The Impact of Escaped Farmed Atlantic Salmon (Salmo salar L.) on Catch Statistics in Scotland

In Scotland and elsewhere, there are concerns that escaped farmed Atlantic salmon (Salmo salar L.) may impact on wild salmon stocks. Potential detrimental effects could arise through disease spread, competition, or inter-breeding. We investigated whether there is evidence of a direct effect of recorded salmon escape events on wild stocks in Scotland using anglers' counts of caught salmon (classified as wild or farmed) and sea trout (Salmo trutta L.). This tests specifically whether documented escape events can be associated with reduced or elevated escapes detected in the catch over a five-year time window, after accounting for overall variation between areas and years. Alternate model frameworks were somewhat inconsistent, however no robust association was found between documented escape events and higher proportion of farm-origin salmon in anglers' catch, nor with overall catch size. A weak positive correlation was found between local escapes and subsequent sea trout catch. This is in the opposite direction to what would be expected if salmon escapes negatively affected wild fish numbers. Our approach specifically investigated documented escape events, contrasting with earlier studies examining potentially wider effects of salmon farming on wild catch size. This approach is more conservative, but alleviates some potential sources of confounding, which are always of concern in observational studies. Successful analysis of anglers' reports of escaped farmed salmon requires high data quality, particularly since reports of farmed salmon are a relatively rare event in the Scottish data. Therefore, as part of our analysis, we reviewed studies of potential sensitivity and specificity of determination of farmed origin. Specificity estimates are generally high in the literature, making an analysis of the form we have performed feasible.     

Disentangling stocking introgression and natural migration in brown trout: survival success and recruitment failure in populations with semi‐supportive breeding

1. Introgression into natural salmonid populations from stocked conspecifics has been widely studied. Outcomes vary from no effect even after decades of stocking, to population replacement after only a couple of generations. Potential introgression caused by semi‐supportive breeding (i.e. using a mixture of local strains as brood stock) is, however, less well studied. 2. We investigated population structure of brown trout (Salmo trutta) in a regulated alpine lake with three natural, environmentally contrasting tributaries used as spawning and rearing habitat. Massive semi‐supportive breeding of admixed local strains has been implemented for decades. Stocked trout represented c. 17% of the total lake population, and a substantial post‐release survival reflects a considerable potential for introgression. However, the mark‐recapture studies indicate no spawning runs of stocked fish. 3. Using 13 polymorphic microsatellite loci, we found natural straying and non‐native reproduction, especially among wild populations inhabiting environmentally unstable habitat. Retained genetic structure across tributaries indicated low reproductive success of wild‐born non‐natives. Moreover, the genetic structure among tributaries has probably not been influenced by semi‐supportive breeding, because of recruitment failure of stocked trout.     

Minimal gene flow from introduced brown trout (Salmo trutta L.) after 30 years of stocking

Lake Tinnsjø, Norway, has been heavily stocked over three decades with two different brown trout stocks. A population genetic survey based on 13 allozyme coding loci was conducted to investigate the genetic composition of the present trout population and the genetic impact of the two donors. Contrary to expectations, highly significant differences in allelic frequencies between Lake Tinnsjø trout and the two donor stocks were found at several loci, suggesting minimal gene flow from stocked trout to the wild populations. Pairwise genetic distance values were several times higher between donor stocks and Lake Tinnsjø samples than between the various spawning inlets to Lake Tinnsjø. F_st increased from 0.024 to 0.090, when samples from the donor stocks were included in the material. There were no deviations from expected Hardy–Weinberg distribution of genotypes in the spawning inlets. It was concluded that the donor stocks contributed little to the trout gene pool in the lake. A possible exception is the trout population below the lake outlet.     

Identifying footprints of selection in stocked brown trout populations: a spatio‐temporal approach

Studies of interactions between farmed and wild salmonid fishes have suggested reduced fitness of farmed strains in the wild, but evidence for selection at the genic level is lacking. We studied three brown trout populations in Denmark which have been significantly admixed with stocked hatchery trout (19–64%), along with two hatchery strains used for stocking. The wild populations were represented by contemporary samples (2000–2006) and two of them by historical samples (1943–1956). We analysed 61 microsatellite loci, nine of which showed putative functional relationships [expressed sequence tag (EST)‐linked or quantitative trait loci]. F_ST‐based outlier tests provided support for diversifying selection at chromosome regions marked by three loci, two anonymous and one EST‐linked. Patterns of differentiation suggested that the loci were candidates for being under diversifying hitch‐hiking selection in hatchery vs. wild environments. Analysis of hatchery strain admixture proportions showed that in one wild population, two of the loci showed significantly lower admixture proportions than the putatively neutral loci, implying contemporary selection against alleles introduced by hatchery strain trout. In the most strongly admixed population, however, there was no evidence for selection, possibly because of immigration by stocked trout overcoming selection against hatchery‐derived alleles or supportive breeding practices allowing hatchery strain trout to escape natural selection. To our knowledge, this is the first study demonstrating footprints of selection in wild salmonid populations subject to spawning intrusion by farmed fish.     

Use of micreosatellite markers for identification of indigenous brown trout in a geographical region heavily influenced by stocked domensticated trout

Based on estimates of genetic differentiation between populations, assignment tests and analysis of isolation by distance, stocked populations of brown trout Salmo trutta of Funen Island, Denmark, had been genetically affected by domesticated trout, whereas the stocking of wild exogenous trout into one of the rivers had little or no impact. At the same time, there were clear indications of remaining indigenous gene pools in the Funen populations. The management implications of these findings are discussed and changes in trout release activity are recommended to avoid further mixing of trout gene pools.     

Evidence for selective angling of introduced trout and their hybrids in a stocked brown trout population

Comparisons of the genetic composition of brown trout Salmo trutta captured by anglers and by electrofishing based on three diagnostic microsatellite loci provided strong evidence that angling is selective in a stocked brown trout population. At two sites, anglers caught significantly younger trout and proportionally more introduced hatchery trout and hybrids than were observed in electrofishing surveys. Selective angling, in combination with a small legal catch size, may have considerably eliminated introduced trout and hybrids before spawning at the study sites, and thus may have reduced the introgression of alien genes into the local gene pool. Angling can be an important factor influencing the genetic structure of fish populations and should be taken into account in studies of introgressive hybridization in stocked fish populations and their management. In this study, demographic consequences of stocking were not assessed. Thus, even though the genetic consequences of stocking may be minimal or largely reversible through angling, resource competition between native and introduced trout, until they reach legal catch size, is expected to have a negative effect on the productivity of the indigenous trout population.     

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.     

Allozyme diversity in brown trout (Salmo trutta) from Central Spain: Genetic consequences of restocking

1.  The brown trout ( Salmo trutta ) represents one of the main freshwater resources in Spain, but habitat alterations and overharvesting have contributed to the decline or disappearance of numerous natural populations. In addition, reinforcement programs of wild populations based on releases of hatchery reared fish of exogenous origin compromise the conservation of remnant native trout resources.
2.  We present allozymic data from Central Spain trout populations including stocked and unstocked populations. Although the levels of genetic variation observed were low and affected by hatchery releases (p = 18.23%, Ho= 3.39%), they were within the range observed in other European areas.
3.  The effective introduction of hatchery reared fish is genetically homogenising the populations in the studied area and disturbing the ancestral pattern of genetic variation that distinguishes the Tajo and Duero basins. Within the eight natural populations analysed, seven had alleles assigned to the foreign trout. The introgression in these populations, following the LDH-5 * 90 allele frequency, ranged between 2% and 29.4%, but those values are not in concordance with the respective stocking effort undertaken in each population. Moreover, the release of hatchery-reared fish does not solve the problems related to the reduced size of wild populations and their recruitment instability.     

Genetic introgression between wild and stocked brown trout in the Douro River basin, Spain

The genetic diversity of Spanish brown trout is currently threatened by stocking with exogenous brown trout from Central and Northern Europe. In the Douro River basin 25% of the analysed populations in the present study showed introgression by genes of hatchery origin. The mean introgression estimated by the single locus approach ( S ) varied from 0 to 22% among populations, with a mean value of 3%. The hatchery allele markers were absent in populations where stocking ceased in 1993. However, the introgression effect was observed in all populations stocked until 1998. It seems that cessation of stocking is a good measure for restoring native populations. A thorough review of published and present data of genetic interactions between wild and stocked brown trout in Spanish rivers indicates different levels of introgression between basins. The absence of a clear geographical pattern in the introgression level suggests that ecological interactions and local stocking programmes may play an important role in stocking success. Finally, several guidelines are provided for conservation and management of native brown trout populations in Spanish rivers.     

Effects of sea trout stocking on the population genetics of landlocked brown trout, Salmo trutta L., in the Conwy River system, North Wales, U.K.

The effect of the introduction of fry of anadromous sea trout, Salmo trutta L., on the genetic integrity of landlocked brown trout populations was evaluated. Samples were taken from six brown trout populations from streams above impassable waterfalls in the Conwy river system (North Wales, U.K.) in 1989 and 1990. Three of these streams had no known stocking history and three had been stocked with sea trout fry from the lower Conwy system over the last few years. Representatives of these sea trout were collected from two streams in the lower Conwy system and from a hatchery. Allele frequencies at 13 loci, six of which were polymorphic, were determined by starch gel electrophoresis.
The stocked populations were intermediate in their allele frequencies between unstocked brown trout and sea trout samples. A principal component analysis suggested significant numbers of hybrids in all of the stocked streams. This shows that some of the introduced sea trout did not migrate down the falls to the sea, but stayed in fresh water and hybridized with the local population. The significance of this finding for the conservation of the genetic resource of brown trout stocks is discussed.     

Genetic differentiation of populations of the copepod sea louse Lepeophtheirus salmonis (Krøyer) ectoparasitic on wild and farmed salmonids around the coasts of Scotland: Evidence from RAPD markers

Sea trout are the sea-going migratory form of the freshwater brown trout (Salmo trutta L.) and since 1989 there have been marked declines in their stocks on the west coasts both of Scotland and Ireland. Various factors have been attributed as possible causal agents in these stock declines, including fresh water acidification, overfishing, climatic fluctuations, habitat degradation and sea lice parasitic burdens. The putative impact of infestations of sea trout by the ectoparasitic copepod sea louse, Lepeophtheirus salmonis (Krøyer), has featured prominently in the controversy, especially with regard to the role of inshore commercial salmon farms as a possible source of infestation of wild salmonids by sea lice. This study focused on the population genetics of L. salmonis around the coasts of Scotland: We sampled fish from wild and cultured stocks and included salmon (Salmo salar L.), rainbow trout (Oncorhynchus mykiss Walbaum) and sea trout as host species. Analyses of allozyme variation of sea lice were confined to data for two polymorphic loci (Fum, Got-2) and conformed to our initial expectation — that the inclusion of a planktonic larval phase in the life cycle of the copepod, in addition to the high mobility of the host fish, would enhance gene flow and preclude genetic differentiation of L. salmonis populations as a result of random drift alone. DNA polymorphism was quantified by means of PCR and RAPD analysis. Six primers were screened for 16 samples (from wild and farmed salmon, wild sea trout and farmed rainbow trout) — including the east, north and west coasts of Scotland — and the data analyzed by AMOVA (Analysis of Molecular Variance). In contrast to the allozyme results, the RAPD analysis showed striking patterns of genetic differentiation around the coasts of Scotland. The overall pattern was one of genetic homogeneity of L. salmonis populations sampled from wild salmon and sea trout. All of the L. salmonis samples taken from farmed salmon and rainbow trout did, however, show highly significant levels of genetic differentiation, both between wild and farmed salmonids and among the various farms themselves. Evidence of high levels of small-scale spatial or temporal heterogeneity of RAPD marker band frequencies was shown for the one farm from which repeat samples (July and November, 1995) were analysed. Samples of sea lice taken from west coast wild sea trout subjected to RAPD analysis also revealed the occurrence of putative “farm markers” in some individual parasites, indicating that they had possibly originated from salmon farms.     

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.     

Genetic population structure in flatfishes and potential impact of aquaculture and stock enhancement on wild populations in Europe

Marine fish wild stocks are known to be heavily depleted by overfishing and flatfish species are no exception. Wild catches being soon insufficient for responding to consumer demand, the cultivation of marine species appeared as a logical response to the need of seafood. Nevertheless, fish aquaculture also entails major impacts on wild populations from which genetic ones are now better known. The hybridization between domestic and native strains potentially have a genetic impact on recipient populations as long as 1) domestic populations are distinct from native wild ones (through domestication process, genetic improvement of captive stocks) and/or 2) the native wild populations are structured (metapopulation structure, local adaptation). Some of the flatfish species exhibit population differentiation and even local adaptation and the release of domestic genetically modified fishes (selected, transgenic) could threaten their survival in case of introgression. The impact of aquaculture on flatfishes is probably still low as land-based farms and low production levels guaranty low rates of escapes and therefore limited contacts between wild and farmed strains. However, flatfish aquaculture is regarded by experts as a rapidly growing domain that will greatly develop soon. In our opinion, this perspective, added to the quite good performances of farmed flatfishes when released into the wild, fully justifies a stronger interest from the scientific community to the conservation of their wild stocks.     

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.     

Phylogeography,genetic structure,and conservation of the endangered Caspian brown trout, <Emphasis Type="Italic">Salmo trutta caspius</Emphasis> (Kessler, 1877), from Iran

The Caspian Sea, the largest inland closed water body in the world, has numerous endemic species. The Caspian brown trout (Salmo trutta caspius) is considered as endangered according to IUCN criteria. Information on phylogeography and genetic structure is crucial for appropriate management of genetic resources. In spite of the huge number of studies carried out in the Salmo trutta species complex across its distribution range, very few data are available on these issues for S. trutta within the Caspian Sea. Mitochondrial (mtDNA control region) and nuclear (major ribosomal DNA internal transcribed spacer 1, ITS-1, and ten microsatellite loci) molecular markers were used to study the phylogeography, genetic structure, and current captive breeding strategies for reinforcement of Caspian trout in North Iranian rivers. Our results confirmed the presence of Salmo trutta caspius in this region. Phylogenetic analysis demonstrated its membership to the brown trout Danubian (DA) lineage. Genetic diversity of Caspian brown trout in Iranian Rivers is comparable to the levels usually observed in sustainable anadromous European brown trout populations. Microsatellite data suggested two main clusters connected by gene flow among river basins likely by anadromous fish. No genetic differences were detected between the hatchery sample and the remaining wild populations. While the current hatchery program has not produced detectable genetic changes in the wild populations, conservation strategies prioritizing habitat improvement and recovering natural spawning areas for enhancing wild populations are emphasized.     

Effects of stocking on the genetic diversity of brown trout populations of the Adriatic and Danubian drainages in Switzerland

This study focuses on genetic variation of brown trout Salmo trutta populations of the Adriatic and Danubian drainages in Switzerland. The allozyme and other protein loci data show a major replacement of native stocks from the Adriatic drainages by introduced hatchery trout of Atlantic basin origin. In most samples, diagnostic alleles for the Adriatic form of Salmo trutta f. fario and for the marbled trout Salmo trutta marmoratus are found at very low frequencies (f<0.15). Taking into account previous genetic studies on brown trout of this basin, the Danubian samples are not heavily contaminated with foreign alleles. The results are consistent with records of local stocking activities which account in part for the high introgression rates of Atlantic alleles into local populations of the Adriatic drainages. In addition, introgression is enhanced by a decrease of natural reproduction which is caused by a deterioration of trout habitats through human activities. Furthermore, a third mechanism is proposed that may contribute to the high introgression rates observed: if Atlantic trout are introduced, the reproductive barriers between the two native forms, marbled trout and Adriatic fario respectively, break down. Atlantic trout apparently hybridize with both native forms and generate gene flow between them. In some parts of Adriatic drainages in Switzerland, the patterns of introgression and hybridization are further complicated by introduction of trout from the Danubian system. Alleles of the marbled trout are also found in the samples of the Danubian drainage system. These are due to stocking activities across the watershed.     

Genetic structure of important resident brown trout breeding lines in Poland

The history of brown trout Salmo trutta L. stocking has long tradition in the European Union and other countries. Hundreds of hatchery facilities on continent have artificial broodstocks used for enhancement of neighbouring and also geographically far river basins. These practices have substantial effect on wild brown trout populations. To illuminate this phenomenon, eleven hatchery stocks and wild populations from northern Poland and Carpathian region were analysed using 13 microsatellite markers. Obtained results revealed high genetic diversity between studied stocks and clear differentiation between northern and southern populations and hybridization between these two major clads. As a recommendation, the principle of treating regions as metapopulations should be applied, which, in the case of Poland, means using the division of the northern and southern genetic lines that were revealed in the present study.     

Escape of farmed tilapiines into the wild and entry of wild forms in fishponds,and the possible interactions between wild and farmed tilapiines from a sample of smallholder farms in Central Uganda

Seven smallholder fishponds in central Uganda were studied between 2000 and 2001 to investigate the interaction of farmed tilapiines with their wild conspecifics. Emphasis was on the features that facilitate escape of fish and/or entry of fish into the farms and interactions between the farmed and the wild. These included number of species, source of seed, connection between pond and natural watercourses, purpose of the farm, destination of cultured fish and interaction between farmed and wild fish. Fishponds had no screens against entry of wild fish into the farm or escape of farmed fish into the wild and occurred within wetlands close to natural watercourses. Ponds stocked with one fish species were found to have multispecies with some individuals that were apparently intermediate morphs between the species. Fry produced within the growout fishponds was shared with other farmers within and outside the watersheds. This study showed that smallholder farms with little or no control of escape or entry of fish out and into the fishponds, and little or no management present circumstances that facilitate continued movement of tilapiines within and across watersheds in Ugandan waters. The study also indicated possible genetic interaction between farmed and their wild conspecifics through interactions within fishponds.     

Generic genetic differences between farmed and wild Atlantic salmon identified from a 7K SNP-chip

Genetic interactions between farmed and wild conspecifics are of special concern in fisheries where large numbers of domesticated individuals are released into the wild. In the Atlantic salmon (Salmo salar), selective breeding since the 1970's has resulted in rapid genetic changes in commercially important traits, such as a doubling of the growth rate. Each year, farmed salmon escape from net pens, enter rivers, and interbreed with wild salmon. Field experiments demonstrate that genetic introgression may weaken the viability of recipient populations. However, due to the lack of diagnostic genetic markers, little is known about actual rates of gene flow from farmed to wild populations. Here we present a panel of 60 single nucleotide polymorphisms (SNPs) that collectively are diagnostic in identifying individual salmon as being farmed or wild, regardless of their populations of origin. These were sourced from a pool of 7000 SNPs comparing historical wild and farmed salmon populations, and were distributed on all but two of the 29 chromosomes. We suggest that the generic differences between farmed and wild salmon at these SNPs have arisen due to domestication. The identified panel of SNPs will permit quantification of gene flow from farmed to wild salmon populations, elucidating one of the most controversial potential impacts of aquaculture. With increasing global interest in aquaculture and increasing pressure on wild populations, results from our study have implications for a wide range of species.