Vertical distribution and substrate preference of brown trout in a littoral zone

Synopsis We studied vertical distribution, substrate preference and food choice of brown trout, Salmo trutta, from benthic gillnet catches at four littoral sampling locations in a Norwegian hydroelectric reservoir. The sampling locations had different bottom substrates; at one location the bottom substrate consisted of sand, while at the other three, substrates consisted of stones ranging 2–5 cm, 10–30 cm and 30–150 cm in diameter, respectively. Small-sized (< 160 cm) and intermediate-size (164–269 mm) brown trout were mainly caught close to the bottom (0–0.5 m above). Small-sized brown trout were caught in the highest frequency at the location with substrate consisting of 10–30 cm large stones. Intermediate-sized brown trout were also caught in highest frequency at this location, but were also caught in a high frequency at the location with sandy substrate. In contrast, the catches of large-sized ( 270 mm) brown trout did not vary with distance from the bottom or with substrate coarseness. The most important food items for the brown trout were aquatic insects, surface insects, Eurycercus lamellatus and crustacean zooplankton, mainly Daphnia longispina, Bythotrephes longimanus, and Holopedium gibberum. In accordance with the differences in vertical distribution, benthic food was more important to small than to large brown trout. We argue that small brown trout stayed close to the bottom to reduce aggressive behaviour from larger specimens, and that small brown trout were therefore more dependent on benthic food items. We also argue that the observed differences in substrate preference between the size groups of brown trout is explained by variation in access to shelter, visual isolation between individuals and benthic feeding conditions between locations.     

Food borne parasites as indicators of trophic segregation between Arctic charr and brown trout

The habitat and diet choice and the infection (prevalence and abundance) of trophically transmitted parasites were compared in Arctic charr and brown trout living sympatrically in two lakes in northern Norway. Arctic charr were found in all main lake habitats, whereas the brown trout were almost exclusively found in the littoral zone. In both lakes the parasite fauna reflected the niche segregation between trout and charr. Surface insects were most common in the diet of trout, but transmit few parasites, and accordingly the brown trout had a relatively low diversity and abundance of parasites. Parasites transmitted by benthic prey such as Gammarus and insect larva, were common in both salmonid host species. Copepod transmitted parasites were much more common in Arctic charr, as brown trout did not include zooplankton in their diets. Parasite species that may use small fish as transport hosts, were far more abundant in piscivorous fish, especially brown trout. The seasonal dynamics in parasite infection were also consistent with the developments in the diet throughout the year. The study demonstrates that the structure of parasite communities of charr and the trout is highly dependent on shifts in habitat and diet of their hosts both on an annual base and through the ontogeny, in addition to the observed niche segregation between the two salmonid species.     

A Bioenergetic Analysis of Factors Limiting Brown Trout Growth in an Ozark Tailwater River

We examined prey utilization and energy consumption by brown trout, Salmo trutta, in a cold tailwater (Little Red River, Arkansas, USA; LRR) having low biodiversity and low availability of fish as prey. Stomach content analysis and age estimation were performed on thirty brown trout (10 each of three size classes for a total of 710 trout) collected monthly from an upstream and downstream site over a 1-year period. Diet diversity was low at both sites, as 80% and 70% of all prey consumed by upstream and downstream brown trout, respectively, were isopods. Piscivory (<0.5% of individuals sampled) and consumption of terrestrial invertebrates were rare. There was no relation between diet diversity and trout age, and a very small ontogenetic shift in brown trout diet. Second, we investigated brown trout growth rates relative to prey consumption and temperature. Temperatures and availability of prey were less than required for maximal trout growth. However, prey availability limited trout growth directly, but sub-optimal temperatures probably buffered the effect of this reduced energy consumption by reducing metabolic energy expenditures. Brown trout growth was 54.8–57.0% of the maximum predicted by a bioenergetics model. Instantaneous growth rates for age 1 and adult brown trout were slightly higher for those downstream (0.195) versus those upstream (0.152). Although isopods are abundant within this tailwater to serve as a forage base, the displacement of native fish fauna and subsequent lack of establishment of cold-tolerant forage fish species due to the thermal regime of hypolimnetic release from Greers Ferry Reservoir probably serves as a major barrier to brown trout growth.     

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.     

The selection of the ‘wild’: A combined molecular approach for the identification of pure indigenous fish from hybridised populations

The identification of pure indigenous fish from hybridised populations represents a key issue in fisheries management and conservation biology. In the present study an approach for selection of purebred marble trout (Salmo trutta marmoratus C.) individuals out of admixed populations was set up and assessed. In a first step, baseline data sets of pure marble trout and pure brown trout specimens based on twelve microsatellite loci were used to simulate five consecutive generations of admixture. The baseline and the resulting simulation data sets were then combined with data of a ‘real’ hybridised marble trout population to perform a single individual assignment test as implemented in STRUCTURE. By this procedure the assignment approach was calibrated and it was possible to compare admixture coefficients obtained for individuals from different populations. The ranking of individual admixture coefficients on a plot and comparison with simulated data revealed that the test population was composed of pure marble trout individuals, first generation hybrids between marble trout and brown trout, and hybrid backcross specimens between both groups. However, by defining a critical q-value of 0.1 and additionally integrating individual sequence data of the mtDNA control region, it was possible to indicate individuals, which could be selected for the establishment of a pure marble trout strain.     

Sacciform cells in the epidermis of the brown trout,Salmo trutta,and the Arctic char,Salvetinus alpinus

Summary Sacciform cells containing an acidophilic, proteinaceous secretion, were identified in the epidermis of the brown trout and Arctic char. This cell type increased in number during the chronic stages of infestation by the ectoparasitic flagellate, Ichthyobodo sp., in immature brown trout, and decreased during sexual maturation in male brown trout and char. It is suggested that the salmonid sacciform cell produces a secretion which protects the fish against infestation or damage by skin parasites.     

Mercury accumulation in five species of freshwater fish in Lake Tyrifjorden,south-east Norway,with emphasis on their suitability as test organisms

Synopsis Mercury concentration in axial muscle of brown trout, Salmo trutta, whitefish, Coregonus lavaretus, Arctic charr, Salvelinus alpinus, smelt, Osmerus eperlanus and pike, Esox lucius, were studied in Lake Tyrifjorden during 1978–1982. Our data demonstrate that older and bigger fish on an average have higher mercury concentration than smaller and younger. Further, complex life histories as in brown trout influence the pattern of mercury accumulation. During young stages accumulation in brown trout is moderate, while accumulation in older stages is highly correlated to lake residency time. Based on our data we suggest the following requirements for a test organism and the collecting procedure; (1) life history should be simple with small sexual differences, (2) ageing should be easy and reliable, and (3) large representative samples should be easily obtained during (4) a fixed biological period i.e. the spawning period. We consider smelt as an appropriate test organism based on these criteria.     

Phylogenetic origin of <Emphasis Type="Italic">Salmo trutta</Emphasis> L 1758 from Sicily,based on mitochondrial and nuclear DNA analyses

The phylogenetic status of brown trout Salmo trutta L 1758 in Sicily is uncertain as some reports describe these trout as S. macrostigma or S. cettii on one hand while other, contradictory reports imply a hatchery origin on the other. In order to clarify this situation, we performed sequence analysis of the mtDNA control region and restriction fragment analysis of the nuclear lactate dehydrogenase (LDH-C1*) gene. A single mitochondrial haplotype (At-s6) found previously in brown trout in Morocco, and two alleles at LDH-C1* (the ancestral*100, at a high frequency, and *90) were revealed. Our results suggest that Sicilian brown trout are native and that they probably colonized Sicily from west to east in an expansion, from the Atlantic Ocean basin, along the North-West African coast. Handling editor: C. Sturmbauer     

Rb-Cs ratio as an indicator of fish diet in lakes of the Patagonia, Argentina

The ratios of Rb to Cs contents were studied in five fish species from seven lakes located in the Patagonia Andean Range, Argentina in order to trace fish diet. The species studied were native velvet catfish (Diplomistes viedmensis) and creole perch (Percichthys trucha), and exotic brown trout (Salmo trutta), rainbow trout (Oncorhynchus mykiss), and brook trout (Salvenilus fontinalis). Rainbow trout specimens from two farms were also studied, as well as fish food items and native mussels (Diplodon chilensis). Rb and Cs concentrations were determined by instrumental neutron activation analysis. A positive correlation of Cs concentration in the muscle of velvet catfish with fish length was observed, probably associated with the long biological half-life of this element in this species, whereas the Rb concentration remained constant, hence inhibiting the use of Rb-Cs ratios as a tracer in this case. Seasonal variations observed for rainbow trout and Cs concentration background bias in one of the lakes studied are also a limiting factor to the use of Rb-Cs ratios as a diet tracer. Rb-Cs ratios allowed clear differentiation of rainbow trout raised in farms from the natural specimens that lived in the same environment, in agreement with Rb-Cs ratios determined in both diets. Rb-Cs ratios in rainbow trout showed significant differences between Rivadavia and Futalaufquen lakes compared to Moreno and Nahuel Huapi lakes, which could be associated with a higher participation of plankton in the diet in the first case. No relevant variations in Rb-Cs ratios of brown trout were observed, probably because of the similarity in the diet.     

Heterochromatins and band karyotypes in three species of salmonids

Summary The heterochromatins of rainbow trout (Salmo gairdneri R.), brown trout (Salmo trutta fario L.) and brook trout (Salvelinus fontinalis M.) were characterized by sequential chromomycin A₃/distamycin A/DAPI (CDD) and DAPI/actinomycin D (DAPI/AmD) fluorescence. On most biarmed chromosomes, an equilocal localization of prominent DAPI/AmD positive, chromomycin A₃ negative, AT-rich blocks at the centromeres were observed in all three species. Band karyotypes of the three species were established. In rainbow trout, several DAPI/AmD positive heterochromatin blocks behaved positive in a silver-staining method. Mitotic and interphase studies proved the presence of inter-individual NOR variation in brown trout. The NORs of brook trout were localized on chromosomes 5, 10, 14, 15 and 29.     

Stock Transfers in Spanish Brown Trout Populations: A Long-term Assessment

Synopsis Stocking of fish from other populations has been commonly employed for enhancement of wild brown trout, Salmo trutta, populations in north Spain. Young hatchery reared brown trout of central European origin were introduced into some Asturian rivers every year since 1984. Based on variation at the isozyme locus LDH-C1* and at the microsatellite locus BFRO 002, two genetic markers race-specific in Salmo trutta, we detected introgression of foreign genomes into native gene pools in some Spanish trout populations where only pure native individuals were present 10 years ago. We strongly suggest development of alternative management policies for conservation of Spanish natural brown trout populations without endangering the traditional recreational fisheries. Jorge I. Izquierdo, Ana G. F. Castillo: These two authors contributed equally to the article.     

Chromosome painting supports lack of homology among sex chromosomes in Oncorhynchus, Salmo, and Salvelinus (Salmonidae)

The sex chromosome pair has been identified previously as the largest submetacentric pair in the genome in several species of the genus Salvelinus (eastern trouts and chars) including S. namaycush (lake trout) and as a large subtelocentric/acrocentric pair in several species of the genus Oncorhynchus (Pacific trouts and salmon). Sex chromosomes have not been identified in Salmo (Atlantic salmon and brown trout). Two paint probes, one specific for the short arm (Yp) and the other for the long arm (Yq) of the sex chromosome pair in Salvelinus namaycush were hybridized to chromosomes of Oncorhynchus mykiss (rainbow trout) and O. tshawytscha (chinook salmon) and Salmo salar (Atlantic salmon) and S. trutta (brown trout). The two probes hybridized to two different autosomal pairs in each of the Oncorhynchus species, supporting lack of homology between the sex chromosomes in the two genera. The Yp probe hybridized to interstitial regions on two different chromosome pairs in S. salar and one pair in S. trutta. The Yq probe hybridized to a different pair in both species.     

Trout (<Emphasis Type="Italic">Salmo trutta</Emphasis>) mitochondrial DNA polymorphism in the centre of the marble trout distribution area

The marble trout, a lineage of the Salmo trutta complex, is endemic to the Southern Alpine region. Although it is endangered throughout its entire distribution range, population genetic data were lacking for the central area, including the upper Etsch/Adige River system (South Tyrol, Northern Italy). A total of 672 Salmo trutta specimens, comprising phenotypic marble trout and phenotypic brown trout, from 20 sampling sites throughout South Tyrol were analysed by sequencing the complete mitochondrial DNA control region. Thirteen distinct haplotypes were identified, which clustered within three major genetic lineages: the Marmoratus (MA), the Atlantic (AT) and the Danubian (DA) lineage. 41.7% of the investigated individuals carried haplotypes of the MA lineage, 47.9% of the AT lineage and 10.4% of the DA lineage. It is noticeable that AT haplotypes were present at all sampling sites and no “pure” marble trout population with exclusively MA haplotypes was found. This points to a considerable impact of stocking with allochthonous brown trout, given that there is no evidence for natural colonisation by individuals of the AT lineage. However, our data indicate, for at least four localities, a limited gene flow between the native marble trout and hatchery-reared strains. Future conservation and rehabilitation measures will thus have to concentrate on the identification of remnant pure marble trout individuals from such mixed populations. Handling editor: C. Sturmbauer     

Genetic introgression on freshwater fish populations caused by restocking programmes

The brown trout (Salmo trutta L.) is one of the best studied native salmonids of Europe. Genetic studies on this species suggest that a large proportion of the evolutionary diversity corresponds to southern European countries, including the Iberian Peninsula, where this study is focused. Stocking activities employing non-indigenous hatchery specimens together with the destruction and fragmentation of natural habitats are major factors causing a decrease of native brown trout populations, mostly in the Mediterranean basins of the Iberian Peninsula. The main aim of the present work is to examine the genetic structure of the brown trout populations of the East Cantabrian region, studying the consequences of the restocking activities with foreign hatchery brown trout specimens into the wild trout populations. We have based our study on the Polymerase Chain Reaction and Restriction Fragment Length Polymorphism technique conducted on a mitochondrial fragment of 2700 base pairs and on the lactate dehydrogenase locus of the nuclear DNA. Our results show higher introgression rates in the Ebro (Mediterranean) basin than in the Cantabrian rivers. This revised version was published online in July 2006 with corrections to the Cover Date.     

Effects of salmon lice infection and salmon lice protection on fjord migrating Atlantic salmon and brown trout post-smolts

Effects of artificial salmon lice infection and pharmaceutical salmon lice prophylaxis on survival and rate of progression of Atlantic salmon (n = 72) and brown trout post-smolts (n = 72) during their fjord migration, were studied by telemetry. The infected groups were artificially exposed to infective salmon lice larvae in the laboratory immediately before release in the inner part of the fjord to simulate a naturally high infection pressure. Groups of infected Atlantic salmon (n = 20) and brown trout (n = 12) were also retained in the hatchery to control the infection intensity and lice development during the study period. Neither salmon lice infection nor pharmaceutical prophylaxis had any effects on survival and rate of progression of fjord migrating Atlantic salmon post-smolts compared to control fish. Atlantic salmon spent on average only 151.2 h (maximum 207.3 h) in passing the 80 km fjord system and had, thus, entered the ocean when the more pathogenic pre-adult and adult lice stages developed. The brown trout, in comparison to Atlantic salmon, remained to a larger extent than Atlantic salmon in the inner part of the fjord system. No effect of salmon lice infection, or protection, was found in brown trout during the first weeks of their fjord migration. Brown trout will, to a larger extent than Atlantic salmon, stay in the fjord areas when salmon lice infections reach the more pathogenic pre-adult and adult stages. In contrast to Atlantic salmon, they will thereby possess the practical capability of returning to freshwater when encountering severe salmon lice attacks.     

High Ag-NOR-site variation associated to a secondary contact in brown trout from the Iberian Peninsula

The analysis of nucleolar organizer regions (NORs) using silver (Ag-) staining and in situ hybridization (ISH) in brown trout (Salmo trutta) from various river basins in the Iberian Peninsula revealed high variation in the number and location of NORs. A total of 17 different Ag-NOR sites were revealed in 10 different chromosome pairs. Three different Ag-NOR patterns clustered by river basins and strongly associated to the internal transcribed spacer 1 (ITS1) variation were detected. The main variability in NOR-sites was found in a secondary contact between two divergent lineages of brown trouts at Duero basin. Our results confirmed the abrupt break in the spatial distribution of genetic variation of brown trout populations previously reported at Duero basin. We hypothesize that NOR-site variation might be a consequence of hybridization between divergent lineages of brown trouts and that NORs could play a major role in the maintenance of a hybrid zone in Duero basin via post-zygotic isolation mechanisms.     

Genetic characterization of brown trout (<Emphasis Type="Italic">Salmo trutta</Emphasis>) populations from the Southern Balkans using mtDNA sequencing and RFLP analysis

In the present study, a polymerase chain reaction-restriction fragment length polymorphism (PCR-RFLP) assay was used to survey variation in a 1450-bp mitochondrial DNA (mtDNA) segment which comprises part of the cytochrome oxidase III (COIII) and ATPase subunit VI genes in 8 brown trout (Salmo trutta L) populations from the southern Balkans. In addition, a 300 bp fragment at the 5′ end of the control region was sequenced from representatives of the populations studied providing the opportunity to assign PCR-RFLP haplotypes into major phylogenetic lineages (i.e. Atlantic, Danubian, marmoratus, Adriatic and Mediterranean). The level of polymorphism found in the 1450 bp segment suggests that this PCR-RFLP assay may be useful for future diagnostic analyses of mitochondrial DNA in brown trout populations. A reduced within-population genetic variability but considerable among-population differentiation was observed. The results are in accordance with previous data on phylogeography of Mediterranean brown trout suggesting that mitochondrial DNA haplotypes are distributed in a mosaic pattern as a consequence of a complex evolutionary history. The present study shows that brown trout populations from the Southern Balkans are highly divergent and possess a unique genetic profile that should be taken into account when establishing conservation management programs. Handling editor: C. Sturmbauer     

Genetic differentiation of southeast Baltic populations of sea trout inferred from single nucleotide polymorphisms

Sea trout (Salmo trutta m. trutta) is a migratory form of brown trout common in the Baltic Sea. Nine populations from the southeast Baltic (Poland; Lithuania; Denmark, Bornholm; Estonia and Russia) were genotyped using iPLEX Gold technology (Sequenom) with 62 informative SNPs. A diagnostic panel of 23 SNPs was applied to estimate genetic differentiation and assess the population structure of Baltic sea trout. The highest level of pairwise F_ST differences was observed between the Russian (East Gulf of Finland) and Polish (Baltic main basin) populations. The lowest differences were between the two Polish and the Polish and Lithuanian populations. A genetic similarity was noted between the Estonian Riguldi River and Danish Bornholm populations, and this finding was supported by a Bayesian and factorial correspondence analysis. Diversity within populations was highest for populations from Estonia and lowest for the Lithuanian population. Genetic structure analysis indicated that individuals from the nine populations were clustered into four groups.     

A shift in the habitat use pattern of a lentic galaxiid fish: an acute behavioural response to an introduced predator

Despite potentially reducing predation mortality, behavioural responses of native species to introduced predators may still have sub-lethal impacts. In video-recorded laboratory trials, we examined the effects of introduced brown trout, Salmo trutta, on the short-term behaviour of a threatened, lake-dwelling galaxiid fish and confirmed a suspected diel pattern in habitat use by this species. We found that Galaxias auratus followed a distinct diel pattern in the use of complex habitats and open water, which was significantly altered by the presence of brown trout. In trials without the introduced predator, G. auratus used complex habitats (rocks or macrophytes) during the day, and open water during the night. In trials with brown trout present, G. auratus spent significantly less time in open water and rarely ventured out of the macrophytes. However, when given the option of using only rocky substrate or open water, which is the more common situation in the lakes to which this galaxiid is endemic, the fish reduced the amount of time they spent in the open water during the night, but still spent more time in open water than when macrophytes were available. Spending the daylight hours amongst the cover of rocks or macrophytes is most likely an adaptation to reduce the risk of predation by visual predators, and the pattern of reduced use of open water habitats in the presence of brown trout is an acute response to the close proximity of the introduced predator. The difference in the nocturnal use of macrophytes and rocks when trout are present may be related to differences in feeding opportunities or success within these habitats.     

Interspecific association of brown trout (Salmo trutta) with non‐native brook trout (Salvelinus fontinalis) at the fry stage

The introduction of non‐native brook trout (Salvelinus fontinalis) in Europe has led to displacement and decreasing populations of native brown trout (Salmo trutta). Some studies have found that brown trout shift to a diet niche similar to brook trout when the two species live in sympatry, which conflicts with the competitive exclusion principle. A change in feeding niche may be a sign of early interspecific association and social learning, leading to behavioral changes. As a first step to address this possibility, it is essential to assess the interspecific association between the species during the early ontogenetic life stages. In this study, we therefore assess whether juvenile brown trout associate with non‐native juvenile brook trout to the same extent as with conspecifics by setting up two experiments: (i) a binomial choice test allowing visual and chemical cues to estimate the species specificity of group preference, and (ii) an association test without physical barriers to estimate the degree of association of a focal brown trout with a group of either conspecifics or heterospecifics. In experiment (1), we found that focal juvenile brown trout preferred to associate with the stimuli groups and did not discriminate either against conspecific or heterospecific groups. Furthermore, more active individuals showed stronger preference for the stimuli group than less active ones, regardless of species. In experiment (2), we found that brook trout groups had a tighter group structure than brown trout groups, and that focal brown trout showed stronger association with brook trout than with brown trout. These results indicate that brown trout may associate with brook trout at an early life stage, which would allow for interspecific social learning to occur. Future studies should look closer into causes and consequences of interspecific association and social learning, including potential effects on the phenotype selection in brown trout populations.