The food web positioning and trophic niche of the non-indigenous round goby: a comparison between two Baltic Sea populations

Native species may show invasiveness toward a recipient ecosystem through increases in abundance as a result of artificial stocking events. Salmonid species are typical examples of native invaders whose abundance is increased after stocking with hatchery fish. This study evaluated the effects of hatchery chum salmon fry on sympatric wild masu salmon fry, benthic invertebrate prey, and algae, after a single stocking event in Mamachi stream, Hokkaido, northern Japan. The results suggested that the stocked hatchery chum salmon fry decreased the foraging efficiency and growth of the wild masu salmon fry through interspecific competition, and depressed the abundance of Ephemerellidae and total grazer invertebrates (Glossosomatidae, Heptageniidae, and Baetidae) through predation. Also, the hatchery chum salmon fry may increase algal biomass through depression of grazer abundance by predation (top-down effect). These results suggested that a single release of hatchery chum salmon fry into a stream may influence the recipient stream ecosystem.     

Intra- and Interspecific Competition in Hatchery Landlocked Salmon and Brown Trout in Semi-Natural Streams

I studied inter- and intraspecific competition in two hatchery stocks: landlocked salmon with long-hatchery background and a heterogenic brown trout stock. These species are potential competitors in the natural environment when landlocked salmon is being restored to wild by stocking hatchery juveniles. Behavioural responses were studied in four indoor laboratory flumes (400 cm long and 37 cm wide) and habitat use in six semi-natural outdoor streams (26 m long and 1.5 m long). Video recordings were used to monitor fish behaviour and electrofishing for fish positioning in the outdoor channels. The study design included five treatments: two densities of brown trout and salmon in solitary and both species together. The results of the study demonstrated that juvenile brown trout changed their behaviour in laboratory streams in response to presence of the landlocked salmon and the density of the conspecifics also tended to alter the habitat use by brown trout in semi-natural streams. Landlocked salmon juveniles showed no response to treatments. I conclude that possible poor adaptive ability to conditions outside hatchery by the hatchery salmon together and more competitive brown trout stocks may limit the success of management action in restoring landlocked salmon back to their natural streams of stocking.     

Hierarchical genetic structure of native masu salmon populations in Hokkaido,Japan

Identification of the spatial extent of genetic structuring that may be influenced by evolutionary, ecological and historical factors is critical for effective conservation or management strategies. Masu salmon Oncorhynchus masou is commonly distributed in Far East, however, many local populations have been under threats of decline due to habitat destruction, overexploitation, and genetic introgression. To reveal the spatial genetic structure of native masu salmon populations in Hokkaido, masu salmon samples were collected from 16 rivers in which there was no official record of artificial releases of any masu salmon stock and were analyzed using 15 microsatellite loci. A Bayesian assignment test revealed that masu salmon populations were divided into two genetically distinct groups: the northeastern and southwestern groups. For within-group genetic structure, all populations, except for geographically proximate populations, were significantly different from each other. AMOVA revealed that genetic variation at among-group level based on groups identified assignment test was greater than that of groups based on geographic locations. There was no significant IBD for the 16 populations. However, the Mantel test revealed significant IBD for the northeastern group, but did not for the southwestern group. This study suggested that native masu salmon populations in Hokkaido exhibit a hierarchical genetic structure that is largely a result of their precise homing behavior. The results of this study also highlight the importance of defining populations by using genetic data rather than by using predefined populations based on geographic locations for the correct determination of genetic structure.     

Comparison of competitive ability between native and introduced salmonids: evidence from pairwise contests

Brown trout, Salmo trutta, and rainbow trout, Oncorhynchus mykiss, have been introduced to freshwaters in Hokkaido, Japan. Today, it is recognized that these introduced salmonids have negative impacts on native salmonids such as white-spotted charr, Salvelinus leucomaenis, and masu salmon, O. masou. In particular, interspecific competition may be an important mechanism that could contribute to the exclusion for native salmonids. In this study, experimental pairwise contests were conducted to compare interference competitive ability between native and introduced salmonids. We demonstrated that brown trout were competitively superior to white-spotted charr and masu salmon whereas rainbow trout were superior to white-spotted charr. We suggest that introduced brown trout negatively impact both white-spotted charr and masu salmon, and introduced rainbow trout negatively impact white-spotted charr.     

Genetic differentiation between collections of hatchery and wild masu salmon (<Emphasis Type="Italic">Oncorhynchus masou</Emphasis>) inferred from mitochondrial and microsatellite DNA analyses

There has been very little effort to understand genetic divergence between wild and hatchery populations of masu salmon (Oncorhynchus masou). In this study, we used mitochondrial (mt) NADH dehydrogenase subunit 5 gene (ND5) and six polymorphic nuclear microsatellite DNA loci to compare the genetic variability in three hatchery broodstocks of masu salmon with the variability in eight putative wild masu populations sampled in five rivers including one known source river for the hatchery broodstocks. Both ND5 and microsatellites showed no significant genetic divergence (based on F_ST estimates) between four annual collections from the source river population, suggesting no change in genetic diversity over this time period. The F_ST estimates, an analysis of molecular variance (AMOVA), and a neighbor-joining tree using both DNA markers suggested significant differentiation between the three hatchery and all eight putative wild populations. We conclude that genetic diversity of hatchery populations are low relative to putative wild populations of masu salmon, and we discuss the implications for conservation and fisheries management in Hokkaido.     

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.     

Lack of trophic competition among wild and hatchery juvenile chum salmon during early marine residence in Taku Inlet,Southeast Alaska

Early marine trophic interactions of wild and hatchery chum salmon (Oncorhynchus keta) were examined as a potential cause for the decline in harvests of adult wild chum salmon in Taku Inlet, Southeast Alaska. In 2004 and 2005, outmigrating juvenile chum salmon were sampled in nearshore habitats of the inlet (spring) and in epipelagic habitat at Icy Strait (summer) as they approached the Gulf of Alaska. Fish were frozen for energy density determination or preserved for diet analyses, and hatchery stocks were identified from the presence of thermal marks on otoliths. We compared feeding intensity, diets, energy density, and size relationships of wild and hatchery stocks (n = 3123) across locations and weeks. Only hatchery fish feeding intensity was negatively correlated with fish abundance. In both years, hatchery chum salmon were initially larger and had greater energy density than wild fish, but lost condition in early weeks after release as they adapted to feeding on wild prey assemblages. Diets differed between the stocks at all inlet locations, but did not differ for hatchery salmon between littoral and neritic habitats in the outer inlet, where the stocks overlapped most. Both diets and energy density converged by late June. Therefore, if density-dependent interactions affect wild chum salmon, these effects must be very rapid because survivors in Icy Strait showed few differences. Our study also demonstrates that hatchery release strategies used near Taku Inlet successfully promote early spatial segregation and prey partitioning, which reduce the probability of competition between wild and hatchery chum salmon stocks.     

The effects of introduced salmonids on two native stream‐dwelling salmonids through interspecific competition

Using an artificial stream, habitat use by two sympatric native salmonids in the presence and absence of introduced salmonid species was investigated experimentally. When only native white‐spotted charr Salvelinus leucomaenis and masu salmon Oncorhynchus masou were sympatric, they occupied different microhabitats. In the presence of introduced brown trout Salmo trutta or rainbow trout Oncorhynchus mykiss , however, white‐spotted charr and masu salmon were observed to use a similar habitat and interspecific competition between white‐spotted charr and masu salmon was initiated. The study suggested that the coexistence of native salmonids was negatively affected through interspecific competition between native and introduced salmonids.     

Evidence for competition at sea between Norton Sound chum salmon and Asian hatchery chum salmon

Increasing production of hatchery salmon over the past four decades has led to concerns about possible density-dependent effects on wild Pacific salmon populations in the North Pacific Ocean. The concern arises because salmon from distant regions overlap in the ocean, and wild salmon populations having low productivity may compete for food with abundant hatchery populations. We tested the hypothesis that adult length-at-age, age-at-maturation, productivity, and abundance of a Norton Sound, Alaska, chum salmon population were influenced by Asian hatchery chum salmon, which have become exceptionally abundant and surpassed the abundance of wild chum salmon in the North Pacific beginning in the early 1980s. We found that smaller adult length-at-age, delayed age-at-maturation, and reduced productivity and abundance of the Norton Sound salmon population were associated with greater production of Asian hatchery chum salmon since 1965. Modeling of the density-dependent relationship, while controlling for other influential variables, indicated that an increase in adult hatchery chum salmon abundance from 10 million to 80 million adult fish led to a 72% reduction in the abundance of the wild chum salmon population. These findings indicate that competition with hatchery chum salmon contributed to the low productivity and abundance of Norton Sound chum salmon, which includes several stocks that are classified as Stocks of Concern by the State of Alaska. This study provides new evidence indicating that large-scale hatchery production may influence body size, age-at-maturation, productivity and abundance of a distant wild salmon population.     

Effects of salmon lice infection and salmon lice protection on fjord migrating Atlantic salmon and brown trout post-smolts

Effects of artificial salmon lice infection and pharmaceutical salmon lice prophylaxis on survival and rate of progression of Atlantic salmon (n = 72) and brown trout post-smolts (n = 72) during their fjord migration, were studied by telemetry. The infected groups were artificially exposed to infective salmon lice larvae in the laboratory immediately before release in the inner part of the fjord to simulate a naturally high infection pressure. Groups of infected Atlantic salmon (n = 20) and brown trout (n = 12) were also retained in the hatchery to control the infection intensity and lice development during the study period. Neither salmon lice infection nor pharmaceutical prophylaxis had any effects on survival and rate of progression of fjord migrating Atlantic salmon post-smolts compared to control fish. Atlantic salmon spent on average only 151.2 h (maximum 207.3 h) in passing the 80 km fjord system and had, thus, entered the ocean when the more pathogenic pre-adult and adult lice stages developed. The brown trout, in comparison to Atlantic salmon, remained to a larger extent than Atlantic salmon in the inner part of the fjord system. No effect of salmon lice infection, or protection, was found in brown trout during the first weeks of their fjord migration. Brown trout will, to a larger extent than Atlantic salmon, stay in the fjord areas when salmon lice infections reach the more pathogenic pre-adult and adult stages. In contrast to Atlantic salmon, they will thereby possess the practical capability of returning to freshwater when encountering severe salmon lice attacks.     

Reproduction of interspecific hybrids of Atlantic salmon and brown trout in a stream environment

1. Reproduction between Atlantic salmon males and interspecific hybrid Salmo salar × Salmo trutta females was monitored in a controlled flow channel diverted from a south European river located at the edge of Atlantic salmon natural geographic distribution in Europe. 2. Post‐F1 hybrids were viable and survived in the wild, at least until dispersal from redds. After transfer to hatchery conditions, 67% survived into the second year. 3. The hybrids possessed 98 chromosomes: two sets of Atlantic salmon(2n = 58) and one set of brown trout (n = 40) chromosomes. 4. The existence of a low proportion of allotriploid individuals can be expected in rivers where Atlantic salmon and brown trout populations coexist.     

Size-structured Interactions between Native and Introduced Species: Can Intraguild Predation Facilitate Invasion by Stream Salmonids?

Dynamics of biological invasions may be complicated in size-structured animal populations. Differences in timing of life history events such as juvenile emergence create complex interaction webs where different life stages of native and non-native species act as predators, competitors, and prey. Stream salmonids are an ideal group for studying these phenomena because they display competition and predation in size-structured populations and have been introduced worldwide. For example, introduced rainbow trout (Oncorhynchus mykiss) are invading streams of Hokkaido Island, Japan and have caused declines in native masu salmon (O. masou) populations. However, age-0 rainbow trout emerge later than age-0 masu salmon and are smaller, which raises the question of why they are able to recruit and therefore invade in the face of a larger competitor. We conducted experiments in laboratory stream channels to test effects of increasing density of age-0 and age-1 rainbow trout on age-0 masu salmon. Age-1 rainbow trout dominated age-0 masu salmon by aggressive interference, relegating them to less favorable foraging positions downstream and reducing their foraging frequency and growth. The age-1 trout also reduced masu salmon survival by predation of about 40% of the individuals overall. In contrast, age-0 rainbow trout had little effect on age-0 masu salmon. Instead, the salmon dominated the age-0 trout by interference competition and reduced their survival by predation of 60% of the individuals. In each case, biotic interactions by the larger species on the smaller were strongly negative due to a combination of interspecific competition and intraguild predation. We predict that together these produce a positive indirect effect in the interaction chain that will allow the recruitment of rainbow trout in the face of competition and predation from age-0 masu salmon, and thereby facilitate their invasion in northern Japan.     

An overview of salmon enhancement and the need to manage and monitor natural spawning in Hokkaido, Japan

The chum and pink salmon catches in Hokkaido, Japan have increased dramatically since the 1970s and the 1990s, respectively. In contrast, masu salmon catches have been steadily decreasing. Despite intensive hatchery development in Hokkaido, naturally spawning salmon populations persist based on results from a recent river survey. This paper focuses on the challenges of maintaining hatchery salmon populations while protecting natural chum, pink and masu salmon populations in Hokkaido. Two important initiatives related to meeting this ambitious goal are managing hatcheries in a way that minimizes negative interactions between natural and hatchery salmon populations, and initiating new efforts at restoring and rehabilitating degraded freshwater habitats. In addition, in order to maintain a balance of demand and supply in the domestic market through the exportation of extra salmon, Hokkaido has decided to enter full assessment to gain Marine Stewardship Council (MSC) certification of the Hokkaido chum salmon trap net fishery. This would involve a fundamental shift in fisheries management as practiced in Japan, specifically elevating the importance of managing the fishery in a way that conserves natural salmon populations. A key component of a new salmon management strategy is the establishment of a zone management framework based on the designation of stream units to spatially separate natural salmon from hatchery salmon to minimize negative effects of hatchery fish and to utilize effectively hatchery salmon for commercial fisheries. This effort is allied with similar initiatives in other Pacific Rim countries that are focusing on management reform to restore natural ecosystem function and maintain the coexistence of wild and hatchery salmon.     

Spatial expansion and increased population density of masu salmon parr independent of river restoration

Spatial expansion and increased population abundance of masu salmon Oncorhynchus masou have been recently reported following habitat restoration in some rivers, but no studies have examined these types of changes under natural conditions. We found spatial expansion and increased population density of masu salmon parr in rivers of eastern Hokkaido where no habitat restoration had occurred during the period examined. Changes in the distribution and population densities of parr are often regarded as effects of river habitat destruction and restoration, yet such changes should be evaluated while taking into account the effects of other ocean environmental conditions as well.     

Feeding of wild and hatchery reared Atlantic salmon (<Emphasis Type="Italic">Salmo salar</Emphasis> L.) smolts during downstream migration

In general, hatchery salmonid smolts experience higher mortality during migration than wild smolts, which is suggested to be due to domestication effects and that hatchery fish lack experience of the natural environment. However, possible differences in feeding during smolt migration between hatchery and wild smolts have rarely been addressed. We compared the number of feeding smolts and stomach fullness among wild Atlantic salmon smolts, hatchery-reared smolts released as 1-year-old parr, and hatchery-reared smolts released as 2-year-old smolts during their descent to sea in River Tornionjoki. In addition, estimations of prey selection among the smolt groups were conducted. A high proportion of wild smolts and smolts stocked as parr actively fed during the smolt migration. A lower proportion of smolts stocked as smolts was feeding and their stomach fullness were much reduced in comparison with the two other groups. The study also indicated that the feeding of migrating smolts is selective rather than opportunistic. In conclusion, this study suggests that stocked 2-year-old smolts may enter sea with an inferior foraging behaviour and it is a possibility that this may contribute to the observed low post-smolt survival in the Baltic Sea.     

New polymorphic microsatellite markers for the masu salmon (Oncorhynchus masou masou) from Korea and their application to wild and hatchery populations

Masu salmon, Oncorhynchus masou masou, is an economically important fish species in the Far East and occurs in two life history forms: sea-run migratory (anadromous) and freshwater resident (non-anadromous). The non-anadromous form has recently become a popular freshwater food and game fish during a well-known Korean winter festival. However, the genetic background of this species remains largely unknown, partly due to a lack of molecular genetic markers. In this study, we developed new polymorphic microsatellite markers for masu salmon using next-generation sequencing technology. From 40 primer sets, 11 primer sets (27.5% of the primer sets selected) were successfully amplified with 106 alleles (range 2–9) in 64 individuals from different populations: two wild and one hatchery. Observed and expected heterozygosities ranged from 0.304 to 0.947 and 0.278 to 0.865, respectively. Significant departures from the Hardy–Weinberg equilibrium were detected for four markers (OMM11, OMM17, OMM28, and OMM33) in a single population. All pair-wise F_ST values were highly significant between the wild and hatchery populations (range 0.084–0.183, P < 0.0001). We identified a set of robust microsatellite markers that worked well even in formalin-fixed samples, which will be suitable for biogeographical and population structure analyses of the masu salmon.     

Relationship between eyed-egg percentage and levels of cortisol and thyroid hormone in masu salmon Oncorhynchus masou

A possible cause of the low eyed-egg percentage that afflicts pond-reared masu salmon Oncorhynchus masou at a fish hatchery (Kumaishi Fish Hatchery, Hokkaido, Japan) was investigated. Serum cortisol levels of masu salmon during the spawning period were compared between individuals from Kumaishi and those from another station (Mori). Cortisol, thyroxine (T₄) and triiodothyronine (T₃) hormones were also measured in eyed eggs and their levels were compared to the eyed-egg percentage. Serum cortisol levels were significantly higher at Kumaishi than at Mori in May and July of the breeding season. At the Kumaishi station, there was a linear positive relationship between serum cortisol and fertilized-egg cortisol levels of female masu salmon. As cortisol levels in both ovulating females and eyed eggs increased, the eyed-egg percentage decreased, indicating a highly significant negative relationship between cortisol levels and eyed-egg percentage. In contrast, as T₃ and T₄ levels in eyed eggs increased, the eyed-egg percentage increased. The strong positive correlation between high cortisol levels in serum and in eyed eggs at Kumaishi indicates that the frequently higher cortisol levels may have caused the lowering of the eyed-egg percentage.     

Effects of body size and predators on intracohort competition in wild and domesticated juvenile salmon in a stream

Competition between masu salmon (Oncorhynchus masou) of wild and aquaculture origin was investigated. Fry were individually marked and released in stream enclosures with and without a piscivorous predator. The aim was to assess the effects of predators and salmon body size on survival and growth of the two types of fish under natural conditions. The presence of predaceous Japanese huchen (Hucho perryi) resulted in lower mean growth rates of surviving fry. Relatively large fish survived and grew better than relatively small fish in the absence of predators, but not in their presence. This probably indicates an indirect effect of predation risk on within-cohort competitive behavior among salmon juveniles, with larger fish forced to give up their position as superior competitors. Domesticated fish survived in larger numbers and grew much faster than wild fish, irrespective of predator presence. Comparison with similar field studies revealed a pattern that the pre-experimental environment influenced the outcome of competition between wild and domesticated juvenile salmon. Domesticated fish were superior competitors even in the absence of an initial size advantage, which commonly gives a further advantage to hatchery-raised fish in natural streams. Therefore, caution dictates to avoid the release or escape of such fish into the wild.     

Differential dispersal patterns of male and female masu salmon fry

Behavioural experiments using artificial and natural stream channels were undertaken to determine whether there were differences in dispersal between newly emerged male and female masu salmon. Eyed eggs from a cross with wild spawners were planted in the middle pool of an artificial channel. After emergence, more males than females moved into an upstream trap, while fewer males moved downstream. In a natural stream, eyed eggs were marked with alizarin complexone to distinguish them from naturally spawned eggs and these were planted into artificial redds. More newly emerged male fry remained at the planted site than female fry. In contrast, more females moved downstream than males. These results imply that differences in dispersal patterns between male and female masu salmon fry are genetically controlled.     

Consequences to fitness‐related traits of hybridization between farmed and wild Atlantic salmon, Salmo salar

Increasing concern has been expressed about the genetic effects of cultured salmonid fishes on natural populations. Avoidance of extreme negative outcomes was one reason for the establishment of a genetic management policy for the State of Alaska. However, domestication within the hatchery may still cause divergence from the wild donor population. This divergence could potentially lead to adverse impacts on wild stocks through straying and introgression. This study examines potential domestication in two Alaskan chinook salmon stocks. The Little Port Walter (LPW) Hatchery Chickamin River stock resulted from a small collection of wild broodstock in 1976. The LPW Unuk stock was founded with a larger number of individuals in 1976 and has had subsequent infusion of wild gametes. These lines have been maintained at LPW through ocean ranching of tagged smolts. Comparisons are made between the hatchery lines, progeny of wild chinook collected from the Chickamin and Unuk Rivers, and hybrids between the hatchery and wild groups. Mature ocean‐ranched female chinook salmon returning to the facility were periodically graded for ripeness and spawned. Body size and meristic measurements were collected from these mature spawners. Maturation timing, fecundity, and individual egg size of these fourth generation hatchery fish are compared with that of offspring of wild fish from the same donor stock. Stock of origin is confirmed for all spawners and offspring using microsatellite DNA analysis.