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.     

High genetic introgression in alpine brown trout (Salmo trutta L.) populations from Hardangervidda, Norway

Abstract –  We studied the effect of stocking brown trout in three alpine lakes (Skavatn, Midtre Grøndalsvatn and Nordmannslågen) of the Hardangervidda mountain plateau by comparing microsatellite data from historical (1933 and 1967) and contemporary (2003) population samples. Historical and contemporary samples from a control lake (Krokavatn) and a contemporary sample from Gjuvsjå (suspected donor lake) were included as well. A total of 331 brown trout were genotyped with 11 microsatellites. Very low level of genetic diversity is observed in the historical samples from Nordmannslågen and Midtre Grøndalsvatn but not in the contemporary samples. In Skavatn historical and contemporary samples show similar genetic diversity as in the other contemporary samples. Admixture analyses with the structure program indicate high levels of introgression by nonnative brown trout in all three lakes. The stocking source for two lakes (i.e. from Gjuvsjå) is partially verifiable via our analyses. The high introgression in these lakes is discussed and the management issues are briefly addressed.     

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.     

Do dams increase genetic diversity in brown trout (Salmo trutta)? Microgeographic differentiation in a fragmented river

Abstract – Local genetic differentiation may potentially arise in recently fragmented populations. Brown trout is a polytypic species exhibiting substantial genetic differentiation, which may evolve in few generations. Movement (semi‐)barriers in rivers may cause fragmentation, isolation and genetic differentiation in fish. In the Måna River (28 km) flowing from the alpine Lake Møsvatn to the boreal Lake Tinnsjø, construction of four hydropower dams during the period 1906–1957 have fragmented the previously (since last Ice Age) continuous wild resident brown trout population. Samples from the two lakes (N = 40) and six sites in the river (N = 30) isolated at different times were analysed at nine microsatellite loci. All populations showed substantial genetic variation (mean number of alleles per locus 5.3–8.9, observed heterozygosity 0.57–0.65 per population, overall F_st = 0.032). Pairwise multilocus F_st estimates indicated no significant differentiation between populations in the two lakes, and no or little differentiation in the lower river (F_st = 0.0035–0.0091). The microgeographic differentiation among wild resident trout at these sites was less than expected based on similar previous studies. However, results from the upper river, in particular the site immediately below the Lake Møsvatn outlet and dam, indicated isolation (F_st > 0.035). Calculation of genetic distances and assignment tests corroborated these results, as did a significant correlation between years of isolation (since dam construction) and F_st. The population structuring is most likely a result of fragmentation by dams, which has increased overall genetic diversity. This increased local differentiation may be caused by natural selection, but more likely by genetic drift in small, recently fragmented populations. Increased local genetic diversity by genetic drift does not justify conservation measures aiming at preserving genetic diversity.     

Recaptures and resource use of native and non-native brown trout, Salmo trutta L., released in a Norwegian lake

Abstract Rate of recapture (gill netting), habitat use, and diet of three strains of stocked brown trout, Salmo trutta L., were compared with resident brown trout in a Norwegian lake. The strains originated from an alpine lake, from a boreal lake, and from the native brown trout population in the lake. Overall recapture rate was 5–8% for all strains. The low recapture rate could be due to the relatively small size at stocking; mean fish length varied between 13.1 and 14.5 cm with strain and stocking method. Two years after release, the frequency of the different strains decreased from about 12% in the first year to stabilize at about 1%. The alpine strain showed the highest overall recapture rate, whereas the native strain was recaptured at an intermediate rate. The overall recapture rate of scatter-planted brown trout was higher than that of spot-planted brown trout. Immediately after being stocked, introduced fish ate less and had a less-varied diet than resident trout; however, stocked fish adopted a natural diet within the first summer. The distribution of trout between the pelagic and the upper epibenthic habitat was similar for both the resident and the stocked brown trout. Results indicate that the habitat use of stocked brown trout is adaptive and becomes similar to that of indigenous fish.     

Genetic persistence of an initially introduced brown trout (Salmo trutta L.) population despite restocking of foreign conspecifics

The persistence of the initial genetic structure despite later stocking with foreign conspecifics is influenced by several factors, leading to different levels of introgression. Stream spawning has been assumed to be the prevailing recruitment strategy of brown trout (Salmo trutta L.). However, in lakes with limited stream spawning habitat, but still with high natural recruitment, lake spawning has been proposed. Using fourteen microsatellites, we assessed the genetic structure in a small allopatric brown trout population in the Lake Skavatn, located on the Hardangervidda mountain plateau, Norway. A total of 265 brown trout were obtained from the lake itself, the outlet stream, two tributaries, a littoral sample from a presumed lake spawning site, a suggested stocking source, and a lake sample from 1967, representing the initial lake population. The six Skavatn samples were best represented by three genetic components, showing a shift from the initial population, but with no genetic signal from the suggested stocking source. The littoral sample had the largest similarity to the initial lake population, possibly indicating the importance of lake spawning in a system where streams offer unpredictable spawning and rearing conditions. Due to large annual variations in recruitment contributions from the different spawning and rearing locations, the genetic structure of the lake population probably vary over time.     

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.     

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 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.     

Effects of stocked trout on native fish communities in boreal foothills lakes

Nasmith LE, Tonn WM, Paszkowski CA, Scrimgeour GJ. Effects of stocked trout on native fish communities in boreal foothills lakes.
Ecology of Freshwater Fish 2010: 19: 279–289. © 2010 John Wiley & Sons A/S Abstract – Ecological effects of stocking nonnative trout into lakes are receiving increased attention, especially in alpine environments. We assessed effects of stocked trout on native forage fishes in the boreal foothills of Alberta (Canada) by comparing fish density, population size structure and spatial and temporal activities in stocked and unstocked lakes over 3 years (2005–2007). The numerically dominant dace (primarily Phoxinus spp.) were larger in stocked lakes, consistent with size‐limited predation. Dace were also more crepuscular and concentrated on the lake‐bottom in stocked lakes, compared to more daytime activity in the water column in unstocked lakes. There were, however, no demonstrable effects of trout on the abundance of forage fish. The lack of major population‐level impacts of stocked trout suggests that current stocking practices, characteristics of boreal foothill lakes (e.g. thermal structure, abundant invertebrates, dense macrophytes) and/or behavioural adjustments of forage fish contribute to healthy native fish populations in our stocked lakes.     

Population characteristics of channel catfish near the northern edge of their distribution: implications for management

Channel catfish, Ictalurus punctatus (Rafinesque), populations in six lakes in northern Idaho, USA, were sampled to describe their population characteristics. During the summers of 2011 and 2012, 4864 channel catfish were sampled. Channel catfish populations had low to moderate catch rates, and length structure was dominated by fish <400 mm. Channel catfish were in good body condition. All populations were maintained by stocking age‐1 or age‐2 fish. Growth of fish reared in thermally enriched environments prior to stocking was fast compared to other North American channel catfish populations. After stocking, growth of channel catfish declined rapidly. Once stocked, cold water temperatures, prey resources and (or) genetic capabilities limited growth. Total annual mortality of age 2 and older channel catfish was generally <40%. Tag returns indicated that angler exploitation was low, varying from 0 to 43% among lakes. This research provides insight on factors regulating channel catfish population dynamics and highlights important considerations associated with their ecology and management.     

Baseline demographics of a non‐native lake trout population and inferences for suppression from sensitivity‐elasticity analyses

Management agencies in several western states of the United States are implementing suppression programmes to control non‐native lake trout, Salvelinus namaycush (Walbaum), for the conservation of native species. This study was implemented to ascertain the population demographics of an expanding lake trout population and use those data to construct an age‐structured model to inform suppression efforts. Population projection matrices were used to model population growth and identify age or stage classes with the greatest influence on population growth. The size and age structure of lake trout sampled was skewed towards juveniles, indicating strong recruitment and a growing population. Matrix‐model simulations corroborated the observed size and age structure, as the lake trout population was predicted to grow exponentially (λ = 1.35, 95% CL: 1.25–1.43) with no suppression efforts. Elasticity analysis of matrix models indicated the relative contribution of survival rates to population growth among immature age classes was equal from age 0 to age at first maturity, but immature survival rates contributed more than adult survival and fertility rates. These results emphasise the importance of targeting juvenile lake trout for suppression efforts during exponential growth in recently established populations.     

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.     

基于线粒体控制区的江苏湖鲚群体遗传多样性和遗传结构分析#br#

为掌握江苏省重要湖泊湖鲚(Coilia nasus taihuensis)群体的遗传多样性和遗传结构,利用线粒体控制区(D-loop)全序列分析了6个湖泊(太湖、滆湖、高邮湖、白马湖、洪泽湖和骆马湖)湖鲚野生群体的遗传多样性水平和群体分化情况。结果表明,6个群体共214尾样本的D-loop序列中,共发现103个变异位点,92种单倍型。6个群体的单倍型多样性为0.726~0.951,核苷酸多样性为0.00552~0.01036,6个群体整体的单倍型和核苷酸多样性分别为0.857和0.00729,表明湖鲚群体的遗传多样性较高,且符合高单倍型多样性和低核苷酸多样性特点。分子方差分析(AMOVA)结果表明,群体间变异百分比为6.20%,群体内变异百分比为93.80%,说明遗传变异主要来自群体内部。群体总的遗传分化系数(F_st)为0.06199(P<0.01),两两群体间的F_st显示,滆湖群体与其他群体间存在极显著的遗传分化(P<0.001),而其他群体间无显著分化(P>0.05)。单倍型分子系统进化树和网络进化图显示,6个群体的单倍型形成了2个谱系,但谱系组成与群体地理分布无相关性。中性检验分析结果显示,湖鲚群体进化过程中经历过种群扩张,扩张时间大约发生在0.089~0.160百万年前。研究结果表明,湖鲚群体具有较高的遗传多样性,滆湖群体与其他群体具有极显著的遗传分化,且拥有多个独享单倍型,应将滆湖群体单独作为一个管理单位,其他5个群体作为一个整体进行管理和利用。     

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.     

Genetic structure of brown trout,Salmo trutta,populations from differently sized tributaries of Lake Mjøsa in south‐east Norway

This study was based on genotyping eight microsatellite loci of 463 brown trout, Salmo trutta L., sampled in nine differently sized tributaries in three areas on the eastern shore of Lake Mjøsa, south‐eastern Norway. The populations were genetically structured, and Mantel's test showed that genetic distance correlated positively with geographical distance. Temporal differentiation F_ST over a 2‐year period was estimated in five streams and was non‐significant after Bonferroni correction. Effective population size N_e was positively correlated with the habitat length available from the lake (0.3–22 km) and negatively with the number of full sib pairs in the sample. There was no correlation between N_e and genetic diversity, and private alleles were recorded in three medium‐sized streams, but not in the largest two. The importance of small spawning and nursery streams for the maintenance of genetic diversity of brown trout was demonstrated.     

A genetic comparison of sympatric anadromous and resident Atlantic salmon

Many studies have identified the importance of local adaptation in Atlantic salmon (Salmo salar) and the strong genetic differences that exist between allopatric or parapatric resident and anadromous populations. However, as truly sympatric migratory phenotypes of Atlantic salmon have not been studied, it remains unclear whether distinct genotypes previously associated with life history differences are maintained through reproductive isolation and subsequent genetic drift or through natural selection induced by different life history requirements. In this study, sympatric anadromous and resident Atlantic salmon were sampled from three Newfoundland (Canada) watersheds to evaluate the genetic divergence of these life history forms. Eight microsatellite loci were used to quantify genetic variation within and among populations. Metrics of genetic differentiation (exact tests for population differentiation, pairwise θ values) provide no evidence of genetic differentiation between some sympatric anadromous and resident phenotypes within a system with no history of segregation. In the remaining two watersheds, the observed differentiation appears to be a consequence of historical segregation rather than life history form. Nonetheless, these differences have been maintained in contemporary times for several generations. At broader spatial scales, resident salmon were more genetically divergent from one another than anadromous life history forms and were more closely related to anadromous salmon from within their watershed than to resident salmon from other watersheds. The study indicates that both life history forms can be maintained within a single population, but that sympatric populations of different life histories can maintain genetic differences for at least several generations after being reconnected.     

Long-term variation in brown trout, Salmo trutta L., stocking success in a large lake: interplay between availability of suitable prey and size at release

Abstract –  Factors affecting long-term variation in brown trout, Salmo trutta L., stocking success were examined in a large lake, Lake Oulujärvi, in central Finland. Brown trout were stocked in spring (late May to early June) in 1974–1991 and in summer (late June to early July) in 1992–2001. The biomass of the vendace, Coregonus albula (L.), population (prey) at release time had the largest positive effect on stocking success within both periods: biomass of adult vendace in spring and both 0+ and adult vendace in summer. Increasing the size of stocked fish had a positive effect if the vendace available at release were only adults. The increasing trend of predator-catch-per-unit-effort (CPUE) [combined CPUE of northern pike Esox lucius L., burbot Lota lota (L.), and pike-perch Stizostedion lucioperca (L.)] through the study period and its negative effect on trout stocking success suggested an increasing effect of predation within the entire time series.     

Response of resident brown trout, Salmo trutta L., and rainbow trout, Oncorhynchus mykiss (Walbaum), to the stocking of hatchery-reared brown trout

Two strains of hatchery-reared adult brown trout, Salmo trutta L., [208–334 mm total length (TL); n =  591] were individually marked and released into a limestone stream. The estimated survival after one month (86%; n =  508) was comparable to that for resident brown trout and rainbow trout, Oncorhynchus mykiss (Walbaum), (89%; n =  771), but declined to 14% ( n =  83) after 8 months compared with 52% ( n =  451) for resident trout. The movement of resident trout out of stocked stretches was higher (14%) than from control sites (5%), but the population size in both individual sites and the overall study area were unaffected. The growth of resident brown trout was unaffected by stocking, but rainbow trout showed lower growth rates in stocked versus unstocked stretches both one and 8 months after stocking ( P <  0.002).