Thermal preference and performance provide the physiological frame within which fish species seek strategies to cope with the challenges raised by the low temperatures and low levels of oxygen and food that characterize winter. There are two common coping strategies: active utilization of winter conditions or simple toleration of winter conditions. The former is typical of winter specialist species with low preferred temperatures, and the latter is typical of species with higher preferred temperatures. Reproductive strategies are embodied in the phenology of spawning: the approach of winter conditions cues reproductive activity in many coldwater fish species, while the departure of winter conditions cues reproduction in many cool and warmwater fish species. This cuing system promotes temporal partitioning of the food resources available to young-of-year fish and thus supports high diversity in freshwater fish communities. If the zoogeographic distribution of a species covers a broad range of winter conditions, local populations may exhibit differences in their winter survival strategies that reflect adaptation to local conditions. Extreme winter specialists are found in shallow eutrophic lakes where long periods of ice cover cause winter oxygen levels to drop to levels that are lethal to many fish. The fish communities of these lakes are simple and composed of species that exhibit specialized adaptations for extended tolerance of very low temperatures and oxygen levels. Zoogeographic boundaries for some species may be positioned at points on the landscape where the severity of winter overwhelms the species' repertoire of winter survival strategies. Freshwater fish communities are vulnerable to many of the shifts in environmental conditions expected with climate change. Temperate and northern communities are particularly vulnerable since the repertoires of physiological and behavioural strategies that characterize many of their members have been shaped by the adverse environmental conditions (e.g. cool short summers, long cold winters) that climate change is expected to mitigate. The responses of these strategies to the rapid relaxation of the adversities that shaped them will play a significant role in the overall responses of these fish populations and their communities to climate change.

Here, we demonstrate a contrasting effect of terrestrial coloured dissolved organic material on the secondary production of boreal nutrient poor lakes. Using fish yield from standardised brown trout gill-net catches as a proxy, we show a unimodal response of lake secondary productivity to dissolved organic carbon (DOC). This suggests a trade-off between positive and negative effects, where the initial increase may hinge upon several factors such as energy subsidising, screening of UV-radiation or P and N load being associated with organic carbon. The subsequent decline in production with further increase in DOC is likely associated with light limitations of primary production. We also show that shallow lakes switch from positive to negative effects at higher carbon loads than deeper lakes. These results underpin the major role of organic carbon for structuring productivity of boreal lake ecosystems.

Increased concentrations of dissolved organic carbon (DOC), often labelled “browning”, is a current trend in northern, particularly boreal, freshwaters. The browning has been attributed to the recent reduction in sulphate (S) deposition during the last 2 to 3 decades. Over the last century, climate and land use change have also caused an increasing trend in vegetation cover (“greening”), and this terrestrially fixed carbon represents another potential source for export of organic carbon to lakes and rivers. The impact of this greening on the observed browning of lakes and rivers on decadal time scales remains poorly investigated, however. Here, we explore time-series both on water chemistry and catchment vegetation cover (using NDVI as proxy) from 70 Norwegian lakes and catchments over a 30-year period. We show that the increase in terrestrial vegetation as well as temperature and runoff significantly adds to the reduced SO4-deposition as a driver of freshwater DOC concentration. Over extended periods (centuries), climate mediated changes in vegetation cover may cause major browning of northern surface waters, with severe impact on ecosystem productivity and functioning.

1. Anthropogenic disturbances of the physical habitat and corresponding effects on fish performance are key issues in stream conservation and restoration. Reduced habitat complexity because of increased sediment loadings and canalization is of particular importance, but it is not clear to what extent fish populations are influenced directly by changes in the physical environment, or indirectly through changes in the biotic environment affecting the food availability. 2. Here, we test for the direct effect of habitat complexity on the performance (growth) of juvenile Atlantic salmon by manipulating shelter availability (interstitial spaces in the substrate) across 20 semi-natural stream channels without altering the substrate composition, and stocking each channel with a common density of fish. A simple method for measuring salmonid shelters using flexible PVC tubes was developed and tested. Daytime sheltering behaviour and growth rates were compared across the channels differing in shelter availability. 3. Measured shelter availability was strongly negatively correlated with observed number of fish not finding shelters and mass loss rates of the fish (growth performance) increased with decreasing number of measured shelters. Number and mean depth of interstitial spaces explained up to 68% and 24% of the among-channel variation in sheltering behaviour and growth performance, respectively. Furthermore, negative effects of shelter reduction increased with fish body size. Thus, changes in habitat structure may even influence the size selection gradients. 4. Shelter availability is an easily measured variable, possibly affecting the population demographics and long-term evolutionary processes, and is therefore a key habitat factor to be considered in stream restoration and habitat classification.

By comparing the population frequency distributions for specific somatic energy between samplings using quantile–quantile (QQ) plots, we tested for energy-related mortality of juvenile (2- and 3-year-old) Atlantic salmon (Salmo salar) sampled at monthly intervals throughout three consecutive winters in a Norwegian river located at 70°N. Between several of the sampling periods, changes in the distributions of specific energy were observed corresponding to removal of low-energy individuals. By using energetic modelling we demonstrated that metabolic processes or feeding could not be responsible for the shifts in the shape of the energy distributions and that negative-energy-dependent mortality was the most likely explanation for the observations. No changes in mean size of the fish or in the shape of the size distributions were observed between successive sampling periods, indicating that mortality was linked to levels of storage energy rather than to body size per se. Our study indicated a critical body energy level for survival of juvenile salmon at approximately 4400–4800 J·g–1, corresponding to a depletion of storage lipids.

We tested the importance of thermal adaptations and energy efficiency in relation to the geographical distribution of two competing freshwater salmonid fish species. Presence-absence data for Arctic char and brown trout were obtained from 1502 Norwegian lakes embracing both temperature and productivity gradients. The distributions were contrasted with laboratory-derived temperature scaling models for food consumption, growth and energy efficiency. Thermal performances of the two species were almost identical. However, Arctic char exhibited double the growth efficiency (per unit of food) and appear to have out-competed brown trout from cold, low-productivity lakes, perhaps by scramble competition. Brown trout, for which previous reports have shown to be aggressive and dominant, have likely excluded the more energy-efficient Arctic char from relatively warm, productive lakes, perhaps by contest competition. Competitive interaction changing in outcome with lake productivity, rather than thermal performance, is likely a major determinant of the range distribution of the two species. Our study highlights the need for more focus on choice of relevant ecophysiological traits in ecological climate impact studies and species distribution modelling.

Summary 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.

Summary 1.Under benign laboratory tank conditions we compared food consumption and metabolism of Atlantic salmon (Salmo salar) juveniles exposed to simulated ice cover (darkness) with fish in natural short, 6 h light, day length (without ice). Three different populations along an ice-cover gradient were tested (59°N−70°N). 2. Resting metabolism was on average 30% lower under simulated ice cover (6·6 J g −1 day−1 ) than under natural day length (9·4 J g −1 day −1 ), and the response was similar for all populations. Northern salmon grew equally well in dark and light conditions, whereas the southern grew significantly poorer in the dark. Fish from all populations fed more under natural day length than in the dark and the northern population had higher consumption than the southern. The relative high growth of fish from the northern population in the dark compared to the southern populations was due partly to higher consumption and partly to higher growth efficiency. Fish from the southern populations had negative growth efficiency in the dark. 3. We also studied the importance of ice cover under more hostile conditions in stream channels using the northern population only. Juveniles held in channels with simulated ice cover lost less energy (20 J g −1 day −1 ) than those held in channels with transparent cover (26 J g −1 day −1 ). This difference in energy loss was due partly (50%) to higher food consumption under simulated ice (4·5 and 1·6 J g −1 , respectively) and partly (30%) to light-induced differences in resting metabolic rate. 4. In conclusion, both experiments showed lower metabolic costs in darkness under simulated ice cover than without ice. Under benign laboratory conditions the response to light (ice cover) varied among populations and only the northern population were able to attain positive growth in the dark. Under semi-natural conditions the lack of ice cover induced strong negative effects on the energy budget. Because energetic deficiencies are assumed to be an important cause of winter mortality, our study indicates that ice break-ups or removal following climatic change may affect winter survival significantly, particularly in northern populations.