Context drives movement patterns in a mobile marine predator.

Intra-specific variability in movement behaviour occurs in all major taxonomic groups. Despite its common occurrence and ecological consequences, individual variability is often overlooked. As a result, there is a persistent gap in knowledge about drivers of intra-specific variability in movement and its role in fulfilling life history requirements. We apply a context-focused approach to bull sharks (Carcharhinus leucas), a highly mobile marine predator, incorporating intra-specific variability to understand how variable movement patterns arise and how they might be altered under future change scenarios. Spatial analysis of sharks, acoustically tagged both at their distributional limit and the centre of distribution in southern Africa, was combined with spatial analysis of acoustically tagged teleost prey and remote-sensing of environmental variables. The objective was to test the hypothesis that varying resource availability and magnitude of seasonal environmental change in different locations interact to produce variable yet predictable movement behaviours across a species' distribution. Sharks from both locations showed high seasonal overlap with predictable prey aggregations. Patterns were variable in the centre of distribution, where residency, small- and large-scale movements were all recorded. In contrast, all animals from the distributional limit performed 'leap-frog migrations', making long-distance migrations bypassing conspecifics in the centre of distribution. By combining multiple variables related to life history requirements for animals in different environments we identified combinations of key drivers that explain the occurrence of differing movement behaviours across different contexts and delineated the effects of environmental factors and prey dynamics on predator movement. Comparisons with other taxa show striking similarities in patterns of intra-specific variability across terrestrial and marine species, suggesting common drivers.

Redefining the oceanic distribution of Atlantic salmon.

Determining the mechanisms driving range-wide reductions in Atlantic salmon marine survival is hindered by an insufficient understanding of their oceanic ecology and distribution. We attached 204 pop-up satellite archival tags to post-spawned salmon when they migrated to the ocean from seven European areas and maiden North American salmon captured at sea at West Greenland. Individuals migrated further north and east than previously reported and displayed increased diving activity near oceanographic fronts, emphasizing the importance of these regions as feeding areas. The oceanic distribution differed among individuals and populations, but overlapped more between geographically proximate than distant populations. Dissimilarities in distribution likely contribute to variation in growth and survival within and among populations due to spatio-temporal differences in environmental conditions. Climate-induced changes in oceanographic conditions will alter the location of frontal areas and may have stock-specific effects on Atlantic salmon population dynamics, likely having the largest impacts on southern populations.

Selection against individuals from genetic introgression of escaped farmed salmon in a natural population of Atlantic salmon.

The viability of wild Atlantic salmon populations is threatened by genetic introgression from escaped farmed salmon. Farmed Atlantic salmon are genetically improved for important commercial traits and a life in captivity but are poorly adapted to the natural environment. The rate of gene flow from escaped farmed to wild salmon depends on their spawning success and on offspring survival at various life stages. We here investigate relative survival of introgressed juvenile Atlantic salmon (parr) in a river in northern Norway. The studied population has experienced genetic introgression from farmed salmon for about four generations (20 years). We followed two cohorts of parr from the year of hatching (0+) to the age of 2 years (2+). Farmed genetic introgression was quantified at the individual level and on a continuous scale using diagnostic SNPs. Population-level genetic introgression decreased from 0+ to 2+ by 64% (2011 cohort) and 37% (2013 cohort). This change was driven by a 70% (2011 cohort) and 49% (2013 cohort) lower survival from age 0+ to 2+ in introgressed parr compared to parr of wild origin. Our observations show that there is natural selection against genetic introgression with a potential cost of lower productivity.

Factors influencing return rate and marine residence duration in sea trout populations in Central Norway.

Brown trout (Salmo trutta) display extensive plasticity in marine migratory behaviours, with marine migrations considered to be an adaptive strategy which enables sea trout to maximize growth and reproductive potential. However, marine migrations are not without associated costs, including threats posed by ever-increasing salmon lice (Lepeophtheirus salmonis) infestations. In the present study, we used passive integrated transponder technology to characterize variability in sea trout migration behaviour amongst three catchments situated in a region of intensive salmon farming in central Norway. Specifically, we investigate how lice infestation, out-migration date and body size alter sea trout return rate and marine residence duration during the first out-migration to sea from each catchment. Distinct catchment-specific differences in sea trout out-migration size and the number of cohorts were observed, but larger body size did not guarantee the successful return of migrating trout. The marine residence duration of individuals that successfully returned to freshwater was positively correlated with lice infestation risk, suggesting for these individuals the lethal infestation threshold had not been reached. Our results also suggest that sea trout populations from lotic-dominated catchments are potentially at greater risk from size-related threats to their survival encountered during their marine migrations than sea trout from lentic-dominated catchments. The variability in sea trout migratory behaviour amongst catchments observed here emphasizes the challenges fisheries managers face when deciding the best actions to take to protect the anadromous portion of brown trout populations.

Implications of the movement behaviour of African tigerfish Hydrocynus vittatus for the design of freshwater protected areas.

African tigerfish Hydrocynus vittatus (n = 35) were tagged with external radio-transmitters in the Kavango River, Namibia, to determine whether freshwater protected areas could be an effective tool for the management and conservation of this species. They were manually tracked in the core study area of 33 km every c. 12 days from July-October 2016 to May 2017 for between 123 to 246 days. In addition, 14 extended surveys were carried out for up to 680 km to determine the total area use of the tagged individuals. Tigerfish displayed at least two behavioural patterns either having high site fidelity with shorter movements or using larger areas with longer movements. Twenty-three (66%) of the tigerfish had high site fidelity using an area of less than 33 km of river, whereas 12 tigerfish (34%) undertook long distance movements of up to 397 km upstream and 116 km downstream from their tagging locations. During the long-distance movements tigerfish crossed the territorial boundaries of Angola, Namibia and Botswana. Of the 35 fish that were monitored, 14 (40%) spent more than 80% of the monitored time in the 33 km study area and 18 (51%) stayed within the study area for at least 50% of the monitored time. These findings suggest that freshwater protected areas may be a useful management tool and we predict that a protected river area of 2-5 km river length could protect 25.9-34.6% of the population for at least 75% of the time whereas protection of 10 km river length could protect at least 50% of tigerfish for at least 75% of the time.

Gene flow from domesticated escapes alters the life history of wild Atlantic salmon.

Interbreeding between domesticated and wild animals occurs in several species. This gene flow has long been anticipated to induce genetic changes in life-history traits of wild populations, thereby influencing population dynamics and viability. Here, we show that individuals with high levels of introgression (domesticated ancestry) have altered age and size at maturation in 62 wild Atlantic salmon Salmo salar populations, including seven ancestral populations to breeding lines of the domesticated salmon. This study documents widespread changes to life-history traits in wild animal populations following gene flow from selectively bred, domesticated conspecifics. The continued high abundance of escaped, domesticated Atlantic salmon thus threatens wild Atlantic salmon populations by inducing genetic changes in fitness-related traits. Our results represent key evidence and a timely warning concerning the potential ecological impacts of the globally increasing use of domesticated animals.

Sex-dependent dominance at a single locus maintains variation in age at maturity in salmon.

Males and females share many traits that have a common genetic basis; however, selection on these traits often differs between the sexes, leading to sexual conflict. Under such sexual antagonism, theory predicts the evolution of genetic architectures that resolve this sexual conflict. Yet, despite intense theoretical and empirical interest, the specific loci underlying sexually antagonistic phenotypes have rarely been identified, limiting our understanding of how sexual conflict impacts genome evolution and the maintenance of genetic diversity. Here we identify a large effect locus controlling age at maturity in Atlantic salmon (Salmo salar), an important fitness trait in which selection favours earlier maturation in males than females, and show it is a clear example of sex-dependent dominance that reduces intralocus sexual conflict and maintains adaptive variation in wild populations. Using high-density single nucleotide polymorphism data across 57 wild populations and whole genome re-sequencing, we find that the vestigial-like family member 3 gene (VGLL3) exhibits sex-dependent dominance in salmon, promoting earlier and later maturation in males and females, respectively. VGLL3, an adiposity regulator associated with size and age at maturity in humans, explained 39% of phenotypic variation, an unexpectedly large proportion for what is usually considered a highly polygenic trait. Such large effects are predicted under balancing selection from either sexually antagonistic or spatially varying selection. Our results provide the first empirical example of dominance reversal allowing greater optimization of phenotypes within each sex, contributing to the resolution of sexual conflict in a major and widespread evolutionary trade-off between age and size at maturity. They also provide key empirical evidence for how variation in reproductive strategies can be maintained over large geographical scales. We anticipate these findings will have a substantial impact on population management in a range of harvested species where trends towards earlier maturation have been observed.

Basin-scale phenology and effects of climate variability on global timing of initial seaward migration of Atlantic salmon (Salmo salar).

Migrations between different habitats are key events in the lives of many organisms. Such movements involve annually recurring travel over long distances usually triggered by seasonal changes in the environment. Often, the migration is associated with travel to or from reproduction areas to regions of growth. Young anadromous Atlantic salmon (Salmo salar) emigrate from freshwater nursery areas during spring and early summer to feed and grow in the North Atlantic Ocean. The transition from the freshwater ('parr') stage to the migratory stage where they descend streams and enter salt water ('smolt') is characterized by morphological, physiological and behavioural changes where the timing of this parr-smolt transition is cued by photoperiod and water temperature. Environmental conditions in the freshwater habitat control the downstream migration and contribute to within- and among-river variation in migratory timing. Moreover, the timing of the freshwater emigration has likely evolved to meet environmental conditions in the ocean as these affect growth and survival of the post-smolts. Using generalized additive mixed-effects modelling, we analysed spatio-temporal variations in the dates of downstream smolt migration in 67 rivers throughout the North Atlantic during the last five decades and found that migrations were earlier in populations in the east than the west. After accounting for this spatial effect, the initiation of the downstream migration among rivers was positively associated with freshwater temperatures, up to about 10 °C and levelling off at higher values, and with sea-surface temperatures. Earlier migration occurred when river discharge levels were low but increasing. On average, the initiation of the smolt seaward migration has occurred 2.5 days earlier per decade throughout the basin of the North Atlantic. This shift in phenology matches changes in air, river, and ocean temperatures, suggesting that Atlantic salmon emigration is responding to the current global climate changes.

Does boat traffic cause displacement of fish in estuaries?

Estuaries are increasingly under threat from a variety of human impacts. Recreational and commercial boat traffic in urban areas may represent a significant disturbance to fish populations and have particularly adverse effects in spatially restricted systems such as estuaries. We examined the effects of passing boats on the abundance of different sized fish within the main navigation channel of an estuary using high resolution sonar (DIDSON). Both the smallest (100-300 mm) and largest (>501 mm) size classes had no change in their abundance following the passage of boats. However, a decrease in abundance of mid-sized fish (301-500 mm) occurred following the passage of boats. This displacement may be attributed to a number of factors including noise, bubbles and the rapidly approaching object of the boat itself. In highly urbanised estuarine systems, regular displacement by boat traffic has the potential to have major negative population level effects on fish assemblages.

Using mass scaling of movement cost and resource encounter rate to predict animal body size-population density relationships.

The negative relationship between body mass and population abundance was documented decades ago and forms one of the most fundamental scaling-laws in ecology. However, current theory fails to capture observed variations and the subject continues to raise controversy. Here we unify empirically observed size-abundance relationships with theory, by incorporating allometries in resource encounter rate and metabolic costs of movements. Fractal geometry is used to quantify the underlying resources distributions. Our model predicts that in environments packed with resources, body mass to population abundance relationships is less negative than the commonly assumed -3/4 power law. When resources are more patchily distributed, we predict a more negative exponent. These predictions are consistent with empirical observations. The current research provides an important step towards synthesizing metabolism, resource distribution and the global scaling of animal abundance, explaining why size-abundance relationships vary among feeding guilds and ecosystems.

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 degrees 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.

Crossing invisible boundaries: the effectiveness of the Langebaan Lagoon marine protected area as a harvest refuge for a migratory fish species in South Africa.

The application of no-take areas in fisheries remains controversial. Critics argue that many targeted species are too mobile to benefit from area protection and that no-take areas are only appropriate for resident species. The degree of protection does not depend on the size of the no-take area but rather on the time fish reside inside its boundaries during key life-history events (i.e., spawning) and during periods of peak fishing activity. We evaluated the potential of a small no-take marine protected area (MPA) inside a coastal embayment as a harvest refuge for a mobile, possibly migratory, long-lived fish species. We used acoustic telemetry to track movements of 30 transmitter-tagged white stumpnose (Rhabdosargus globiceps) across and on both sides of the boundary of a small (34 km(2)) no-take area over a full year. Being landlocked on 3 sides, the location of the MPA inside the lagoon made it practical to detect all boundary crossings and to calculate the time individual fish used the MPA. We detected frequent movements across the boundary, with strong seasonal and individual variations. There were significant differences in MPA use patterns between fish from different release areas. The time spent in the MPA by individual fish during summer (mean 50%; max 98%) was out of proportion with the size of that area (4% of total habitat). Summer coincided with peak recreational fishing activity and with the spawning season of this species. The small MPA provided a refuge for a part of the spawning stock of white stumpnose. Our findings suggest that if strategically placed, a small no-take area can be effective in protecting mobile species and that models of spillover from no-take areas should account for seasonal and individual variation in area use and the spatiotemporal distribution of fish and fishers.