A shift in habitat use patterns of brown trout (Salmo trutta): A behavioural response to macrophyte removal.

Mass development of macrophytes is an increasing problem worldwide and they are frequently removed where they are in conflict with local waterway users. Yet, macrophytes can provide important refuge and nursery habitats for fish. Little is known about the consequences of macrophyte removal for fish behavioural space use and habitat selection. We hypothesised that macrophyte removal would affect brown trout (Salmo trutta) movement during the partial removal of the aquatic plant Juncus bulbosus (L.) in an oligotrophic impounded Norwegian river.We tagged 94 brown trout and tracked them using passive acoustic telemetry for 10 months and mapped the cover of J. bulbosus. Trout behavioural patterns were quantified as space use (utilisation areas 50% and 95%) which was linked to habitat use and selection for J. bulbosus. Removal of J. bulbosus influenced space use of brown trout by reducing the core utilisation area by 22%. Habitat use and selection were likewise influenced by removal with increased use and selection of areas with low J. bulbosus cover (<25%) with corresponding reduction in high J. bulbosus cover (>25-75%). Finally, diurnal differences in space use and habitat use were found, with 19% larger utilisation areas at night and higher use of areas with low J. bulbosus during daytime. Yet, all effect sizes were relatively small compared to the size of the study area. This research provides a detailed case study on the effects of macrophyte removal on fish behavioural patterns in a section of a large Norwegian river with macrophyte mass development. We found no large effects of removal on trout behaviour but noted an increased use of areas with low macrophyte cover. This research is relevant for water managers and policy makers of freshwater conservation and provides a template for using acoustic telemetry to study the effects of macrophyte removal on fish.

Empirical support for sequential imprinting during downstream migration in Atlantic salmon (Salmo salar) smolts.

The precise homing of Atlantic salmon to their natal river and spawning grounds is the foundation for locally adapted genetically differentiated populations across rivers or across river sections. A sequential imprinting hypothesis states that salmon smolts may imprint on environmental clues along the outward migration route and then use this in reverse order to direct the spawning migration later in life. In this study, we provide empirical support for this hypothesis. PIT-tagged wild Atlantic salmon using a 2 km hydropower tunnel as downstream migrating smolts had a 18% (1SW) and 23% (2SW) lower probability of successfully migrating through the parallel river stretch as adult spawners compared to spawners that migrated through the same river stretch as smolts. These findings highlight how a fine-scale riverine migration route may be imprinted in wild Atlantic salmon smolts. From an applied perspective, these results stress the importance of not depriving smolts from parts of their migration route to ensure successful return of adults to their natal spawning grounds.

Hydropower-induced selection of behavioural traits in Atlantic salmon (Salmo salar).

Renewable energy projects such as hydropower facilities contribute towards meeting the world`s growing energy demands and urgent need for mitigating climate change. However, such infrastructure has the potential to substantially alter the environment which, in turn, can induce new challenges related to for instance fish migration conditions. As a consequence, local adaptations related to pre-development migration conditions may be affected for influenced populations. To explore selection regimes operating at a river hydropower plant, we monitored Atlantic salmon smolt individuals during their seaward migration. When passing the hydropower plant, the smolts chose between a surface fish passage or a submerged turbine intake. Smolts were scored for behavioural type (basal locomotor activity, net restrain (a measure of escape responses) and willingness to leave a familiar environment) prior to their migration choice, and we found that smolts with high basal activity had higher probability of using the fish passage than the turbine intake. In addition, migration route choice was a partly consistent trait in that fish that had previously passed a hydroelectric facility by using a fish passage rather than the turbine intake were significantly more likely to use it again when faced with the same choice. Higher mortality among turbine migrants could potentially reduce or eliminate particular behaviour types within populations- and the corresponding population genetic diversity that is essential to cope with future environmental challenges.

Population consequences of climate change through effects on functional traits of lentic brown trout in the sub-Arctic.

Climate-induced plasticity in functional traits has received recent attention due to the immense importance phenotypic variation plays in population level responses. Here, we explore the effect of different climate-change scenarios on lentic populations of a freshwater ectotherm, the brown trout (Salmo trutta L.), through climate effects on functional traits. We first parameterize models of climate variables on growth, spawning probability and fecundity. The models are utilized to inform a dynamic age-structured projection matrix, enabling long-term population viability projections under climate and population density variation. Ambient temperature and winter conditions had a substantial effect on population growth rate. In general, warmer summer temperatures resulted in faster growth rates for young fish but ended in smaller size at age as fish got older. Increasing summer temperatures also induced maturation at younger age and smaller size. In addition, we found effects of first-year growth on later growth trajectories for a fish, indicating that environmental conditions experienced the first year will also influence size at age later in life. At the population level, increasing temperatures average (up to 4 °C increase in areas with mean summer temperature at approximately 12 °C) resulted in a positive effect on population growth rate (i.e. smaller but more fish) during climate simulations including increasing and more variable temperatures.

Synchrony and multimodality in the timing of Atlantic salmon smolt migration in two Norwegian fjords.

The timing of the smolt migration of Atlantic salmon (Salmo salar) is a phenological trait increasingly important to the fitness of this species. Understanding when and how smolts migrate to the sea is crucial to understanding how salmon populations will be affected by both climate change and the elevated salmon lice concentrations produced by salmon farms. Here, acoustic telemetry was used to monitor the fjord migration of wild post-smolts from four rivers across two fjord systems in western Norway. Smolts began their migration throughout the month of May in all populations. Within-population, the timing of migration was multimodal with peaks in migration determined by the timing of spring floods. As a result, migrations were synchronized across populations with similar hydrology. There was little indication that the timing of migration had an impact on survival from the river mouth to the outer fjord. However, populations with longer fjord migrations experienced lower survival rates and had higher variance in fjord residency times. Explicit consideration of the multimodality inherent to the timing of smolt migration in these populations may help predict when smolts are in the fjord, as these modes seem predictable from available environmental data.

Freezer on, lights off! Environmental effects on activity rhythms of fish in the Arctic.

Polar regions are characterized by acute seasonal changes in the environment, with organisms inhabiting these regions lacking diel photoperiodic information for parts of the year. We present, to our knowledge, the first high-resolution analysis of diel and seasonal activity of free-living fishes in polar waters (74°N), subject to extreme variation in photoperiod, temperature and food availability. Using biotelemetry, we tracked two sympatric ecomorphs of lake-dwelling Arctic charr (Salvelinus alpinus n = 23) over an annual cycle. Charr activity rhythms reflected the above-surface photoperiod (including under ice), with diel rhythms of activity observed. During the dark winter solstice period, charr activity became arrhythmic and much reduced, even though estimated light levels were within those at which charr can feed. When twilight resumed, charr activity ensued as diel vertical migration, which continued throughout spring and with increasing day length, despite stable water temperatures. Diel activity rhythms ceased during the polar day, with a sharp increase in arrhythmic fish activity occurring at ice-break. Despite contrasting resource use, circannual rhythms were mirrored in the two ecomorphs, although individual variability in activity rhythms was evident. Our data support conclusions of functionally adaptive periods of arrhythmicity in polar animals, suggesting maintenance of a circannual oscillator for scheduling seasonal behavioural and developmental processes.

Genetic differences between wild and hatchery-bred brown trout (Salmo trutta L.) in single nucleotide polymorphisms linked to selective traits.

To study effects from natural selection acting on brown trout in a natural stream habitat compared with a hatchery environment, 3,781 single nucleotide polymorphism (SNP) markers were analyzed in three closely related groups of brown trout (Salmo trutta L.). Autumn (W/0+, n = 48) and consecutive spring (W/1+, n = 47) samples of brown trout individuals belonging to the same cohort and stream were retrieved using electrofishing. A third group (H/1+, n = 48) comprised hatchery-reared individuals, bred from a mixture of wild parents of the strain of the two former groups and from a neighboring stream. Pairwise analysis of FST outliers and analysis under a hierarchical model by means of ARLEQUIN software detected 421 (10.8%) candidates of selection, before multitest correction. BAYESCAN software detected 10 candidate loci, all of which were included among the ARLEQUIN candidate loci. Body length was significantly different across genotypes at 10 candidate loci in the W/0+, at 34 candidate loci in the W/1+ and at 21 candidate loci in the H/1+ group. The W/1+ sample was tested for genotype-specific body length at all loci, and significant differences were found in 10.6% of all loci, and of these, 14.2% had higher frequency of the largest genotype in the W/1+ sample than in W/0+. The corresponding proportion among the candidate loci of W/1+ was 22.7% with genotype-specific body length, and 88.2% of these had increased frequency of the largest genotype from W/0+ to W/1+, indicating a linkage between these loci and traits affecting growth and survival under this stream's environmental conditions. Bayesian structuring of all loci, and of the noncandidate loci suggested two (K = 2), alternatively four clusters (K = 4). This differed from the candidate SNPs, which suggested only two clusters. In both cases, the hatchery fish dominated one cluster, and body length of W/1+ fish was positively correlated with membership of one cluster both from the K = 2 and the K = 4 structure. Our analysis demonstrates profound genetic differentiation that can be linked to differential selection on a fitness-related trait (individual growth) in brown trout living under natural vs. hatchery conditions. Candidate SNP loci linked to genes affecting individual growth were identified and provide important inputs into future mapping of the genetic basis of brown trout body size selection.

Contemporary divergence in early life history in grayling (Thymallus thymallus).

BACKGROUND: Following colonization of new habitats and subsequent selection, adaptation to environmental conditions might be expected to be rapid. In a mountain lake in Norway, Lesjaskogsvatnet, more than 20 distinct spawning demes of grayling have been established since the lake was colonized, some 20-25 generations ago. The demes spawn in tributaries consistently exhibiting either colder or warmer temperature conditions during spawning in spring and subsequent early development during early summer. In order to explore the degree of temperature-related divergence in early development, a multi-temperature common-garden experiment was performed on embryos from four different demes experiencing different spring temperatures. RESULTS: Early developmental characters were measured to test if individuals from the four demes respond differently to the treatment temperatures. There was clear evidence of among-deme differences (genotype - environment interactions) in larval growth and yolk-to-body-size conversion efficiency. Under the cold treatment regime, larval growth rates were highest for individuals belonging to cold streams. Individuals from warm streams had the highest yolk-consumption rate under cold conditions. As a consequence, yolk-to-body-mass conversion efficiency was highest for cold-deme individuals under cold conditions. As we observed response parallelism between individuals from demes belonging to similar thermal groups for these traits, some of the differentiation seems likely to result from local adaptation CONCLUSION: The observed differences in length at age during early larval development most likely have a genetic component, even though both directional and random processes are likely to have influenced evolutionary change in the demes under study.

Total mercury in wild fish in Guizhou reservoirs, China.

The health hazard of mercury (Hg) compounds is internationally recognized, and the main pathways for methylmercury (MeHg) intake in humans are through consumption of food, especially fish. Given the large releases of Hg to the environment in China, combined with the fast development of hydropower, this issue deserves attention. Provided similar mobilization pathways of Hg in China as seen in reservoirs in North America and Europe one should expect increased Hg contamination in relation to future hydropower reservoir construction in this country. This study presents total Hg (THg) concentrations in wild fish from six Guizhou reservoirs, China. The THg concentrations in fish were generally low despite high background levels in the bedrock and depositions from local point sources. The over all mean +/- SD concentration of THg was (0.066 +/- 0.078) microg/g (n = 235). After adjusting for among-reservoir variation in THg, there were significant differences in THg among functional groups of the fish, assumed to reflect trophic levels. Predicted THg-concentration ratios, retrieved from a mixed linear model, between the functional groups were 9:4:4:1 for carnivorous, omnivorous, planktivorous and herbivorous fish. This result indicated that MeHg accumulation may prevail even under circumstances with short food chains as in this Chinese water system. No fish exceeded recommended maximum THg limit for human consumption set by World Health Organization and the Standardization Administration of China (0.5 microg/g fish wet weight (ww)). Only six fish (2.5%) exceeded the maximum THg limit set by US Environmental Protection Agency (0.3 microg/g fish ww).

Contemporary temperature-driven divergence in a Nordic freshwater fish under conditions commonly thought to hinder adaptation.

BACKGROUND: Evaluating the limits of adaptation to temperature is important given the IPCC-predicted rise in global temperatures. The rate and scope of evolutionary adaptation can be limited by low genetic diversity, gene flow, and costs associated with adaptive change. Freshwater organisms are physically confined to lakes and rivers, and must therefore deal directly with climate variation and change. In this study, we take advantage of a system characterised by low genetic variation, small population size, gene flow and between-trait trade-offs to study how such conditions affect the ability of a freshwater fish to adapt to climate change. We test for genetically-based differences in developmental traits indicating local adaptation, by conducting a common-garden experiment using embryos and larvae from replicate pairs of sympatric grayling demes that spawn and develop in natural cold and warm water, respectively. These demes have common ancestors from a colonization event 22 generations ago. Consequently, we explore if diversification may occur under severely constraining conditions. RESULTS: We found evidence for divergence in ontogenetic rates. The divergence pattern followed adaptation predictions as cold-deme individuals displayed higher growth rates and yolk conversion efficiency than warm-deme individuals at the same temperature. The cold-deme embryos had a higher rate of muscle mass development. Most of the growth- and development differences occurred prior to hatch. The divergence was probably not caused by genetic drift as there was a strong degree of parallelism in the divergence pattern and because phenotypic differentiation (Q(ST)) was larger than estimated genetic drift levels (microsatellite F(ST)) between demes from different temperature groups. We also document that these particular grayling populations cannot develop successfully at temperatures above 12°C, whereas other European populations can, and that increasing the muscle mass development rate comes at the cost of some skeletal trait development rates. CONCLUSIONS: This study shows that genetically based phenotypic divergence can prevail even under conditions of low genetic variation and ongoing gene flow. Furthermore, population-specific maximum development temperatures along with musculoskeletal developmental trade-offs may constrain adaptation.

Body downsizing caused by non-consumptive social stress severely depresses population growth rate.

Chronic social stress diverts energy away from growth, reproduction and immunity, and is thus a potential driver of population dynamics. However, the effects of social stress on demographic density dependence remain largely overlooked in ecological theory. Here we combine behavioural experiments, physiology and population modelling to show in a top predator (pike Esox lucius) that social stress alone may be a primary driver of demographic density dependence. Doubling pike density in experimental ponds under controlled prey availability did not significantly change prey intake by pike (i.e. did not significantly change interference or exploitative competition), but induced a neuroendocrine stress response reflecting a size-dependent dominance hierarchy, depressed pike energetic status and lowered pike body growth rate by 23 per cent. Assuming fixed size-dependent survival and fecundity functions parameterized for the Windermere (UK) pike population, stress-induced smaller body size shifts age-specific survival rates and lowers age-specific fecundity, which in Leslie matrices projects into reduced population rate of increase (lambda) by 37-56%. Our models also predict that social stress flattens elasticity profiles of lambda to age-specific survival and fecundity, thus making population persistence more dependent on old individuals. Our results suggest that accounting for non-consumptive social stress from competitors and predators is necessary to accurately understand, predict and manage food-web dynamics.

Contemporary isolation-by-distance, but not isolation-by-time, among demes of European Grayling (Thymallus thymallus, Linnaeus) with recent common ancestors.

The development of isolation by distance (IBD) and isolation by time (IBT) was contrasted among demes of European grayling (Thymallus thymallus) that have diverged within the last 25 generations following colonization of a lake (Lesjaskogsvatnet). We find low but significant levels of genetic differentiation among spawning tributaries and a pattern of IBD among them. We do not, however, find evidence for IBT despite an up to four-week difference in spawning date between "warm/early" and "cold/late" spawning demes and differences in the incubation temperatures experienced by offspring. It appears that IBD has developed more rapidly than IBT in this system and that adaptive divergence has been initiated in the absence of IBT. Although analysis of selected loci could reveal reduced recombination in parts of the genome associated with temporal divergence, our analysis of neutral genetic data suggests that IBD is a more important isolating mechanism in the early stages of adaptive divergence in European grayling.

Trait changes in a harvested population are driven by a dynamic tug-of-war between natural and harvest selection.

Selective harvest of large individuals should alter natural adaptive landscapes and drive evolution toward reduced somatic growth and increased reproductive investment. However, few studies have simultaneously considered the relative importance of artificial and natural selection in driving trait changes in wild populations. Using 50 years of individual-based data on Windermere pike (Esox lucius), we show that trait changes tracked the adaptive peak, which moved in the direction imposed by the dominating selective force. Individual lifetime somatic growth decreased at the start of the time series because harvest selection was strong and natural selection was too weak to override the strength of harvest selection. However, natural selection favoring fast somatic growth strengthened across the time series in parallel with the increase in pike abundance and, presumably, cannibalism. Harvest selection was overridden by natural selection when the fishing effort dwindled, triggering a rapid increase in pike somatic growth. The two selective forces appear to have acted in concert during only one short period of prey collapse that favored slow-growing pike. Moreover, increased somatic growth occurred concurrently with a reduction in reproductive investment in young and small female pike, indicating a tradeoff between growth and reproduction. The age-specific amplitude of this change paralleled the age-specific strength of harvest pressure, suggesting that reduced investment was also a response to increased life expectancy. This is the first study to demonstrate that a consideration of both natural selection and artificial selection is needed to fully explain time-varying trait dynamics in harvested populations.

Four decades of opposing natural and human-induced artificial selection acting on Windermere pike (Esox lucius).

The ability of natural selection to drive local adaptation has been appreciated ever since Darwin. Whether human impacts can impede the adaptive process has received less attention. We tested this hypothesis by quantifying natural selection and harvest selection acting on a freshwater fish (pike) over four decades. Across the time series, directional natural selection tended to favour large individuals whereas the fishery targeted large individuals. Moreover, non-linear natural selection tended to favour intermediate sized fish whereas the fishery targeted intermediate sized fish because the smallest and largest individuals were often not captured. Thus, our results unequivocally demonstrate that natural selection and fishery selection often acted in opposite directions within this natural system. Moreover, the two selective factors combined to produce reduced fitness overall and stronger stabilizing selection relative to natural selection acting alone. The long-term ramifications of such human-induced modifications to adaptive landscapes are currently unknown and certainly warrant further investigation.

The ideal free pike: 50 years of fitness-maximizing dispersal in Windermere.

The ideal free distribution (IFD) theory is one of the most influential theories in evolutionary ecology. It predicts how animals ought to distribute themselves within a heterogeneous habitat in order to maximize lifetime fitness. We test the population level consequence of the IFD theory using 40-year worth data on pike (Esox lucius) living in a natural lake divided into two basins. We do so by employing empirically derived density-dependent survival, dispersal and fecundity functions in the estimation of basin-specific density-dependent fitness surfaces. The intersection of the fitness surfaces for the two basins is used for deriving expected spatial distributions of pike. Comparing the derived expected spatial distributions with 50 years data of the actual spatial distribution demonstrated that pike is ideal free distributed within the lake. In general, there was a net migration from the less productive north basin to the more productive south basin. However, a pike density-manipulation experiment imposing shifting pike density gradients between the two basins managed to switch the net migration direction and hence clearly demonstrated that the Windermere pike choose their habitat in an ideal free manner. Demonstration of ideal free habitat selection on an operational field scale like this has never been undertaken before.

Contemporary fisherian life-history evolution in small salmonid populations.

The relative importance of natural selection and random drift in phenotypic evolution has been discussed since the introduction of the first population genetic models. The empirical evidence used to evaluate the evolutionary theories of Fisher and Wright remains obscure because formal tests for neutral divergence or sensitive attempts to separate the effects of selection and drift are scarce, subject to error, and have not been interpreted in the light of well-known population demography. We combined quantitative genetic and microsatellite DNA analyses to investigate the determinants of contemporary life-history evolution in isolated populations of grayling (Thymallus thymallus, Salmonidae) that originated from a common source 80-120 years ago. Here we show that natural selection was the dominant diversifying agent in the evolution of the quantitative traits. However, the populations were founded by a small number of individuals, exhibit very low microsatellite-based effective sizes and show genetic imprints of severe 'bottlenecks'; which are conditions often suggested to constrain selection and favour drift. This study demonstrates a very clear case of fisherian evolution in small natural populations across a contemporary timescale.