Genetic determination of the contribution of stocked and wild Atlantic salmon, Salmo salar L., to the angling fisheries in two Spanish rivers

From 1983 to 1986 the Rivers Ason and Nansa (northern Spain) were stocked with over 100 000 eyed ova year⁻¹ from a Scottish hatchery in an effort to increase angling catches of Atlantic salmon. An inherent genetic marker was used to estimate the contributions of stocked and wild fish from these year-classes to the 1988 rod-and-line fishery. Stocking with foreign ova yielded 0.9 fish per 10 000 ova in the R. Asón and zero fish in the R. Nansa. Natural egg deposition yielded 4.1 and 4.9 fish per 10000 ova, respectively. Reasons for the difference in performance between stocked and wild fish are examined.     

Genetic implications of hatchery rearing in Atlantic salmon: effects of rearing environment on genetic composition

In an experiment to investigate genetic consequences of hatchery rearing in salmon, allozyme variation at five polymorphic loci was examined in Atlantic salmon of known initial genetic composition, which were reared throughout freshwater life in the hatchery or stocked into the wild as swim-up fry. The genetic composition of the juveniles in the hatchery remained homogeneous from fertilization up to stocking, and from stocking to 2+ in the wild, however, those remaining at the hatchery developed genetic differences among smolting and nonsmolting 1+ parr. These differences were attributed to conditions leading to early smolting at 1+ among the hatchery fish, with 1+ smolts diverging from the gene pool from which they were derived, whereas those stocked into the wild did not smolt until a year later and retained the original genetic composition. The results are discussed in relation to hatchery rearing of salmon and implications for the use of reared fish in stocking and enhancement programmes.     

The timing, sex and age composition of the wild and reared Atlantic salmon ascending the Simojoki River, northern Finland

In the Simojoki River in the northern Gulf of Bothnia, reared salmon stocked as smolts produced considerable numbers of ascending one‐sea‐winter (1 SW) males, whereas the proportion of male 1 SW salmon was low among spawning migrants of wild or reared parr origin. The sex ratio among ascending wild fish and reared salmon stocked as parr was similar, with females predominating, while reared salmon stocked as smolts were mainly males. The multi‐sea‐winter (MSW) salmon entered the river annually within a fairly short time period from the beginning of the migration season, independent of their sex or origin. 1 SW males migrated into the river significantly later in the season than MSW males. The results indicate that the delayed opening of the fishing season in the Gulf of Bothnia is effective in reducing the harvest of MSW salmon at sea. However, as the timing of the ascent may vary by several weeks from year to year, the effect of this regulation bound to certain calendar days may also vary considerably from year to year.     

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.     

Interactions between natural and farmed fish populations: information from genetic markers

Studies on genetic changes in farmed fish populations are reviewed, and the potential interactions between wild and farm escapee, and between wild and stocked, fish populations are discussed. Examples of the application of genetic markers in studies concerning survival and reproduction of stocked fish, and genetic and ecological interactions between stocks, are given for brook trout, Salvelinus fontinalis , brown trout, Salmo trutta , rainbow trout, Salmo gairdneri , cod, Gadus morhua , Guadalupe bass, Micropterus treculi , walleye, Stizostedion vitreum vitreum and chum salmon, Oncorhynchus keta . The various studies produced different results. Evidence for successful reproduction and genetic interactions between released and wild stocks have been found in a few cases. Stocked genetic material sometimes had a lower reproductive success than wild material. In one case the transplanted genetic material failed to acclimatize, and was apparently lost from the genepool in two generations. Investigations on the genetic and ecological interactions between wild and farmed populations are of great importance to the preservation of wild populations and their genetic resources.     

The survival of semi‐wild, wild and hatchery‐reared Atlantic salmon smolts of the Simojoki River in the Baltic Sea

The recapture rate and survival of hatchery‐reared Atlantic salmon Salmo salar stocked as 1 year‐old parr (semi‐wild) with that of hatchery‐reared Atlantic salmon stocked as 2 year‐old smolts and wild smolts of Atlantic salmon in the northern Baltic Sea were compared. This was done through tagging experiments carried out in 1986–1988 and 1992. The recapture rate of the semi‐wild groups varied from 1·0 to 13·1%, being similar in 3 tagging years and lower in 1 year than that of the wild groups (1·7–17·0%). The recapture rate of the semi‐wild groups was similar (in 2 years) or higher (in 2 years) than that of the hatchery‐reared groups stocked as smolts (1·3–6·3%). The survival of semi‐wild smolts during the sea migration was as high as that of wild Atlantic salmon of an equal size and two to three times higher than hatchery‐reared Atlantic salmon stocked as smolts. The survival rate was positively associated with smolt size. The suitability of hatchery‐reared parr and smolts in the management of reduced Atlantic salmon stocks is compared.     

Maladaptation and phenotypic mismatch in cultured Atlantic salmon used for stocking

Compared to wild populations, cultured Atlantic salmon often sustain higher mortality rates and lower adult return rates when stocked as juveniles into natural streams. The ultimate causes for such differences in fitness, however, are not clear. Here we tested if relaxed levels of natural selection and improved survival in the hatchery environment could account for the observed degree of maladaptation among stocked fish. To do this, we assessed the degree of phenotypic mismatch between wild and cultured fish in three populations over five consecutive years. Significant differences were found in several phenotypic traits that are likely to have fitness implications. Thus, if the objective is to mimic wild individuals for restoration purposes, current hatchery practices aimed at maximising juvenile survival and enhancing growth may need to be revised.     

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.     

Changes in the genetic structure of Atlantic salmon populations over four decades reveal substantial impacts of stocking and potential resiliency

While the stocking of captive‐bred fish has been occurring for decades and has had substantial immediate genetic and evolutionary impacts on wild populations, its long‐term consequences have only been weakly investigated. Here, we conducted a spatiotemporal analysis of 1428 Atlantic salmon sampled from 1965 to 2006 in 25 populations throughout France to investigate the influence of stocking on the neutral genetic structure in wild Atlantic salmon (Salmo salar) populations. On the basis of the analysis of 11 microsatellite loci, we found that the overall genetic structure among populations dramatically decreased over the period studied. Admixture rates among populations were highly variable, ranging from a nearly undetectable contribution from donor stocks to total replacement of the native gene pool, suggesting extremely variable impacts of stocking. Depending on population, admixture rates either increased, remained stable, or decreased in samples collected between 1998 and 2006 compared to samples from 1965 to 1987, suggesting either rising, long‐lasting or short‐term impacts of stocking. We discuss the potential mechanisms contributing to this variability, including the reduced fitness of stocked fish and persistence of wild locally adapted individuals.     

Dispersal and survival of stocked cyprinids in a small English river: comparison with wild fishes using a multi-method approach

Mark–recapture and fixed-station passive integrated transponder (PIT) telemetry were used to compare movements, distribution and survival of stocked juvenile chub Leuciscus cephalus and roach Rutilus rutilus with those of wild conspecifics. Daily activity of wild fish activity was affected by a combination of river flow and temperature, whereas stocked fishes were not influenced by environmental factors. PIT telemetry recorded exploratory movements of stocked L. cephalus immediately after stocking, a substantial number of stocked fish moved both downstream and upstream during periods of elevated flow, and proportionally more stocked fish moved during the first 6 weeks after release than later on. Proportionally more stocked fish than wild fish moved through PIT antennae, stocked L. cephalus moved greater distances than wild L. cephalus and were more widely distributed than wild fish. Minimum estimates of survival after 5 months were 50·5% for stocked R. rutilus and 28·0% for stocked L. cephalus . Ultimately, stocked cyprinids appeared to be able to cope with elevated flows and most remained in the river section local to the stocking location.     

Determinants of hierarchical genetic structure in Atlantic salmon populations: environmental factors vs. anthropogenic influences

Disentangling the effects of natural environmental features and anthropogenic factors on the genetic structure of endangered populations is an important challenge for conservation biology. Here, we investigated the combined influences of major environmental features and stocking with non‐native fish on the genetic structure and local adaptation of Atlantic salmon (Salmo salar) populations. We used 17 microsatellite loci to genotype 975 individuals originating from 34 French rivers. Bayesian analyses revealed a hierarchical genetic structure into five geographically distinct clusters. Coastal distance, geological substrate and river length were strong predictors of population structure. Gene flow was higher among rivers with similar geologies, suggesting local adaptation to geological substrate. The effect of river length was mainly owing to one highly differentiated population that has the farthest spawning grounds off the river mouth (up to 900 km) and the largest fish, suggesting local adaptation to river length. We detected high levels of admixture in stocked populations but also in neighbouring ones, implying large‐scale impacts of stocking through dispersal of non‐native individuals. However, we found relatively few admixed individuals suggesting a lower fitness of stocked fish and/or some reproductive isolation between wild and stocked individuals. When excluding stocked populations, genetic structure increased as did its correlation with environmental factors. This study overall indicates that geological substrate and river length are major environmental factors influencing gene flow and potential local adaptation among Atlantic salmon populations but that stocking with non‐native individuals may ultimately disrupt these natural patterns of gene flow among locally adapted populations.     

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.     

Observations on the dietary habits of wild and stocked brown trout, Salmo trutta L., in Irish lakes

The dietary habits and feeding rates of wild and stocked brown trout were compared for populations in a number of Irish lakes. Wild trout and stocked fish, which had been present in a fishery for 12 months or longer, tend to feed on the same dietary items at similar rates. Stocked fish in their immediate post-planting period (1–14 days) ate less than both the wild trout and established planted fish. In some instances recently stocked fish appear to have a preference for surface food items. They also consumed stones and detritic material. Data indicate that stocked fish adopted a natural diet in less than 5 months. Results are discussed in relation to angling crops of wild and stocked fish and the comparative success of autumn and spring plantings of salmonids.     

Genetic and phenotypic changes in an Atlantic salmon population supplemented with non-local individuals: a longitudinal study over 21 years

While introductions and supplementations using non-native and potentially domesticated individuals may have dramatic evolutionary effects on wild populations, few studies documented the evolution of genetic diversity and life-history traits in supplemented populations. Here, we investigated year-to-year changes from 1989 to 2009 in genetic admixture at 15 microsatellite loci and in phenotypic traits in an Atlantic salmon (Salmo salar) population stocked during the first decade of this period with two genetically and phenotypically distinct source populations. We detected a pattern of temporally increasing introgressive hybridization between the stocked population and both source populations. The proportion of fish returning to the river after a single winter at sea (versus several ones) was higher in fish assigned to the main source population than in local individuals. Moreover, during the first decade of the study, both single-sea-winter and multi-sea-winter (MSW) fish assigned to the main source population were smaller than local fish. During the second decade of the study, MSW fish defined as hybrids were lighter and smaller than fish from parental populations, suggesting outbreeding depression. Overall, this study suggests that supplementation with non-local individuals may alter not only the genetic diversity of wild populations but also life-history traits of adaptive significance.     

Stocking hatchery‐reared brown trout in different densities into a wild population – a comparison of growth and movement

In spring 2001 and 2002 a small stream was stocked with tagged hatchery‐reared yearling brown trout ( Salmo trutta ), in order to study their influence on the resident brown trout population. The stream was separated into six sections: two sections without stocking, two sections where stocking doubled the trout population and two sections where the fish population was quadrupled. The working hypothesis was that due to food limitation (competition) growth of the wild fish will be negatively influenced by stocking, and wild fish will be displaced by the (possibly more aggressive) hatchery fish. Surprisingly, growth rate of wild and stocked fish of the same age was similar and independent of stocking density. Two main reasons may be responsible for this finding: only a low percentage of the stocked fish remained in the stream, and food was not limited during summer. Only 12–19% of the stocked fish were recaptured after six months, in contrats to 40–70% of one‐year old and up to 100% of older wild trout. The wild fish were not displaced by hatchery‐reared fish: During summer the wild fish remained more or less stationary, whereas most of the stocked trout had left their release site. The results indicate that in a natural stream stocking of hatchery reared brown trout does not influence negatively growth and movement of the wild fish independent of stocking density.     

Testing predictions of the critical period for survival concept using experiments with stocked Atlantic salmon

Two separate field experiments were performed in the U.S.A. and Norway with stocked Atlantic salmon Salmo salar . In the Norwegian experiment, the offspring of early‐spawning fish which had larger eggs and emerged a few days before offspring of later spawning fish had consistently higher survival rates. In the U.S.A. experiment, stream sections with higher proportions of favourable foraging locations during the critical period (the transition from dependence on maternally‐derived yolk reserves to independent feeding) had lower loss rates of fish stocked as unfed fry. These results provide support for the critical period concept (CPC) in Atlantic salmon, underscores the utility of a manipulative approach to achieve further advances in knowledge of Atlantic salmon ecology and provide additional guidance to management and restoration. A mechanistic, conceptual model for density dependence is presented to identify important knowledge gaps that remain to further evaluate the importance of the CPC for Atlantic salmon population regulation.     

Divergence in smolt production from the stocking of 1-summer-old and 1-year-old Atlantic salmon parr in a northern Baltic river

One‐year‐old and 1‐summer‐old salmon parr have been stocked in the Simojoki, a northern Baltic river, to protect the natural salmon stock from extinction. This enhancement practice was studied during a 5‐year stocking period, when an average of 14.3 1‐summer‐old parr and 5.4 1‐year‐old parr was needed to produce one sea running smolt. The mean yield from stocking was 70 smolts per 1000 stocked 1‐summer‐old parr and 186 smolts per 1000 stocked 1‐year‐old parr. The mean cost of smolts produced by stocking 1‐year‐old and 1‐summer‐old parr was 1.2 and 2.6 times the cost of producing one 50 g smolt in the hatchery, respectively. There was a negative relationship between density of wild parr and length of smolts stocked as 1‐summer‐old parr in the river, but no relationship between wild parr density and length of smolts stocked as 1‐year‐old parr. This indicated that 1‐summer‐old parr were more susceptible than the older parr to competition with wild parr. Considering production costs and smolt yield, stocking of 1‐year‐old parr appeared to be more profitable than that of 1‐summer‐old parr in the enhancement of the endangered salmon stock.     

Infectious salmon anaemia virus in wild fish from Scotland.

Following the outbreak of infectious salmon anaemia (ISA) at salmon farms in Scotland, UK, a survey was established to determine the extent of infection in wild fish. All fish tested were free from the clinical symptoms of ISA. Isolations of ISAV were made from 5 sea trout within areas where ISA affected salmon farms were located. Evidence for ISAV in other sea trout was provided by ISA RT-PCR diagnostic tests. Results from ISA RT-PCR tests reveal evidence for ISAV being present in salmon parr, adult salmon and juvenile brown trout in rivers distant from salmon farms and indicate that, at the time of the survey (1998-1999), ISAV may have been widely distributed. Nucleotide sequence analysis of segments 2 and 8 showed that for most sequences from wild fish there was 100% homology with ISAV isolated from clinically affected farmed fish although evidence is presented which indicates variability in ISAV sequences from wild fish. Modelling the RT-PCR findings indicates that ISAV among salmonid fish was spatially non-random. Brown trout, sea trout and salmon (adult and parr) show a pattern of occasionally large numbers of positive samples against a background of very low numbers.     

Has stocking contributed to an increase in the rod catch of anadromous trout (Salmo trutta L.) in the Shetland Islands,UK?

The stocking of hatchery-origin fish into rivers and lakes has long been used in fisheries management to try to enhance catches, especially for trout and salmon species. Frequently, however, the long-term impacts of stocking programmes have not been evaluated. In this study, the authors investigate the contribution of a stocking programme undertaken to support the rod catch of sea trout in the Shetland Islands, UK. Once a highly productive recreational fishery, Shetland sea trout catches crashed in the mid-1990s. Around the time that stocking began, increases in rod catches were also reported, with advocates of the stocking highlighting the apparent success of the programme. Using a suite of genetic markers (microsatellites), this study explores the contribution of the stocking programme to the Shetland sea trout population. The authors found that the domesticated broodstock and wild spawned brown trout from seven streams were genetically distinct. Despite extensive stocking, wild spawned brown trout dominated, even in those streams with a long history of supplementation. The majority of sea trout caught and analysed were of wild origin – only a single individual was of pure stocked origin, with a small number of fish being of wild × stocked origins. This study suggests that stocking with a domesticated strain of brown trout has made only a very limited contribution to the Shetland Islands rod catch, and that the revival of sea trout numbers appears to be driven almost exclusively by recovery of trout spawned in the wild.     

Density‐dependent growth in hatchery‐reared brown trout released into a natural stream

Hatchery‐reared brown trout Salmo trutta stocked in a natural stream in addition to resident wild brown trout grew more slowly than those stocked with an experimentally reduced density of brown wild trout. In both cases, hatchery‐reared brown trout grew more slowly than resident wild fish in control sections. Mortality and movements did not differ among the three categories of fish. The results showed that growth of stocked hatchery‐reared brown trout parr was density‐dependent, most likely as a consequence of increased competition. Thus, supplementary release of hatchery‐reared fish did not necessarily increase biomass.