Current status of the brown trout (Salmo trutta) populations within eastern Pyrenees genetic refuges

Since the end of the 20th century, some headwaters of rivers in the eastern Pyrenees have been designated as genetic refuges to protect remaining native brown trout (Salmo trutta) diversity. The declaration was based on limited or no evidence of genetic impact from released non‐native Atlantic hatchery fish. Hatchery releases were completely banned into the genetic refuges, but pre‐existing fishing activities were maintained. Specific locations in each refuge have been monitored every 2–3 trout generations to update genetic information to accurately assess the contribution of these reservoirs to the preservation of native brown trout gene pools. This work updates genetic information to year 2014 in three of these locations (in Ter, Freser and Flamisell rivers). Previous studies identified hatchery introgressed populations within refuges and suggested discrepancies between the underlying intention of the genetic refuges and the gene pools detected. Therefore, we also examined genetic divergences among locations inside refuge river segments. Combined information at five microsatellite and the lactate dehydrogenase C (LDH‐C*) loci showed reduced but significant temporal native allele frequency fluctuations in some of the above specific locations that did not modify overall levels of local diversity and river divergences. Bayesian clustering analyses confirmed the presence of differentiated native units within each genetic refuge. Some locations of the Freser River within the genetic refuge area showed high hatchery impact of non‐native fish (over 20%). We discuss additional local actions (releases of native fish, selective removals and fishery reinforcement with sterile individuals) to improve the conservation objective of genetic refuges.     

Assessment of three strategies practiced by fishery managers for restoring native brown trout (Salmo trutta) populations in Northern French Alpine Streams

Abstract – The efficiency of three strategies carried out by fisheries managers to restore native Mediterranean brown trout populations threatened by non‐native Atlantic populations were assessed. The strategies tested were (i) genetic refuge area where stocking is banned, (ii) stocking with Mediterranean fry and (iii) translocation of wild Mediterranean spawners. Using two discriminatory microsatellite loci between Atlantic and Mediterranean alleles, we compared the genetic composition of samples before and after the changes of practices. The three strategies had several detectable effects in the standing populations causing strong temporal changes in departures from Hardy–Weinberg equilibrium and linkage equilibrium between loci and the apparition of new Mediterranean alleles. The significant reductions in the proportions of Atlantic alleles observed over time can mostly be explained by the disappearing of the pure non‐native Atlantic trout after the stopping of hatchery releases. The results, however, also suggest that the active strategies carried out by managers led to intraspecific introgression between both non‐native Atlantic and native Mediterranean strains.     

Erosion of the native genetic resources of brown trout in Spain

Abstract– We analyzed the introduction of hatchery-reared trout in the Riutort Creek, a small stream in the eastern Spanish Pyrennees. We used gene correlation matrices between individuals to analyze the fish coancestry in the Riutort Creek samples and in the hatchery stock. Hatchery fish disturbed the single ancestry in the native population of the creek, and were clearly detected with principal coordinate analysis of the gene correlation matrix. The amount of introgression produced by successful introductions was estimated from the principal coordinate analysis projections of the matrix of F_ST values between the putative native Riutort Creek population, the hatchery stock and the introgressed population. In only two years the amount of introgression rose to 10%, indicating that 5% of the native ancestry is lost each year as a result of the stocking program. Based on these results, we review the present understandings on the genetic impact of hatchery fish on indigenous Spanish brown trout populations. The stocking of these populations involves a non-native broodstock widespread through the Spanish hatcheries, but successful stockings appear to be limited to wild populations subjected to occasional releases in protected or unfished areas. Surprisingly, extensive stocking in fished areas result in a more limited genetic impact on the recipient native population.     

Stocking impact,population structure and conservation of wild brown trout populations in inner Galicia (NW Spain), an unstable hydrologic region

1. Brown trout (Salmo trutta) is an important conservation resource in the Iberian Peninsula. The Atlantic is considered the most hydrologically stable region for the species, although inner Galicia (NW Spain) shows Mediterranean (unstable) climatic conditions. The Galician region, threatened by past releases of brown trout individuals from central European origin, harbours two native lineages, one of them endemic to the Iberian Peninsula. These populations are thus highly valuable for conservation, as well as being important for recreational fisheries. 2. In total, 546 individuals from 16 sampling sites (15 natural locations from inner Galicia and one from a central European hatchery stock) were genotyped for 11 nuclear markers (10 microsatellite loci and the LDH‐C* locus) to analyse genetic variability, population structure and introgression impact from stocking in order to assess the conservation status of brown trout in the region. Moreover, correlation among hatchery introgression and environmental variables relevant for species population dynamics was also investigated. 3. Genetic variability was within the range of Iberian brown trout (He = 0.500–0.600). Stocking impact was higher than previously reported values for the Atlantic region and was related to environmental instability. Highly significant native population differentiation was observed in the whole region (F_ST = 0.283), at least four main genetic groups being detected across the geographic distribution studied. 4. Conservation strategies at local level (including the creation of genetic refuges and temporal monitoring of genetic composition) are suggested to agencies and administrations for the sustainable management of brown trout.     

Genetic effects of introduced hatchery stocks on indigenous brown trout (Salmo trutta L.) populations in Spain

Abstract– To assess the levels of gene introgression from cultured to wild brown trout populations, four officially stocked locations and four nonstocked locations were sampled for one to three consecutive years and compared to the hatchery strain used for stocking. Allozyme analysis for 25 loci included those previously described as providing allelic markers distinguishing hatchery stocks and native populations. Different levels of hybridization and introgression with hatchery índividuals were detected in stocked drainages as well as in protected locations. These findings indicate that new policies for stocking and monitoring hatchery fish are needed if gene pools of wild Spanish brown trout populations are to be preserved.     

Predation on wild and hatchery salmon by non‐native brown trout (Salmo trutta) in the Trinity River,California

Non‐native predators may interfere with conservation efforts for native species. For example, fisheries managers have recently become concerned that non‐native brown trout may impede efforts to restore native salmon and trout in California's Trinity River. However, the extent of brown trout predation on these species is unknown. We quantified brown trout predation on wild and hatchery‐produced salmon and trout in the Trinity River in 2015. We first estimated the total biomass of prey consumed annually by brown trout using a bioenergetics model and measurements of brown trout growth and abundance over a 64‐km study reach. Then, we used stable isotope analysis and gastric lavage to allocate total consumption to specific prey taxa. Although hatchery‐produced fish are primarily released in the spring, hatchery fish accounted for most of the annual consumption by large, piscivorous brown trout (>40 cm long). In all, the 1579 (95% CI 1,279–1,878) brown trout >20 cm long in the study reach ate 5,930 kg (95% CI 3,800–8,805 kg) of hatchery fish in 2015. Brown trout predation on hatchery fish was ca. 7% of the total biomass released from the hatchery. Brown trout only ate 924 kg (95% CI 60–3,526 kg) of wild fish in 2015, but this was potentially a large proportion of wild salmon production because wild fish were relatively small. As large brown trout rely heavily on hatchery‐produced fish, modifying hatchery practices to minimise predation may enhance survival of hatchery fish and potentially reduce the abundance of predatory brown trout.     

Electrophoretic evaluation of a supplemental stocking programme for brown trout, Salmo trutta L.

Abstract. Over a 15-year period hatchery brown trout, Salmo trutta L., have been added 10 Lower Lough Erne, Northern Ireland to supplement declining native populations. Introductions have mainly comprised eyed ova and fingerlings, stocked into a number of rivers in the Erne drainage. Utilizing a natural genetic tag an electrophoretic assessment of the stocking programme was undertaken. The percentage hatchery genetic contribution in trout populations varied widely from river to river (19%-91%). Lough-caught brown trout (3+ and older) showed a substantial (21·5%) hatchery genetic component. Introgression of native-and hatchery stocks was evident. The resultant deleterious genetic consequences for the conservation of the unique Lough Erne brown trout gene pool arc discussed and alternative management strategies are proposed.     

Importance of life‐history and landscape characteristics for genetic structure and genetic diversity of brown trout (Salmo trutta L.)

Abstract – Investigating the influence of evolutionary forces on the genetic structure and genetic diversity remains a major challenge. Yet, it is of considerable interest for conservation and management of a species. This study investigates the influence of life‐history and landscape features, such as altitude, connectivity and habitat size, on genetic diversity and genetic structure of brown trout (Salmo trutta L.) with stream‐resident, lake‐dwelling and sea‐migrating life‐history in two river systems in northern Sweden. Using regression tree analysis including ecological and landscape characteristics, we show that life history is the most important variable explaining genetic diversity and population differentiation. Sea‐migrating populations show high diversity and low differentiation, and lake‐ and stream‐resident populations show low diversity and high population differentiation, among all samples. No overall genetic correlation with geographical distance was noted; however, among sea‐migrating populations within the River Vindelälven drainage, this pattern was observed. This study illustrates that life‐history and landscape features help to explain genetic structure and genetic variation. The information is important for conservation and management actions, such as fisheries regulations, habitat restorations, stocking of hatchery fish, defining management units and introducing genetic monitoring programmes.     

Benefits of repeated stocking with adult,hatchery‐reared brown trout,Salmo trutta,to recreational fisheries?

Abstract  Recaptures of adult, hatchery-reared, brown trout, Salmo trutta L., and fishing time from anglers were used to evaluate the benefits of stocking programmes with repeated releases of adult brown trout. The recapture rate varied between 17% and 29%. The time between stocking and capture (referred to as residence time) varied between 1 and 160 days (median 3–49 days). Between 67% and 84% of trout caught in the river were recently released fish. Fishing effort increased after stocking, thereby increasing the impact of angling on wild stocks. Stocking with adult brown trout decreases the impact of angling on wild trout only if the time spent fishing by all anglers is kept stable. Furthermore, because of the short residence time of stocked trout, long-term impacts through competition for space and food, or genetic impact through introgression, are limited.     

Survival and growth of hatchery and wild brown trout (Salmo trutta) parr in three Austrian headwater streams

Brown trout populations of three headwater streams in the Northern Limestone Alps of Austria were supplemented by three‐month‐old hatchery‐reared parr from a wild and locally adapted strain and a nonresident domesticated hatchery strain. Growth and survival were monitored with three surveys over a period of 16 months after stocking. Fish descending from the wild reared origin strain demonstrated higher survival rates than the hatchery strain. Differences in growth were found among the investigated streams but not among the investigated strains. The differing temperature regimes of the streams were considered as the primary factor causing those disparities. We conclude that stocking measures had little or no additive effect on successful natural reproduction, as the resident wild brown trout performed significantly better than the stocked fish.     

Foreign brown trout in protected landscapes as a consequence of connectivity

Abstract To preserve the local genetic patrimony of wild aquatic species, population enhancement should be limited to supportive breeding of native lineages and controlled releases in target areas with depleted population size. This is the case of brown trout, Salmo trutta L., in northern Iberia, a species of high economic and cultural value in the region. Employing a combination of conserved and hypervariable genetic markers, illegal stocking of domestic brown trout in protected areas (National Park, Natural Reserve) within the Cantabric chain was detected. The most likely origin of the domestic individuals was determined by assignment tests to hatchery stocks as a result of past releases conducted downstream and out of the protected areas. Where connectivity exists along a water course, the entire drainage should be considered as a single management unit to preserve the native genetic diversity of upstream protected areas.     

Demographic changes following mechanical removal of exotic brown trout in an Intermountain West (USA), high‐elevation stream

Exotic species present a great threat to native fish conservation; however, eradicating exotics is expensive and often impractical. Mechanical removal can be ineffective for eradication, but nonetheless may increase management effectiveness by identifying portions of a watershed that are strong sources of exotics. We used mechanical removal to understand processes driving exotic brown trout (Salmo trutta) populations in the Logan River, Utah. Our goals were to: (i) evaluate the demographic response of brown trout to mechanical removal, (ii) identify sources of brown trout recruitment at a watershed scale and (iii) evaluate whether mechanical removal can reduce brown trout densities. We removed brown trout from 2 km of the Logan River (4174 fish), and 5.6 km of Right Hand Fork (RHF, 15,245 fish), a low‐elevation tributary, using single‐pass electrofishing. We compared fish abundance and size distributions prior to, and after 2 years of mechanical removal. In the Logan River, immigration to the removal reach and high natural variability in fish abundances limited the response to mechanical removal. In contrast, mechanical removal in RHF resulted in a strong recruitment pulse, shifting the size distribution towards smaller fish. These results suggest that, before removal, density‐dependent mortality or emigration of juvenile fish stabilised adult populations and may have provided a source of juveniles to the main stem. Overall, in sites demonstrating strong density‐dependent population regulation, or near sources of exotics, short‐term mechanical removal has limited effects on brown trout populations but may help identify factors governing populations and inform large‐scale management of exotic species.     

Otter (Lutra lutra) predation on stocked brown trout (Salmo trutta) in two Danish lowland rivers

Abstract –  This study aimed to evaluate otter predation on stocked trout. Large hatchery-reared trout (16–30 cm) were stocked into two Danish rivers with different fish populations. Otter diet before and after trout stocking was determined by analysing 685 spraints, collected regularly during the 35-day study period. Fish composition in the rivers before stocking was assessed by electrofishing. In River Trend, a typical trout river, the proportion of trout in the otter diet increased from 8% before stocking to 33% a few days after stocking. Moreover, trout lengths in the diet changed significantly towards the lengths of stocked trout, indicating that newly stocked trout were preferred to wild trout. In River Skals, dominated by cyprinids, there was no change in otter diet after stocking of hatchery trout, i.e., these were ignored by otter. Otter predation should be taken into account together with fish and bird predation, when stocking is used as a measure for conserving endangered salmonid populations.     

Genetic diversity within sea trout population from an intensively stocked southern Baltic river,based on microsatellite DNA analysis

Sea trout, Salmo trutta L., populations in the Slupia River basin have been affected by mass stocking with smolts and fry. This work is focused on a small‐scale differentiation in sea trout populations from one basin with a strong emphasis on the relationship between wild and hatchery origin fish. A total of 216 sea trout were genotyped at 10 microsatellite loci. Global F_ST obtained by amova was low at 0.0165. Pairwise F_ST were significant for all tests except wild and stocked adults. The highest pairwise difference was found between the hatchery sample and Kwacza (F_ST = 0.038). Analysis of the genetic structure revealed micro‐geographical differentiation with four subpopulations. The quality of the artificial spawning was found not to be adequate with a high risk of adverse effects to the whole population. All future stocking actions in the basin should consider the existing population structures.     

Effects of stocking on the genetic structure of brown trout,Salmo trutta,in Central Europe inferred from mitochondrial and nuclear DNA markers

Abstract Stocking has had a considerable effect on wild brown trout, Salmo trutta L., populations throughout Europe. To elucidate this impact and to outline further management strategies, the genetic structure of 25 wild populations and five hatchery stocks from Czech Republic and Slovakia were analysed using mitochondrial (control region) and nuclear DNA (microsatellites, LDH‐C1*) markers. Stocking practices have caused massive hybridisation between the Atlantic and Danube brown trout strains in the central Danube basin and have lead to a loss of among‐population divergence in Slovakia and the eastern part of Czech Republic. Comparison with studies from neighbouring countries revealed substantial differences in haplotype, allele frequencies and genetic diversity across Central Europe. Differences in stocking management and origin of breeding stocks appear to be crucial factors for the spatial variability of the genetic structure of brown trout.     

Stocking of brown trout, Salmo trutta L.: effects of acclimatization

One-year-old hatchery brown trout, Salmo trutta L., ( n  = 16 520) from a sea-run, local brood stock were marked and released (scatter-planted) into the River Laisälven in northern Sweden. Eight different groups were created using Alcian blue and Visible Implant Elastomer tags. Half the fish were kept in small enclosures in four stocking areas for 6 days before release. The other half were released just after transportation. To evaluate the effect of acclimatization on post-stocking performance, the areas were electric fished 2 months later. During the electric fishing survey, a higher number of the acclimatized hatchery fish were recaptured than those released immediately. The growth rate of stocked fish differed significantly between stocking areas and fish held in enclosures grew more than those released directly. The rate of recapture of hatchery fish varied between stocking areas (6.4–17.4%). Movements of juveniles within and between the stocking areas were low, and only 3.6% of the recaptured fish were found in an area not originally stocked. These results showed that acclimatization of fish before release increases the number and size at recapture within a stocking area.     

Parallelism in thermal growth response in otoliths and scales of brown trout (Salmo trutta L.) from alpine lakes independent of genetic background

Low density in natural populations of salmonids has predominantly been managed by stocking of non‐native conspecifics. Due partly to domestication, introduced non‐native fish may be maladapted under natural conditions. Interbreeding between introduced and wild individuals may therefore impair local adaptation and potentially population viability. Brown trout (Salmo trutta L.) from three headwaters (with stocked fish) and three interconnected lakes (with native fish) on the Hardangervidda mountain plateau, southern Norway, were tested for differences in thermal effects on scale and otolith growth. Otolith and scale annuli widths from immature brown trout showed positive correlation with mean annual summer temperature for all six sampled populations. In mature individuals, a similar positive thermal correlation was evident for the otoliths only. Interannuli width measurements from scales indicate a halt in somatic growth for brown trout in this alpine environment when reaching ages between 7 and 9 winters, coinciding with age at maturity. Our study indicates that otolith growth follows summer temperature even when individuals do not respond with somatic growth in these populations and that introduced brown trout and introgressed populations have similar thermal growth responses. Due to the continued otolith growth after stagnation in somatic growth and the impact of fluctuations in summer temperature, the utilisation of otolith annuli widths for back calculation of length at age should be treated with caution.     

Failure of predator conditioning: an experimental study of predator avoidance in brown trout (Salmo trutta)

Many studies have documented that hatchery‐reared salmonids generally have inferior survival after being stocked compared with wild conspecifics, hatchery and wild salmonids have been observed to differ in their antipredator responses. The response of brown trout (Salmo trutta) juveniles (0+) of differing backgrounds to a live predator was compared in two experiments. First, the antipredator behaviour of predator‐naïve hatchery‐reared brown trout and wild‐exposed brown trout were assessed in behavioural trials which lasted for eight days. Second, predator‐naïve and predator‐conditioned hatchery‐reared brown trout were assessed in identical behavioural trials. Brown trout were ‘predator‐conditioned’ by being held in a stream‐water aquarium with adult Atlantic salmon (Salmo salar) and adult brown trout for two days prior to behavioural trials. Predator‐conditioned hatchery‐reared brown trout spent more time in shelters in the trial aquaria than predator‐naïve hatchery‐reared fish, but did not differ in time spent in the predator‐free area. Predator conditioning may account for the increased time spent in the shelter, but does not appear to have affected time spent in the predator‐free area. However, even if significant alteration in behaviour can be noted in the laboratory, the response might not be appropriate in the wild.     

Ecological genetics: introductory note

Abstract– The eight papers resulting from the Ecological Genetics session of the symposium Ecology of Stream Fish: State of the Art and Future Prospects (Luarca, Spain, April 1998) describe the use of molecular genetic markers to investigate questions of population distinctness, interbreeding, and adaptation in five salmonid species. Widely contrasting ecological distinctions described in four species indigenous to western North America [steelhead (rainbow) trout, sockeye salmon, pink salmon and bull trout] reflect past natural and human-induced activities, serving to guide future management and conservation actions through optimizing adaptive opportunities, and preventing genetic losses through localized extinctions. Studies of brown trout contrast widespread introgressions from exogenous hatchery introductions that threaten the integrity of native Spanish populations with a reduced threat based on use of indigenous fish in Portuguese hatcheries. A 9-year study of natural progeny of two genetically distinct Swedish brown trout populations introduced to a previously trout-free area identifies apparent local adaptations as a model to guide other translocations.     

Competition between hatchery‐raised and wild brown trout Salmo trutta in enclosures – do hatchery releases have negative effects on wild populations?

Abstract –  We compared the growth and prey consumption of juvenile hatchery and wild brown trout of similar genetic origin in 12 wire mesh cages in a natural river in North-Eastern Finland. Wild trout started feeding shortly after the start of the experiment, and clearly earlier than novel hatchery trout, but ate less in the presence of hatchery trout. When accompanied with wild trout, novel hatchery trout started to feed earlier, consumed more live prey, and lost less weight than when in allopatry. Hatchery trout grew more slowly than wild trout. To our knowledge, this is the first study to show that hatchery trout benefit from the presence of wild, experienced trout in a complex semi-natural environment. However, our results also indicate that the benefits to hatchery fish are not transferred to wild fish, and ultimately that care should be exercised in management actions when using hatchery trout to supplement wild populations. On the other hand survival potential of the novel hatchery brown trout could be better in the wild when also undomesticated trout are present.