Stocking of native Mediterranean brown trout (Salmo trutta) into French tributaries of Lake Geneva does not contribute to lake-migratory spawners

Abstract –  We studied the contribution of stocking with Mediterranean autochthonous brown trout ( Salmo trutta ) to the lake-migratory spawners in French tributaries of Lake Geneva. This singular lake system is inhabited by both the resident and the migratory life-history form, and by trout of Atlantic and Mediterranean origins. Using two discriminatory microsatellite loci between the Atlantic and Mediterranean alleles, we compared the genetic composition of the migratory Atlantic populations of the four main tributaries before (1999; N  = 113) and after (2004, 2005 and 2006; N  = 137) recent introductions of native Mediterranean trout. No significant modification of the genetic composition was detected in all four populations over time and only two individuals were statistically assigned to the Mediterranean strain by the Bayesian method implemented in the structure program. These results suggest a negligible contribution of the stockings with autochthonous Mediterranean trout to the lake-migratory spawners migrating into the four tributaries.     

Can female preference drive intraspecific diversity dynamics in brown trout (Salmo trutta,L.)?

Intraspecific diversity can be strongly impacted by the dynamics of reproductive isolation during secondary contacts. The high levels of hybridisation and introgression between Atlantic and Mediterranean lineages in contemporary populations of brown trout in the Northern part of the French Alps are a good case in point. After a long period of allopatry, which one assumes has facilitated their divergent phenotypic and genetic evolution, man has removed a geographical reproductive barrier via stocking the Atlantic lineage in the Mediterranean area, thereby potentially enabling gene flow between native and non native populations. We investigate how much a prezygotic reproductive behaviour, that is female preference for male phenotype, can influence hybridisation in natural environments, using subaquatic video recording of reproduction. Our statistical model indicates that female preference appears to be largely heterogamous: females tend to select dissimilar males with respect to their own phenotype; thus, rather than acting as a barrier to gene flow, female preference is favouring gene flow between lineages that have been artificially placed in sympatry. This finding based on observational data is in agreement with previously reported genetic data, high levels of hybridisation between the lineages. We suggest that a knowledge of reproductive behaviour in natural environments is an essential tool for biodiversity managers to assess the potential risks associated with the introduction into recipient populations of non‐native lineage fish at the intraspecific level.     

Investigating the genetic structure of trout from the Garden of Ninfa (central Italy): Suggestions for conservation and management

Mediterranean populations of brown trout (Salmo trutta L. complex) have lost a large part of their genetic distinctiveness, mostly due to massive restocking, and the waters of the Gardens of Ninfa (province of Latina, central Italy, Site of Community Importance since 2013) are regarded as one of a few potential reservoirs of autochthonous trout lineages in the Tyrrhenian drainage of the Italian peninsula. In this study, nuclear and mitochondrial markers were used on brown trout samples from Ninfa to estimate non‐Mediterranean influence in the population gene pool, potential changes of genetic structure over time and genetic relationships with other sites known (or suspected) to host native trout gene pools. Striking changes in both microsatellite and mtDNA allele frequencies over a 9‐year time span were found and provided evidence of unrecorded stocking from the nearby Lake Fibreno. Results are analysed in the light of potential ecological consequences of such events on a longer time scale and provide a scientific background for fisheries management and conservation programmes in the area.     

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 refuges for a self‐sustained fishery: experience in wild brown trout populations in the eastern Pyrenees

Abstract – Management policies balancing harvest and conservation of natural populations of fish are difficult to establish, both scientifically and politically. This issue is particularly difficult when those populations represent native genetic resources. Since 1997, several brown trout populations in the eastern Pyrenees Mountains (Spain) were designated as ‘genetic refuges’ under varying fishing regulations, where releases of hatchery‐origin fish are not permitted. We analysed genetic variation in samples of brown trout from six of those refuge populations and four non‐refuge populations within the same region. Each population was sampled in four separate years: 1993, 1999, 2004 and 2006. Our analyses were based on a diagnostic allele (LDH‐C*90) that distinguishes native and exogenous hatchery populations. Comparisons were based on stocking histories before and after refuge designations and on three management strategies: fished, unfished and catch‐and‐release. Overall, we detected significant genetic introgression resulting from past stocking practices despite the current restriction of hatchery releases imposed by the recent genetic refuge policy. However, this new policy has prevented detectable introgression from increasing throughout the region and together with additional measures on length and number of captured fish is contributing to self‐sustained fisheries that are achieving conservation goals. Quick acceptance of ‘genetic refuges’ by anglers in one particular river, the Ter River basin, has been a key factor in protecting native gene pools compared with the Segre River basins where refuges were not readily accepted.     

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.     

Genetic and morphological divergences between wild and captive‐bred populations of Salmo trutta abanticus

Salmo trutta abanticus is a non‐anadromous trout species native to Lake Abant and Seven Lakes in Turkey. A restocking programme by captive breeding was initiated in 1999 to support S. trutta abanticus population. Reared 2‐year‐old juveniles from randomly caught wild parental individuals in Maçka breeding farm were introduced into Lake Abant. We aimed to compare genetic and morphological divergences between wild‐ and captive‐bred populations using seven microsatellite loci and geometric morphometric measurements. A significant genetic and morphological divergences were detected between all population in F_st and canonical variate analysis based on geometric morphometric with 10 homolog landmark. Eighty‐six microsatellites alleles were recorded across loci. Number of private alleles, observed alleles and observed heterozygosity are statistically significant higher in Maçka captive‐bred population than Lake Abant and Seven Lakes populations. Of 42 tests, three departures from Hardy–Weinberg equilibrium were detected in all populations after Bonferroni correction. Two pairs of loci (Ssa85 – Str73 and Str73‐Str543) in Maçka, one pairs of loci (Ssa85‐Str73) in Abant and two pairs of loci (Ssa85‐Str60 and Str73‐Str543) in Seven Lakes populations show linkage disequilibrium. Population structure analysed with Structure software showed three genetic groups (?K = 3) in our studied populations. Relatedness estimates show higher mean relatedness values (r = 0.220 ± 0.230) for Maçka captive‐breed population than wild populations of Abant Lake and Seven Lakes (r = 0.140 ± 0.210 and r = 0.170 ± 0.200 respectively).     

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.     

Microsatellite markers reveal population structure in gilthead sea bream Sparus auratus from the Atlantic Ocean and Mediterranean Sea

ABSTRACT:   This study was carried out to investigate the genetic variability and the population genetics of Sparus auratus. In fact, despite its importance in Mediterranean fisheries and aquaculture, very little it is known concerning its population structure. Samples of wild gilthead sea bream were collected in seven different localities along the Mediterranean and Atlantic coasts and were genetically characterized by means of microsatellite markers. Genotyping at four microsatellite loci revealed high polymorphism (7–38 alleles/locus) and expected heterozygosities, which ranged from 0.80 to 0.85. A slight but significant population structure was found ( F _ST = 0.010). In fact, at least three populations of gilthead sea bream within the Western Mediterranean Sea were identified (Sardinian Sea, Sardinian Channel and Central Tyrrhenian Sea), which are also genetically differentiated from those of the Atlantic Ocean and the Adriatic Sea.     

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.     

Genetic characterization of naturalized populations of brown trout Salmo trutta L. in southern Chile using allozyme and microsatellite markers

This study describes the genetic structure of five naturalized populations of brown trout in southern Chile using allozyme and microsatellite markers to establish levels of intra‐ and interpopulation genetic variability and divergence. Fourteen enzymatic systems were used comprising 20 loci and three microsatellite loci specific to brown trout. The genetic variability values (allozymes, P=20–35%, average=27%, H_O=0.118–0.160, average=0.141; microsatellites, P=33.3–100%, average=66.66%, H_O=0.202–0.274, average=0.229) are similar to values described in other naturalized populations of brown trout present in Chile, but higher than those observed in European populations of this species. Values of total genetic diversity (H_T) (allozymes=0.1216 and microsatellites=0.3504) and relative genetic divergence (G_ST) (allozymes=9.5% and microsatellites=15%) were also similar to the results obtained in previous studies of Chilean populations of brown trout. These values, when compared with those obtained in Europe, proved to be similar for H_T but lower for G_ST. The low interpopulational genetic differentiation was in accordance with the small genetic distance observed between the populations analysed (D Nei=0.004–0.025). On the other hand, the high frequency of one of the two alternative alleles of the phylogeographic marker locus LDH‐5* in the populations analysed (LDH‐5*90>0.84) would indicate a European origin, in particular Atlantic as opposed to Mediterranean, for the brown trout introduced into Chile. The high levels of genetic variability suggest a mixed origin for the naturalized brown trout in Chile, which could have originated either before or during the introduction process. Nevertheless, the low level of genetic differentiation between populations could reflect the short lapse of time in evolutionary terms, during which populations introduced into Chile have been exposed to different evolutionary forces, and which has not been sufficiently long to produce greater genetic differentiation between populations.     

The effect of competition among three salmonids on dominance and growth during the juvenile life stage

Although non‐native species can sometimes threaten the value of ecosystem services, their presence can contribute to the benefits derived from the environment. In the Great Lakes, non‐native brown trout (Salmo trutta) and rainbow trout (Oncorhynchus mykiss) support substantial recreational fisheries. With current efforts underway to restore once‐native Atlantic salmon (Salmo salar) to Lake Ontario, there is some concern that Atlantic salmon will impede non‐native contributions to the recreational fishery because Atlantic salmon exhibit niche overlap with brown trout and rainbow trout, particularly during the juvenile life stage. We therefore examined competition and growth of juvenile Atlantic salmon, brown trout and rainbow trout in semi‐natural streams. We found that brown trout were the most dominant and had the greatest growth rate regardless of what other species were present. Rainbow trout were more dominant than Atlantic salmon and consumed the most food of the three species. However, in the presence of brown trout, rainbow trout fed less frequently and exhibited negative growth as compared to when the rainbow trout were present with only Atlantic salmon. These data suggest that, outside of density‐dependent effects, Atlantic salmon will not impact stream production of brown trout and rainbow trout.     

Competitive interactions among multiple non‐native salmonids and two populations of Atlantic salmon

Competitive interactions with non‐native species can have negative impacts on the conservation of native species, resulting in chronic stress and reduced survival. Here, juvenile Atlantic salmon (Salmo salar) from two allopatric populations (Sebago and LaHave) that are being used for reintroduction into Lake Ontario were placed into semi‐natural stream tanks with four non‐native salmonid competitors that are established in Ontario streams: brown trout (S. trutta), rainbow trout (Oncorhynchus mykiss), Chinook salmon (O. tshawytscha) and coho salmon (O. kisutch). Brown trout and rainbow trout reduced the survival and fitness‐related traits of Atlantic salmon, whereas Chinook salmon and coho salmon had no impact on these traits. These data support theories on ecological niche overlap and link differences in observed aggression levels with competitive outcomes. Measurements of circulating hormones indicated that the Atlantic salmon were not chronically stressed nor had a change in social status at the 10‐month time point in the semi‐natural stream tanks. Additionally, the Sebago population was better able to coexist with the non‐native salmonids than the LaHave population. Certain populations of Atlantic salmon may thus be more suitable for some environments of the juvenile stream phase for the reintroduction into Lake Ontario.     

Allozyme diversity in Australian rainbow trout, Oncorhynchus mykiss

Rainbow trout, Oncorhynchus mykiss (Walbaum), were first introduced into Australia over 100 years ago, and forms the basis of important recreational inland fisheries and an aquaculture industry in south‐eastern Australia. This paper investigates the genetic variation within and between samples of Australian rainbow trout using allozyme electrophoresis. The levels of genetic diversity within Australia do not show marked differences from those observed in hatchery and wild populations from throughout North America, New Zealand and South Africa, but there is evidence for the loss of some rare alleles during translocation from California to Australia via New Zealand. No appreciable difference in genetic diversity was apparent between hatchery and self‐sustaining wild populations of rainbow trout from mainland Australia. However, significant differences in allelic frequencies were observed, with consistent genetic differences between Victorian and New South Wales samples most likely reflecting state‐based hatchery and stocking policies.     

Competitive effects between rainbow trout and Atlantic salmon in natural and artificial streams

Competition with non‐native species may impede the restoration of native species, but differences in competitive abilities among intraspecific native populations may make some populations more suitable for reintroduction than others. Here, juvenile Atlantic salmon (Salmo salar) from two allopatric populations (LaHave and Sebago) being used for reintroduction into Lake Ontario were placed into two natural stream sites differing in the presence of ecologically similar rainbow trout (Oncorhynchus mykiss). We assessed the effects of competition in the natural streams on fitness‐related traits and habitat use of the Atlantic salmon. We then compared these effects to those observed in artificial streams from a previous study. Atlantic salmon in natural streams had reduced fitness‐related traits that were associated with suboptimal microhabitats in the presence of rainbow trout, but utilised optimal microhabitats in their absence. In the presence of rainbow trout, the two Atlantic salmon populations exhibited comparable recapture proportions to each other, but the individuals from the Sebago population had better performance (body size and condition) than those from the LaHave population. Responses of both Atlantic salmon populations to competition with rainbow trout were generally similar in both direction and magnitude when compared to results from the artificial stream study. The combined results suggest that native species restoration efforts should be focused on candidate populations that are ecologically suitable to reintroduction environments, as well as on suitable habitats that do not contain exotic competitors. Moreover, this study highlights the value of controlled experiments in artificial environments for predicting fitness‐related performance in natural environments.     

Microsatellite diversity assessment of brown trout (Salmo trutta) population structure indicate limited genetic impact of stocking in a Norwegian alpine lake

Abstract – Releases of non-native trout often result in introgression into natural populations and negative genetic effects. The causal ecological mechanisms for a wide range of reported outcomes are poorly understood. Brown trout population structure in an alpine lake with three major recruitment streams was assessed by analysis for eight DNA microsatellite markers and compared with the non-native population. The lake is subject to a 40-year recorded history of stocking with exogenous trout. No certain deviations from Hardy–Weinberg equilibrium were found. Tests for population differentiation and genetic distance indicated separate populations for all the sampled areas, and with the exogenous population as a cluster quite different from the others. Assignment tests indicated that only a small fraction of the fish sampled from the lake originated from the introduced trout strain (<3%). Wild discriminate, naturally reproducing populations characterize this alpine lake ecosystem, in spite of 40 years of stocking, which appears to have had a limited impact. It is unlikely that this population structure can ultimately be explained by trout movement patterns. Genetic analysis needs to be supplemented by studies of local life history strategies, to evaluate the relative importance of local adaptation versus random genetic differentiation, because implications for conservation and management are different.     

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.     

Assessing the long-term genetic impact of historical stocking events on contemporary populations of Atlantic salmon, Salmo salar

Abstract  Supplementation of wild fish with non-native or domesticated fish is common practice. However, these stocked and native fish differ both ecologically and genetically and, in the wild, they interact in a multitude of ways, often with negative repercussions for the native population. This study assessed the long-term genetic impact of historical stocking activities on a contemporary population of Atlantic salmon, Salmo salar L. During the 1960s salmon from hatcheries in Scotland and Iceland were transplanted to the River Dart, England. Microsatellite loci were used to assess the current level of population admixture between samples taken from the source location of the stocked fish during the 1960s and contemporary Dart populations. After allowances were made for natural genetic relationships between donor and recipient populations, the long-term impact of the historical stocking events on a catchment scale appears minimal. However, one tributary consistently reflected closer genetic relationships with the donor populations, indicating a possible long-term impact on a localised scale.     

The relationship between the snowmelt flood and the establishment of non‐native brown trout (Salmo trutta) in streams of the Chitose River,Hokkaido, northern Japan

Flow regime is one of the major determinants of establishment success for non‐native aquatic organisms. Here, we examine the influence of flow variability associated with snowmelt flood on the establishment success of non‐native brown trout in 10 streams in northern Japan. We regarded the presence of Age‐0 brown trout as the index of the successful establishment. The emergence of Age‐0 brown trout in our study region begins in May, a time that overlaps with the occurrence of snowmelt flood. The presence of Age‐0 brown trout was negatively associated with flow variability, and it was also negatively associated with summer water temperature. Our results indicate that the non‐native brown trout tends to establish in the streams with smaller snowmelt floods and lower summer water temperatures. Brown trout is an invasive, non‐native species that is problematic all over the world, and effective management strategies for preventing their further expansion are urgently needed. This study suggests that river managers should recognise that stable streams such as spring‐fed streams (i.e., low flow and summer water temperature) and flow‐regulated streams, have a higher potential risk of brown trout invasion.     

Genetic effects of selection on polygenic traits with examples from Atlantic salmon, Salmo salar L.

Abstract. Recent attempts to apply selective breeding strategies in fish populations have focused on the genetic effects of selection. The basic mechanisms of long-term response to selection are discussed with reference to experience from laboratory and farm animal populations. A simplified polygenic model is presented, and it is shown that considerable response may be obtained for polygenic traits without essential changes in allele frequencies and genetic variability, and that the probability of fixation of alleles or genotypes because of selection will be low when the trait is affected by a larger number of loci. The span between the extreme fixed genotypes for such traits may easily become wider than that which may be realized under normal biological conditions. This is illustrated with some examples from Atlantic salmon. Salmo salar L. The effects of divergent fitness selection of populations on genetic differentiation and allele frequencies are also considered, and some examples from experiments with Norwegian river strains of Atlantic salmon are discussed. It is shown that the river strains are not expected to differ much in frequencies of fitness alleles if the number of loci affecting the fitness trait is large, and that considerable overlapping of genetic values for fitness between strains may be expected.