parasites. For Pacific salmon and the parasitic sea louse, Lepeophtheirus salmonis, analysis of the exposure period may resolve conflicting observations of epizootic mortality in field studies and parasite rejection in experiments. This is because exposure periods can differ by 2–3 orders of magnitude, ranging from months in the field to hours in experiments. We developed a mathematical model of salmon–louse population dynamics, parametrized by a study that monitored naturally infected juvenile salmon held in ocean enclosures. Analysis of replicated trials indicates that lice suffer high mortality, particularly during pre-adult stages. The model suggests louse populations rapidly decline following brief exposure of juvenile salmon, similar to laboratory study designs and data. However, when the exposure period lasts for several weeks, as occurs when juvenile salmon migrate past salmon farms, the model predicts that lice accumulate to abundances that can elevate salmon mortality and depress salmon populations. The duration of parasite exposure is probably critical to salmon–louse population dynamics, and should therefore be accommodated in coastal planning and management where fish farms are situated on wild fish migration routes.     

The effects of turbidity and vegetation on the risk of juvenile salmonids, Oncorhynchus spp., to predation by adult cutthroat trout, O. clarkii

Synopsis During their seaward migration, juvenile salmonids encounter structural and visual cover which varies between and within watersheds. In this study, the effects of two types of cover (turbidity and artificial vegetation) on the predation mortality of juvenile salmonids exposed to fish piscivores was investigated in outdoor concrete ponds. During experiments, adult coastal cutthroat trout, Oncorhynchus clarkii clarkii, were allowed to feed on juvenile salmonid prey — chinook salmon, O. tshawytscha, chum salmon, O. keta, sockeye salmon, O. nerka, and cutthroat trout — in separate trials. Daily instantaneous per capita predation rate was determined for each turbidity and vegetation treatment, within each trial. Mean predation rates varied between 1% and 76% daily. In the presence of cover, mean daily predation rates were 10–75% lower than those in controls (no vegetation and clear water), depending on prey species. Predation rates were significantly lower in the presence of vegetation cover and did not covary with prey size or species. The effects of turbidity were generally not significant and were not additive with the effects of vegetation. However, turbidity appeared to significantly reduce the effectiveness of vegetation as cover for juvenile chinook and sockeye salmon. We suggest that these two forms of cover do not affect risk of predation by fish piscivores to juvenile salmonids via the same mechanism.     

Sea louse infection of juvenile sockeye salmon in relation to marine salmon farms on Canada's west coast

Background

Pathogens are growing threats to wildlife. The rapid growth of marine salmon farms over the past two decades has increased host abundance for pathogenic sea lice in coastal waters, and wild juvenile salmon swimming past farms are frequently infected with lice. Here we report the first investigation of the potential role of salmon farms in transmitting sea lice to juvenile sockeye salmon (Oncorhynchus nerka).

Methodology/Principal Findings

We used genetic analyses to determine the origin of sockeye from Canada''s two most important salmon rivers, the Fraser and Skeena; Fraser sockeye migrate through a region with salmon farms, and Skeena sockeye do not. We compared lice levels between Fraser and Skeena juvenile sockeye, and within the salmon farm region we compared lice levels on wild fish either before or after migration past farms. We matched the latter data on wild juveniles with sea lice data concurrently gathered on farms. Fraser River sockeye migrating through a region with salmon farms hosted an order of magnitude more sea lice than Skeena River populations, where there are no farms. Lice abundances on juvenile sockeye in the salmon farm region were substantially higher downstream of farms than upstream of farms for the two common species of lice: Caligus clemensi and Lepeophtheirus salmonis, and changes in their proportions between two years matched changes on the fish farms. Mixed-effects models show that position relative to salmon farms best explained C. clemensi abundance on sockeye, while migration year combined with position relative to salmon farms and temperature was one of two top models to explain L. salmonis abundance.

Conclusions/Significance

This is the first study to demonstrate a potential role of salmon farms in sea lice transmission to juvenile sockeye salmon during their critical early marine migration. Moreover, it demonstrates a major migration corridor past farms for sockeye that originated in the Fraser River, a complex of populations that are the subject of conservation concern.     

Transmission dynamics of parasitic sea lice from farm to wild salmon

Marine salmon farming has been correlated with parasitic sea lice infestations and concurrent declines of wild salmonids. Here, we report a quantitative analysis of how a single salmon farm altered the natural transmission dynamics of sea lice to juvenile Pacific salmon. We studied infections of sea lice (Lepeophtheirus salmonis and Caligus clemensi) on juvenile pink salmon (Oncorhynchus gorbuscha) and chum salmon (Oncorhynchus keta) as they passed an isolated salmon farm during their seaward migration down two long and narrow corridors. Our calculations suggest the infection pressure imposed by the farm was four orders of magnitude greater than ambient levels, resulting in a maximum infection pressure near the farm that was 73 times greater than ambient levels and exceeded ambient levels for 30 km along the two wild salmon migration corridors. The farm-produced cohort of lice parasitizing the wild juvenile hosts reached reproductive maturity and produced a second generation of lice that re-infected the juvenile salmon. This raises the infection pressure from the farm by an additional order of magnitude, with a composite infection pressure that exceeds ambient levels for 75 km of the two migration routes. Amplified sea lice infestations due to salmon farms are a potential limiting factor to wild salmonid conservation.     

Alternative developmental pathways and the propensity to migrate: a case study in the Atlantic salmon

Migratory behaviour with its associated phenotypic changes is generally viewed as an adaptive strategy because it incurs survival or reproductive advantages to migrants. The development of a migrant phenotype is believed to be controlled by threshold mechanisms, where individuals emigrate only after surpassing a particular body size but delay migration if below. For such a strategy to respond to natural selection, part of the phenotypic variance in the propensity to migrate must be explained by variation in additive genetic effects. Here, we use data gathered in the field and from a common rearing experiment to test for a genetic basis associated with seaward migration in Atlantic salmon (Salmo salar L.). We document a high heritability of the liability trait underlying the propensity to emigrate in juvenile salmon, and significant differences between offspring grouped according to their sires in body-size threshold values above which emigration takes place. The presence of additive genetic variance in both the liability and thresholds makes the onset of migration a process sensitive to selection and may therefore constitute an important explanatory mechanism for the interpopulation differences in the size at seaward migration observed in this species.     

Quantifying the ocean, freshwater and human effects on year-to-year variability of one-sea-winter Atlantic salmon angled in multiple Norwegian rivers

Many Atlantic salmon, Salmo salar, populations are decreasing throughout the species' distributional range probably due to several factors acting in concert. A number of studies have documented the influence of freshwater and ocean conditions, climate variability and human impacts resulting from impoundment and aquaculture. However, most previous research has focused on analyzing single or only a few populations, and quantified isolated effects rather than handling multiple factors in conjunction. By using a multi-river mixed-effects model we estimated the effects of oceanic and river conditions, as well as human impacts, on year-to-year and between-river variability across 60 time series of recreational catch of one-sea-winter salmon (grilse) from Norwegian rivers over 29 years (1979-2007). Warm coastal temperatures at the time of smolt entrance into the sea and increased water discharge during upstream migration of mature fish were associated with higher rod catches of grilse. When hydropower stations were present in the course of the river systems the strength of the relationship with runoff was reduced. Catches of grilse in the river increased significantly following the reduction of the harvesting of this life-stage at sea. However, an average decreasing temporal trend was still detected and appeared to be stronger in the presence of salmon farms on the migration route of smolts in coastal/fjord areas. These results suggest that both ocean and freshwater conditions in conjunction with various human impacts contribute to shape interannual fluctuations and between-river variability of wild Atlantic salmon in Norwegian rivers. Current global change altering coastal temperature and water flow patterns might have implications for future grilse catches, moreover, positioning of aquaculture facilities as well as the implementation of hydropower schemes or other encroachments should be made with care when implementing management actions and searching for solutions to conserve this species.     

Ecological Risk Assessment Paradigm for Salmon: Analyzing Immune Function to Evaluate Risk

Wild Pacific salmon populations are in serious decline, and as a result, a number of salmon stocks are listed as threatened or endangered under the Endangered Species Act. Our research identifies and supports the possibility that certain environmental contaminants can alter salmon survival, and as a result may contribute to these species being at risk. We have shown that juvenile chinook salmon (Oncorhynchus tshawytscha) are exposed to polychlorinated biphenyls (PCBs) and polycyclic aromatic hydrocarbons (PAHs) as they migrate through a contaminated urban estuary in Puget Sound WA (the Duwamish Waterway estuary). Immune function was analyzed in these fish by examining the ability of their anterior kidney and splenic leukocytes to produce a primary and secondary in vitro plaque-forming cell (PFC) response to the hapten, trinitrophenyl (TNP), and by determining their susceptibility to a marine pathogen, Vibrio anguillarum. We found that fish outmigrating from the urban estuary produced a significantly lower PFC response to TNP and were more susceptible to the pathogen, compared to juvenile salmon collected from a rural estuary during their outmigration. In the laboratory, we exposed juvenile chinook salmon collected from a hatchery to either a PCB technical mixture or a PAH compound to determine if these contaminants have the potential to alter immune function in salmon. Indeed, we found that salmon exposed in the laboratory to either the PCB mixture or the PAH also produced lower PFC responses and were more susceptible to disease compared to animals treated with the solvent vehicle. In summary, contaminants such as PAHs and PCBs are demonstrated to influence salmon health, and thus have the potential to adversely impact salmon populations.     

Density‐dependent mortality in Pacific salmon: the ghost of impacts past?

Conservation biologists often ignore density dependence because at‐risk populations are typically small relative to historical levels. However, if populations are reduced as a result of impacts that lower carrying capacity, then density‐dependent mortality may exist at low population abundances. Here, we explore this issue in threatened populations of juvenile chinook salmon (Oncorhynchus tshawytscha). We followed the fate of more than 50 000 juvenile chinook in the Snake River Basin, USA to test the hypothesis that their survival was inversely associated with juvenile density. We also tested the hypotheses that non‐indigenous brook trout and habitat quality affect the presence or strength of density dependence. Our results indicate that juvenile chinook suffer density‐dependent mortality and the strength of density dependence was greater in streams in which brook trout were absent. We were unable to detect an effect of habitat quality on the strength of density dependence. Historical impacts of humans have greatly reduced population sizes of salmon, and the density dependence we report may stem from a shortage of nutrients normally derived from decomposing salmon carcasses. Cohorts of juvenile salmon may experience density‐dependent mortality at population sizes far below historical levels and recovery of imperiled populations may be much slower than currently expected.     

Using "travel time" data to characterize the behavior of migrating animals

For migratory species, duration of migration, or "travel time," is often a critical variable in determining the cost of migration. Observed travel times are the result of both environmental factors such as air or water currents and the behavior of individuals. In an effort to distinguish among these components, I developed a migration model based on an advection-diffusion equation that characterizes population movements in terms of two biologically meaningful parameters: migration rate and rate of population spread. I applied the model to travel time data from juvenile chinook salmon (Onchorhynchus tshawytscha), which were tagged during their seaward migration. The tagged fish originated from three separate evolutionarily significant units (ESUs) as classified by the U. S. National Marine Fisheries Service. The model was expanded by allowing migration and diffusion rates to vary with fish length and river flow. Variability in travel times explained by these factors was strikingly similar from year to year within ESUs, and the migratory behavior revealed by the analysis was consistent with the life-history patterns that distinguish the ESUs. The approach presented here is easily adaptable to a wide range of migratory species and may be particularly useful for predicting how at-risk populations respond to variable conditions in regulated or otherwise disturbed migration habitats.     

Fecundity and egg size variation in North American Pacific salmon (Oncorhynchus)

We examined regional and latitudinal variation in fecundity and egg weight for five species of Pacific salmon ( Oncorhynchus ) along the Pacific coast of North America. Data were examined for 24 chum salmon, 15 pink salmon, 34 sockeye salmon, 44 chinook salmon, and 40 coho salmon populations from published sources, unpublished Canadian hatchery records, our own laboratory investigations, and other unpublished sources. Substantial regional variation in fecundity and egg weight was observed, with salmon on the Queen Charlotte Islands and Vancouver Island in British Columbia generally having lower fecundity and larger egg size than nearby mainland populations. The relative distance of freshwater migration to the spawning grounds generally had a marked effect on both fecundity and egg size, with populations spawning in the upper portions in the drainages of large rivers like the Fraser River in British Columbia having reduced fecundity and egg size compared with coastal spawning populations. Fecundity was generally higher and egg size generally lower in more northern populations of sockeye, chinook, and coho salmon compared with southern ones. We suggest that egg size tends to be lower in northern populations of some species as a result of increased fecundity due to their older ages at maturity and a limited amount of energy that can be expended on egg production.     

Non-indigenous brook trout and the demise of Pacific salmon: a forgotten threat?

Non-indigenous species may be the most severe environmental threat the world now faces. Fishes, in particular, have been intentionally introduced worldwide and have commonly caused the local extinction of native fish. Despite their importance, the impact of introduced fishes on threatened populations of Pacific salmon has never been systemically examined. Here, we take advantage of several unique datasets from the Columbia River Basin to address the impact of non-indigenous brook trout, Salvelinus fontinalis, on threatened spring/summer-run chinook salmon, Oncorhynchus tshawytscha. More than 41 000 juvenile chinook were individually marked, and their survival in streams without brook trout was nearly double the survival in streams with brook trout. Furthermore, when brook trout were absent, habitat quality was positively associated with chinook survival, but when brook trout were present no relationship between chinook survival and habitat quality was evident. The difference in juvenile chinook survival between sites with, and without, brook trout would increase population growth rate (lambda) by ca. 2.5%. This increase in lambda would be sufficient to reverse the negative population growth observed in many chinook populations. Because many of the populations we investigated occur in wilderness areas, their habitat has been considered pristine; however, our results emphasize that non-indigenous species are present and may have a dramatic impact, even in remote regions that otherwise appear pristine.     

Physiological consequences of the salmon louse (Lepeophtheirus salmonis) on juvenile pink salmon (Oncorhynchus gorbuscha): implications for wild salmon ecology and management, and for salmon aquaculture

Pink salmon, Oncorhynchus gorbuscha, are the most abundant wild salmon species and are thought of as an indicator of ecosystem health. The salmon louse, Lepeophtheirus salmonis, is endemic to pink salmon habitat but these ectoparasites have been implicated in reducing local pink salmon populations in the Broughton Archipelago, British Columbia. This allegation arose largely because juvenile pink salmon migrate past commercial open net salmon farms, which are known to incubate the salmon louse. Juvenile pink salmon are thought to be especially sensitive to this ectoparasite because they enter the sea at such a small size (approx. 0.2 g). Here, we describe how 'no effect' thresholds for salmon louse sublethal impacts on juvenile pink salmon were determined using physiological principles. These data were accepted by environmental managers and are being used to minimize the impact of salmon aquaculture on wild pink salmon populations.     

In situ and laboratory studies on the behaviour and survival of Pacific salmon (genus Oncorhynchus)

Juvenile Pacific salmon display a marked surface water orientation during downstream migration, estuarine and nearshore coastal rearing phases. Many estuaries in British Columbia are vertically stratified with a shallow, well-defined halocline which can restrict the dispersion of wastes discharged into less saline surface waters and impose constraints upon aquatic organisms. In situ experiments in an estuary receiving a surface discharge of treated pulp mill wastes, revealed conditions which were lethal to underyearling salmon at, and below the halocline (4.0–6.5 m depth). Behavioural bioassays determined that juvenile chinook salmon were biased towards the water surface and avoided waters at depth. Dissolved oxygen was the variable which affected this distribution most significantly. Surface waters receiving effluent from another pulp mill were lethal to juvenile salmon within 350 m, and a significant vertical avoidance response occurred within 350–950 m of the outfalls. The behavioural response was significantly correlated with in situ temperature, pH and colour (effluent).As a complement to field experiments we developed a 4500 l water column simulator (WCS) to examine salmon behaviour in the laboratory. We investigated the surface water orientation behaviour of juvenile salmon in relation to variations in salinity and dissolved oxygen. Under simulated vertically stratified estuarine conditions, the fish moved freely between overlying fresh water and salt water. Induction of hypoxic conditions in fresh water elicited a downward distribution shift towards the halocline and oxygenated, but more saline, waters. Avoidance reactions (50% level) occurred consistently up to 7–8 mg · l^–1 dissolved oxygen. Salmon continued to examine the hypoxic freshwater zone despite sub-optimal conditions.     

New tetranucleotide microsatellites for fine‐scale discrimination among endangered chinook salmon (Oncorhynchus tshawytscha)

The unambiguous identification of Central Valley spring‐run chinook salmon has become imperative since their proposed listing in 1998. The accuracy of methods used to assign individuals to their stock of origin is critical for understanding juvenile migration patterns and determining the success of protection measures. Existing microsatellites discriminate between the endangered winter‐run and other chinook but are insufficient to characterize phylogenetically less distinct runs. Here, we isolated and developed highly variable tetranucleotide microsatellites for the specific goal of increasing discriminatory power among closely related populations, providing a new power towards the reliable differentiation of nonwinter runs     

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.     

Migration patterns and navigation cues of Atlantic salmon post-smolts migrating from 12 rivers through the coastal zones around the Irish Sea

The freshwater phase of the first seaward migration of juvenile Atlantic salmon (Salmo salar) is relatively well understood when compared with our understanding of the marine phase of their migration. In 2021, 1008 wild and 60 ranched Atlantic salmon smolts were tagged with acoustic transmitters in 12 rivers in England, Scotland, Northern Ireland and Ireland. Large marine receiver arrays were deployed in the Irish Sea at two locations: at the transition of the Irish Sea into the North Atlantic between Ireland and Scotland, and between southern Scotland and Northern Ireland, to examine the early phase of the marine migration of Atlantic salmon smolts. After leaving their natal rivers' post-smolt migration through the Irish Sea was rapid with minimum speeds ranging from 14.03 to 38.56 km.day⁻¹ for Atlantic salmon smolts that entered the Irish Sea directly from their natal river, to 9.69–39.94 km.day⁻¹ for Atlantic salmon smolts that entered the Irish Sea directly from their natal estuary. Population minimum migration success through the study area was strongly correlated with the distance of travel, populations further away from the point of entry to the open North Atlantic exhibited lower migration success. Post-smolts from different populations experienced different water temperatures on entering the North Atlantic. This was largely driven by the timing of their migration and may have significant consequences for feeding and ultimately survivorship. The influence of water currents on post-smolt movement was investigated using data from previously constructed numerical hydrodynamic models. Modeled water current data in the northern Irish Sea showed that post-smolts had a strong preference for migrating when the current direction was at around 283° (west-north-west) but did not migrate when exposed to strong currents in other directions. This is the most favorable direction for onward passage from the Irish Sea to the continental shelf edge current, a known accumulation point for migrating post-smolts. These results strongly indicate that post-smolts migrating through the coastal marine environment are: (1) not simply migrating by current following (2) engage in active directional swimming (3) have an intrinsic sense of their migration direction and (4) can use cues other than water current direction to orientate during this part of their migration.     

Effects of host migration, diversity and aquaculture on sea lice threats to Pacific salmon populations

Animal migrations can affect disease dynamics. One consequence of migration common to marine fish and invertebrates is migratory allopatry-a period of spatial separation between adult and juvenile hosts, which is caused by host migration and which prevents parasite transmission from adult to juvenile hosts. We studied this characteristic for sea lice (Lepeophtheirus salmonis and Caligus clemensi) and pink salmon (Oncorhynchus gorbuscha) from one of the Canada's largest salmon stocks. Migratory allopatry protects juvenile salmon from L. salmonis for two to three months of early marine life (2-3% prevalence). In contrast, host diversity facilitates access for C. clemensi to juvenile salmon (8-20% prevalence) but infections appear ephemeral. Aquaculture can augment host abundance and diversity and increase parasite exposure of wild juvenile fish. An empirically parametrized model shows high sensitivity of salmon populations to increased L. salmonis exposure, predicting population collapse at one to five motile L. salmonis per juvenile pink salmon. These results characterize parasite threats of salmon aquaculture to wild salmon populations and show how host migration and diversity are important factors affecting parasite transmission in the oceans.