Variation of brown trout Salmo trutta young-of-the-year growth along environmental gradients in Europe

This study analysed the influence of temperature and other environmental factors on the growth of brown trout Salmo trutta YOY in Europe. Air temperature accounted for the greatest proportion of the variance in maximum total length, but the inclusion of other factors significantly increased the proportion of the variance explained.     

Distribution, growth and movement of River Usk brown trout (Salmo trutta)

The River Usk catchment in South Wales supports mainly freshwater resident brown trout, with few anadromous fish. Electric fishing and netting revealed that age-class distribution differed between main river and tributary habitats, the latter environment acting as a nursery area for young fish. Few fry were found at main river sites. Age-class distribution also differed between tributary systems, and possible reasons are discussed. Trapping experiments indicated that trout move to main river habitat at 2+ yr. Lengths at age (back-calculated from scale reading) were similar for main river and tributary resident fish at 1 and 2 year, but main river fish were larger at 3 and 4 yr. This habitat shift and size difference is discussed with reference to current angling regulations.     

Latitudinal clines in life-history variables of anadromous brown trout in Europe

Based on published data, we reviewed clinal variations in life-history characteristics of anadromous brown trout Salmo trutta from 102 European rivers at latitudes between 54 and 70° N. Growth rate in fresh water, mean smolt age, mean sea age at first sexual maturity, proportion of repeal spawners among adults, longevity, and length of adult life span exhibited latitudinal clines. Brown trout grew faster in fresh water, smolted and matured younger, lived fewer years but spawned more times in the south than in the north. The life-history traits studied were often correlated. Longevity, smolt age and sea age at maturity were negatively and smolt length and proportion of repeat spawners among adults were positively correlated with growth rate in fresh water. Longevity was positively correlated with smolt age and sea age at maturity. The latter also increased with increasing smolt age. None of these significant correlations among life history variables, except for those between smolt age and parr growth and proportion of repeat spawners and parr growth, are latitudinal effects. We do not know to what extent the latitudinal variation in life–history traits reflects phenotypic plasticity and to what extent it is caused by inherited differences among populations.     

Thermal dependence of embryonic growth and development in brown trout

Fertilized eggs from a brown trout Salmo trutta population in northern Spain were incubated in the laboratory at 4, 6, 8, 10, 12, 14, 16 and 18° C. Developmental stage and embryo size were monitored by taking samples at regular intervals. Survival was maximal at 8 and 10° C and decreased at higher and lower temperatures. Despite starting development, no embryo hatched at 16 and 18° C, which suggests an upper thermal limit for development between 14 and 16° C. Time required to reach a given ontogenetic stage decreased with increasing temperature. Embryos incubated at lower temperatures were larger at 50% hatching, and these differences persisted throughout the subsequent embryonic period until the start of exogenous feeding. A comparison with previously published data indicates low interpopulation variability in thermal sensitivity of embryonic development, even in consideration of the great latitudinal range of the studies.     

Recruitment variability of resident brown trout in peripheral populations from southern Europe

1. Population regulation was studied for seven consecutive years (1992–98) in five rivers at the periphery of the distribution of Salmo trutta, where the fish were living under environmental constraints quite different from those of the main distribution area. 2. Recruitment is naturally highly variable and the populations had been earlier classified as overexploited. Thus we expected that densities of young trout in most populations would be too low for density‐dependent mortality to operate. We tested this by fitting the abundance of recruits to egg densities over seven consecutive years (stock–recruitment relationship), and used the results to judge whether exploitation should be restricted in the interests of conserving the populations. 3. The density of 0+ trout in early September, as well as the initial density of eggs and parents, varied greatly among localities and years. The data for all populations fitted the Ricker stock–recruitment model. The proportion of variance explained by the population curves varied between 32% and 51%. However, in most cases the observations were in the density‐independent part of the stock–recruitment curve, where densities of the recruits increased proportionally with egg densities. 4. Our findings suggest that recruitment densities in most rivers and years were below the carrying capacity of the habitats. Although density‐dependent mechanisms seemed to regulate fish abundance in some cases, environmental factors and harvesting appeared generally to preclude populations from reaching densities high enough for negative feedbacks to operate. The findings thus lend support to Haldane’s (1956) second hypothesis that changes in population density are primarily due to density‐independent factors in unfavourable areas and areas with low density due to exploitation. Exploitation should be reduced to allow natural selection to operate more effectively.     

Variation in mitochondrial DNA and post-glacial colonization of north western Europe by brown trout

A purified mitochondrial DNA (mtDNA) probe was used to examine restriction fragment length polymorphisms produced by six restriction enzymes ( Xba I, Eco RV, Ava II, Hinf I, Hae III, Mbo I) in 915 brown trout from western Europe. A total of 20 composite haplotypes were found with one to seven haplotypes in individual populations. Icelandic trout samples from north, south, east, and west coast drainages showed only a single common haplotype in contrast to the high level of polymorphism found in Irish and Scottish populations. The phylogeny of mtDNA haplotypes and the pattern of haplotype distribution suggests that post-glacial colonization of brown trout in NW Europe was more complex than the dual colonization model which has been proposed on the basis of differential LDH-5* allele distribution. For example, Lough Melvin (Ireland) appears to have been independently     

Demogenetic structure of brown trout Salmo trutta Linnaeus, 1758 populations in mountain headwaters: implications for conservation management

A demogenetic analysis based on 7 years of observation (2005–2011) was conducted to examine the population structure of brown trout Salmo trutta in pristine dendritic headwaters. The value of genetic divergence (F_ST) among sampling units ranged from ?0.03 to 0.16. Demographic synchrony was low or moderate, and the average correlation coefficient of population growth between sampling units () ranged from 0.28 to 0.66. No isolation by distance was observed, but genetic divergence was negatively correlated with demographic synchrony among sampling units. Variance in the population growth rate (i.e. local extinction probability) increased with distance from the mainstream and from other sampling units. In contradiction with the usual model of stream‐dwelling salmonids, the upstream sections of headwaters holds only ephemeral subpopulations, whereas the mainstream played a role in the source area of the metapopulation. These findings stress the importance of the mainstream in management conservation for brown trout in low productive mountain headwaters.     

Individual variation in habitat use and growth of male and female brown trout

We examined variation in growth and habitat use of individually PIT‐tagged brown trout Salmo trutta in three stream enclosures, each divided into a fine substrate, deep pool habitat and a coarse substrate, shallow habitat. Habitat use and movements of individual fish were monitored continually by placing PIT detectors between habitats. All fish were measured and weighed biweekly over a three month period. There was no significant relationship between habitat use and initial body size, nor was there a consistent relationship between habitat use and densities of benthic macroinvertebrates or abundance of drifting invertebrates in the two habitats. Most habitat changes occurred at night, with activity peaks just prior to sunrise and after sunset. Trout used pools more at night than during the day. Within any given day, diurnal and nocturnal habitat use of individual fish varied little, with variation greater at night than during the day. Partial habitat segregation by sex was observed; only males used pools extensively during daytime, whereas males and females used riffles.
Growth rate was positively related to use of pools during daytime but not at night. Growth rate was also affected by enclosure, with growth rates being highest in the most downstream enclosure, which had the deepest pool (mean of 42 cm) and lowest in the most upstream enclosure, which had the shallowest pool (mean of 28 cm). A complete exchange of trout between the most upstream and downstream enclosure indicated that the enclosure effect was due to physical differences and not to individual fish differences between enclosures. The effect appears to have been caused by differences in depth as daytime use of pools was correlated with the area of the pool ≥35 cm deep, and production of trout biomass per enclosure was directly related to mean pool depth. Our results suggest that there is a relationship between habitat use and growth of individuals that is independent of body size, but that this relationship is influenced by sex of the fish and by the physical characteristics of the environment. Further, the data indicate that short‐term behavioral decisions on habitat use by brown trout have a potential effect on longer‐term individual fitness through growth rates.     

Prey consumption rates and growth of piscivorous brown trout in a subarctic watercourse

Prey consumption rates of piscivorous brown trout Salmo trutta were studied in the Pasvik watercourse, which forms the border between Norway and Russia. Estimates of food consumption in the field were similar to or slightly less than maximum values from a bioenergetic model. The piscivore diet consisted mainly of vendace Coregonus albula with a smaller number of whitefish Coregonus lavaretus . Individual brown trout had an estimated mean daily intake of c . 1·5 vendace and 0·4 whitefish, and a rapid annual growth increment of 7–8 cm year⁻¹. The total population of brown trout >25 cm total length was estimated as 8445 individuals (0·6 individuals ha⁻¹), giving a mean ±  s . e . annual consumption of 1553880 ± 405360 vendace and 439140 ± 287130 whitefish for the whole watercourse. The rapid growth in summer of brown trout >25 cm indicated a high prey consumption rate.     

Differences in growth between offspring of anadromous and freshwater brown trout Salmo trutta

In this study, individual growth of juvenile offspring of anadromous and freshwater resident brown trout Salmo trutta and crosses between the two from the River Imsa, Norway, was estimated. The juveniles were incubated until hatching at two temperatures (±S.D. ), either 4.4 ± 1.5°C or 7.1 ± 0.6°C. Growth rate was estimated for 22 days in August–September when the fish on average were c. 8 g in wet mass, and the estimates were standardized to 1 g fish dry mass. Offspring of anadromous S. trutta grew better at both 15 and 18°C than offspring of freshwater resident S. trutta or offspring of crosses between the two S. trutta types. This difference appears not to result from a maternal effect because anadromous S. trutta grew better than the hybrids with anadromous mothers. Instead, this appears to be an inherited difference between the anadromous and the freshwater resident fish lending support to the hypothesis that anadromous and freshwater resident S. trutta in this river differ in genetic expression. Egg incubation temperature of S. trutta appeared not to influence the later growth as reported earlier from the studies of Atlantic salmon Salmo salar.     

Fitness-related effects of growth investment in brown trout under field and hatchery conditions

The mortality of brown trout Salmo trutta over winter in a near-natural stream was not significantly increased by growth hormone (GH) treatment, but lipid reserves were lower in GH-treated fish. As GH-treated trout grew faster than controls, GH appears to promote growth at the expense of investment in maintenance. However, the growth promoting effect of GH was much more pronounced in the hatchery than in the stream, suggesting that the pay-off associated with increased growth investment is higher under hatchery conditions with unrestricted food supply than in wild, where food availability is limited.     

Characteristics of brown trout males influence growth and survival of their offspring

Half-sib groups of juvenile brown trout Salmo trutta (0+ years) having fathers with different characteristics were compared in several respects. Two kinds of experiments were performed, aquarium observations in the laboratory and stocking in a semi-natural stream. It was found in the aquarium experiments that the offspring of fathers with larger adipose fins were more active and were probably better at defending territories. Offspring of males chosen by females had higher growth rate. The feeding rate was higher in juveniles having a less aggressive father or a father that was chosen by the mother. The stream experiments showed that offspring of dominant males had higher growth rates, and that offspring of less aggressive fathers had higher survival. Thus, characteristics of the males (fathers) influences the quality of the offspring, and there seem to be benefits for the females both to choose among potential mates and to spawn with the most dominant male. As these male characteristics are never regarded during the process of artificial breeding in hatcheries, the results indicate a basis for divergence between hatchery and wild populations.     

Daily energy intake and growth of piscivorous brown trout,Salmo trutta

• 1 The chief objectives were to determine the daily energy intake and growth of piscivorous brown trout (Salmo trutta), and to compare the observed values with those expected from models developed previously for brown trout feeding on freshwater invertebrates. Energy budgets for individual fish were obtained from experiments with 40 trout (initial live weight 250–318 g) bred from wild parents, and kept at five constant temperatures (5, 10, 13, 15, 18 °C) and 100% oxygen saturation. Each trout was fed to satiation on freshly killed sticklebacks (Gasterosteus aculeatus) over a period of 42 days.
• 2 Energy intake (C_IN cal day^‐1) and growth (C_G cal day^‐1) were measured directly and energy losses (C_Q cal day^‐1) were estimated by difference (C_Q = C_IN ‐ C_G). All three variables increased with temperature. A model previously used to predict the daily energy intake (C_IN(INV)) of trout feeding to satiation on invertebrates was adapted, by changing only one parameter, to provide an excellent model (R² = 0.998) for predicting the mean daily energy intake (C_IN(ST)) for the piscivorous trout. Values of C_IN(ST) were 58% (range 48–67%) higher than those for C_IN(INV). A simple model was also developed to estimate mean daily energy losses for piscivorous trout (R² = 0.999). Both models were combined to provide excellent estimates of the daily energy gain (growth) of the piscivorous trout, and this was about three times that for trout feeding on invertebrates. The optimum temperature for maximum growth in energy terms increased from 13.9 °C for trout feeding on invertebrates to 17.0 °C (range 16.6–17.4 °C) for piscivorous trout.
• 3 The models are basically an extension of those developed for trout feeding on invertebrates. They show clearly how energy intake, growth, and the optimum temperature for growth increase markedly when trout change their diet from invertebrates to fish. The implications of this are discussed and it is shown that, in theory, these increases should continue if a more energy‐rich diet was utilised by the trout.

     

Forecasting fish distribution along stream networks: brown trout (Salmo trutta) in Europe

Aim

Species inhabiting fresh waters are severely affected by climate change and other anthropogenic stressors. Effective management and conservation plans require advances in the accuracy and reliability of species distribution forecasts. Here, we forecast distribution shifts of Salmo trutta based on environmental predictors and examine the effect of using different statistical techniques and varying geographical extents on the performance and extrapolation of the models obtained.

Location

Watercourses of Ebro, Elbe and Danube river basins (c. 1,041,000 km²; Mediterranean and temperate climates, Europe).

Methods

The occurrence of S. trutta and variables of climate, land cover and stream topography were assigned to stream reaches. Data obtained were used to build correlative species distribution models (SDMs) and forecasts for future decades (2020s, 2050s and 2080s) under the A1b emissions scenario, using four statistical techniques (generalised linear models, generalised additive models, random forest, and multivariate adaptive regression).

Results

The SDMs showed an excellent performance. Climate was a better predictor than stream topography, while land cover characteristics were not necessary to improve performance. Forecasts predict the distribution of S. trutta to become increasingly restricted over time. The geographical extent of data had a weak impact on model performance and gain/loss values, but better species response curves were generated using data from all three basins collectively. By 2080, 64% of the stream reaches sampled will be unsuitable habitats for S. trutta, with Elbe basin being the most affected, and virtually no new habitats will be gained in any basin.

Main conclusions

More reliable predictions are obtained when the geographical data used for modelling approximate the environmental range where the species is present. Future research incorporating both correlative and mechanistic approaches may increase robustness and accuracy of predictions.

     

Thermal sensitivity of growth, food intake and activity of juvenile brown trout

Growth rate, food intake and activity of juvenile brown trout (Salmo trutta L.) were studied at 10 constant temperatures (2.6-22.3 degrees C) and using eight full-sib families. We found between-family differences in growth rate but not significant temperaturexfamily interaction. The effects of temperature were well described by polynomial regressions that allowed the calculation of optima, performance breadths and thermal limits. The obtained values were noticeably higher and the performance breadth wider, than previously reported data for the species.     

Validation of scale-reading estimates of age and growth in a brown trout Salmo trutta population

In this study, we validated a scale-reading method estimating age and growth in brown trout Salmo trutta in wild, landlocked, stream-dwelling populations from mountain headwaters in the Elbe catchment area of the Czech Republic. The values estimated from scale reading were compared with measured values, collected using a mark-recapture program over eight consecutive years. The age-corrected absolute percentage error was 10.71%, primarily because the ages of the oldest individuals according to scale reading were underestimated, and the ages of juvenile individuals were slightly overestimated. The back-calculated length was slightly underestimated (the mean error was ?4.60 mm), but it was not significantly different from the real measured length. This study suggests that in cold mountain headwaters, scale reading is a sufficiently accurate method for age and growth estimation in juvenile brown trout; however, the results for adult individuals must be taken with caution.     

Prey orientation in piscivorous brown trout

Piscivorous brown trout Salmo trutta change their feeding behaviour depending on prey species, prey size and number of prey eaten. In trout which had eaten fish recently, most had one fish in their stomach, but up to 16 prey fish were found. Individuals of the small-sized minnow Phoxinus phoxinus were swallowed chiefly tail first, whereas individuals of the larger Arctic charr Salvelinus alpinus were taken both head and tail first. The largest charr were swallowed head first. In stomachs containing more than one fish prey, prey orientation was likely to be mixed. For all three types of prey orientation (i.e. tail first, head first and mixed), significant and positive correlations existed between prey length and predator length. The maximum prey size eaten tail first or in mixed orientation was about 70–85% of the size of prey eaten head first, indicating morphological advantages in eating the prey head first.