Torbjørn Forseth

The energy budget, niche shift, reproduction and growth in a population of Arctic charr, Salvelinus alpinus

We studied age at niche shift and sexual maturity of Arctic charr Salvelinus alpinus (L.) from a Norwegian lake in relation to their energy budget. Young Arctic charr (age −2) fed successfully on zooplankton giving them a significant energy surplus for somatic growth. Energy intake, however, levelled off at age −3 to −4, and decreased with age among older fish. Similarly, energetic costs increased from age −2 to −3 and then the costs levelled off and decreased, probably as a consequence of a niche shift from zooplankton to zoobenthos feeding in deeper water. This shift did not increase energy intake, but reduced costs, probably through higher feeding efficiency on the larger zoobenthos in the deeper water and reduced metabolism at lower ambient temperature

Ice-cover effects on competitive interactions between two fish species.

1. Variations in the strength of ecological interactions between seasons have received little attention, despite an increased focus on climate alterations on ecosystems. Particularly, the winter situation is often neglected when studying competitive interactions. In northern temperate freshwaters, winter implies low temperatures and reduced food availability, but also strong reduction in ambient light because of ice and snow cover. Here, we study how brown trout [Salmo trutta (L.)] respond to variations in ice-cover duration and competition with Arctic charr [Salvelinus alpinus (L.)], by linking laboratory-derived physiological performance and field data on variation in abundance among and within natural brown trout populations. 2. Both Arctic charr and brown trout reduced resting metabolic rate under simulated ice-cover (darkness) in the laboratory, compared to no ice (6-h daylight). However, in contrast to brown trout, Arctic charr was able to obtain positive growth rate in darkness and had higher food intake in tank experiments than brown trout. Arctic charr also performed better (lower energy loss) under simulated ice-cover in a semi-natural environment with natural food supply. 3. When comparing brown trout biomass across 190 Norwegian lakes along a climate gradient, longer ice-covered duration decreased the biomass only in lakes where brown trout lived together with Arctic charr. We were not able to detect any effect of ice-cover on brown trout biomass in lakes where brown trout was the only fish species. 4. Similarly, a 25-year time series from a lake with both brown trout and Arctic charr showed that brown trout population growth rate depended on the interaction between ice breakup date and Arctic charr abundance. High charr abundance was correlated with low trout population growth rate only in combination with long winters. 5. In conclusion, the two species differed in performance under ice, and the observed outcome of competition in natural populations was strongly dependent on duration of the ice-covered period. Our study shows that changes in ice phenology may alter species interactions in Northern aquatic systems. Increased knowledge of how adaptations to winter conditions differ among coexisting species is therefore vital for our understanding of ecological impacts of climate change.

Spatial distribution of limited resources and local density regulation in juvenile Atlantic salmon

1. Spatial heterogeneity of resources may influence competition among individuals and thus have a fundamental role in shaping population dynamics and carrying capacity. In the present study, we identify shelter opportunities as a limiting resource for juvenile Atlantic salmon (Salmo salar L.). Experimental and field studies are combined in order to demonstrate how the spatial distribution of shelters may influence population dynamics on both within and among population scales. 2. In closed experimental streams, fish performance scaled negatively with decreasing shelter availability and increasing densities. In contrast, the fish in open stream channels dispersed according to shelter availability and performance of fish remaining in the streams did not depend on initial density or shelters. 3. The field study confirmed that spatial variation in densities of 1-year-old juveniles was governed both by initial recruit density and shelter availability. Strength of density-dependent population regulation, measured as carrying capacity, increased with decreasing number of shelters. 4. Nine rivers were surveyed for spatial variation in shelter availability and increased shelter heterogeneity tended to decrease maximum observed population size (measured using catch statistics of adult salmon as a proxy). 5. Our studies highlight the importance of small-scale within-population spatial structure in population dynamics and demonstrate that not only the absolute amount of limiting resources but also their spatial arrangement can be an important factor influencing population carrying capacity.

Chernobyl radioactivity persists in fish.

After the accident at the Chernobyl nuclear reactor in 1986, the concentration of radioactive caesium (134Cs and 137Cs) in fish was expected to decline rapidly. The estimated ecological half-life (the time needed to reduce the average caesium concentration by 50%) was 0.3 to 4.6 years,. Since 1986, we have measured radiocaesium in brown trout (Salmo trutta) and Arctic charr (Salvelinus alpinus), both of which are widely eaten in Scandinavia, in a lake contaminated by Chernobyl fallout,. We have measured radiocaesium in nearly 4,000 fish, taking samples 2-4 times every year from spring to autumn. We find that the decline in radiocaesium was initially rapid for 3-4 years and was then much slower. About 10% of the initial peak radioactivity declines with an ecological half-life of as long as 8-22 years.

Thermal growth performance of juvenile brown trout Salmo trutta: no support for thermal adaptation hypotheses

Using thermal growth data from eight populations of anadromous and lake-feeding brown trout Salmo trutta, hypotheses of adaptation to local optima and countergradient variation in growth were tested. The adaptation to local optima hypothesis suggests that natural selection can shift optimal performance temperatures to match the prevailing temperature in a new or changed thermal niche. In contradiction, the countergradient variation hypothesis suggests that populations from hostile environments perform better than conspecifics from benign environments at all temperatures. In this study, growth capacity varied between populations but there was no significant correlation between any of the estimated thermal performance parameters (e.g. lower and upper thermal growth limits, optimal temperature for growth and maximum growth capacity) and natural climatic conditions among populations. Hence, S. trutta growth response to temperature lends no support for either of the two suggested thermal adaptation hypotheses. Instead, growth capacity among populations tended to correlate positively with female size at maturity.

Adaptive winter survival strategies: defended energy levels in juvenile Atlantic salmon along a latitudinal gradient

Current knowledge suggests that patterns of energy storage and depletion in animals are governed by behavioural trade-offs between risks associated with feeding and future energy demands. However, the length of adverse periods varies over geographical or climatic gradients. To explore the potential for genotypic sources of variation in behavioural trade-offs, we compared the winter energy-depletion patterns among 13 wild populations of juvenile Atlantic salmon (Salmo salar L.) along a latitudinal gradient (58–70°N) and performed common-environment experiments of energy-state-dependent feeding. In the wild, winter lipid-depletion rates were lower for northern than for southern populations. The variation in spring lipid levels among the population was lower than autumn variation, with storage lipid levels clustered close to critical limits for survival. In semi-natural stream channels with natural food supply, hatchery-reared fish originating from northern populations showed a positive scaling of feeding activity with decreasing energy levels, whereas southern populations did not. In conclusion, juvenile Atlantic salmon from northern populations defend their energy levels more strongly than fish from southern populations. Adaptive variation in feeding activity appears important for this difference. Thus, the present study shows a link between geographical patterns in storage energy trajectories and adaptive differences in state-dependent feeding motivation.

Effects of Water Chemistry and Habitat on the Density of Young Brown Trout Salmo trutta in Acidic Streams

We examined the relationship between young brown trout ( Salmo trutta) density in lake tributaries, and water chemistry and habitat variables. The study was carried out during the autumn in three acidic, softwater river systems in western and southwestern Norway; Gaular and Vikedal (1987–1993) and Bjerkreim (1988–1993). The streams had mean calcium concentrations of 0.35 mg L-1 (Gaular), 0.52 mg L-1 (Vikedal) and 0.84 mg L-1 (Bjerkreim). The concentration of inorganic Al was generally low, with mean values of 8.40 (Gaular), 22.22 (Vikedal) and 43.36 μg L-1 (Bjerkreim). In multiple regressions that involved different water chemistry variables, brown trout density correlated best with calcium concentration and with a combination of calcium and pH; the Ca2+:H+ ratio. In Vikedal and Gaular, calcium explained 51 and 57%, respectively, of the variability in brown trout densities. Althoug alkalinity exhibited the best correlation with brown trout density in Bjerkreim ( r2=0.33), it was similar to that of the model that included all major ions plus pH. The Ca2+:H+ ratio had a larger effect for variability in brown trout density in Gaular (r2=0.66) than calcium alone. In Vikedal and Bjerkreim, the Ca2+:H+ ratio also correlated with brown trout density, but considerably less than in Gaular. The predictive power of habitat variables was much lower than that of water chemistry; the single most important factors were altitude in Gaular (r2=0.22), mean water temperature in Vikedal (r2=0.11) and depth SD (index of heterogeneity) in Bjerkreim (r2=0.07). Models that included both habitat and water chemistry variables showed that the density of young brown trout was predicted primarily by calcium concentrations in Gaular (r2=0.75) and Vikedal (r2=0.54), as opposed to pH in Bjerkreim (r2=0.25). Habitat had low effect in all three river systems (r2=0.01–0.04). The final model explained 86, 68 and 32%, respectively, of the variability in brown trout density in the three catchments. Thus, water chemistry variables seem to be factors that limit the density of young brown trout in acidic softwater streams.

A FUNCTIONAL MODEL OF RADIOCESIUM TURNOVER IN BROWN TROUT

We describe a mechanistic model to explore the environmental and metabolic determinants of radiocesium kinetics in a freshwater fish, the brown trout (Salmo trutta). Our model, which incorporates submodels for fish growth, food consumption, ambient temperature, and radiocesium turnover, was validated and calibrated to observations in three year-classes of brown trout from a Norwegian lake contaminated with Chernobyl fallout. Observed growth and independent estimates of food consumption matched simulations. Whereas our model successfully described trends and seasonal dynamics of radiocesium in brown trout, it was highly sensitive to the temperature submodel with model uncertainty largely driven by uncertainties in absorption efficiency, feeding rate, and temperature. We used the model to simulate both fallout and steady-state (constant prey radiocesium) conditions. Peak radiocesium in fish after a fallout depended upon time of year, fish size, rate of food consumption and growth, and water temperature regime. Among the modeled lake types (temperate, boreal, and subalpine), peak radioactivity was highest for brown trout living in the warmest lake, feeding and growing at a maximum rate, and for fallout occurring early in spring. In these lakes, increasing growth rate from 80 to 100% of the maximum nearly doubled peak radioactivity. The model predicted that the relationship between body size and fish radioactivity changed from negative to positive within 2 yr after a fallout. At steady state, the model predicted seasonal dynamics in accumulation of radiocesium, with maximum biomagnification in autumn and minimum in early spring. Maximum biomagnification was related to fish size (weak negative slope), growth rate, and temperature regime, and a metamodel was developed for predicting biomagnification.

A concept for improving Atlantic salmon Salmo salar smolt migration past hydro power intakes.

In this study, cost effective (in terms of reducing loss of power production) measures for increasing bypass migration of Atlantic salmon Salmo salar were developed and tested by establishing statistical models for timing of smolt migration and favourable diversion of water to the bypass. Initial tracking of radio-tagged smolts showed very low bypass migration under normal hydropower operations. Bypass migration increased when bypass discharge was experimentally increased and a model was developed that described relationships between total river discharge, bypass diversion and smolt migration route. Further improvements were obtained by installing two strobe lights at the power-production tunnel entrance that increased bypass migration during the night, but not during daytime. According to the behaviour of radio-tagged fish, the implemented measures contributed to increasing the annual percentage of bypass migration from 11 to 64%, and according to model predictions to 60-74% when the hydropower facilities were operated according to the developed models. To ensure correct timing of discharge diversion a smolt migration model was developed based on environmental variables that could successfully predict the general pattern of migration timing. The concept presented for improving smolt migration past hydropower intakes should be applicable in many systems where migration past hydropower installations cannot easily be solved by screening systems.

Habitat use and life history of brown trout (Salmo trutta) and Arctic charr (Salvelinus alpinus) in some low acidity lakes in central Norway

Habitat utilization and the life history of browntrout Salmo trutta and Arctic charr Salvelinus alpinus were investigated in fivesympatric populations and five allopatric brown troutpopulations in Høylandet catchment, a atmosphaericlow deposition area in Mid Norway. There was asignificant inverse correlation in abundance ofepibenthic Arctic charr and brown trout in theselakes, indicating that the latter species is dominant.The largest numbers of sympatric brown trout andArctic charr were caught in epibenthic habitat. In twolakes, brown trout to some extent also occurredpelagically, while pelagic individuals of Arctic charrwere found in all five lakes. The main food items forboth epibenthic and pelagic brown trout wereterrestrial surface insects and chironomid pupae.Zooplankton was the primary food item for Arctic charrin both habitats. Although the age distribution wasvery different in the populations, neither speciesseem to suffer from recruitment failure. There was nosignificant difference in survival rates betweensympatric populations of brown trout and Arctic charr.We found a significant inverse correlation betweenepibenthic catches of brown trout and the mean weightof 4+ fish, the most abundant age group. However, ifusing weight data for three-year-old fish, no suchrelationship was found for Arctic charr. Brown troutand Arctic charr reached asymptotic lengths of197–364 mm and 259–321 mm, respectively. Both speciestypically reached sexual maturity at age 2–3, and nomaturation-induced mortality was evident. We concludethat fish populations in Høylandet lakes areregulated throughout their lifes by inter- andintraspecific competition.