Comparing dominance relationships and movement of native marble trout (Salmo marmoratus) and introduced rainbow trout (Oncorhynchus mykiss).

Behaviour observations of the endangered native marble trout (Salmo marmoratus, Cuvier, 1829) and introduced rainbow trout (Oncorhynchus mykiss, Walbaum, 1792) in the laboratory and in a tributary to the Idrijca River in Slovenia were combined to study the movements and dominance relationships between individuals of the two species in an open field test. Under laboratory conditions, no difference between the species was detected for neither time spent actively or distance moved. In species paired tests, rainbow trout initiated more aggressive behaviours towards marble trout than vice versa, and rainbow trout were clearly the dominant individuals. After simultaneous release in the river, marble trout immediately left the release area and spent twice as long time as rainbow trout until they settled in an area of the river; therefore, the release site was immediately occupied exclusively by rainbow trout. Thus, the dominant and aggressive behaviour of rainbow trout seen in the laboratory before release might have influenced marble trout's subsequent behaviour in the river, by marble trout leaving the areas occupied by rainbow trout and moving to locations further away from the release site. In the field, the marble trout occupied sites individually, whereas rainbow aggregated at a few locations. Rainbow trout showed higher movement activity in the morning compared to marble trout. There was a positive correlation between swimming speed in the laboratory and movement in the field for marble trout but not for rainbow trout. In conclusion, the results in this study support the need to end stocking of rainbow trout in rivers with native marble trout. To better understand the interaction between the species, and to develop efficient management plans to protect the native marble trout, reference behaviours should first be understood, and future research in sites where the two species do not co-exist is needed. This is especially important for marble trout for which behavioural research and data are lacking.

Linking omega-3 polyunsaturated fatty acids in natural diet with brain size of wild consumers.

Omega-3 long-chain polyunsaturated fatty acids (n-3 LC-PUFA) are key structural lipids and their dietary intake is essential for brain development of virtually all vertebrates. The importance of n-3 LC-PUFA has been demonstrated in clinical and laboratory studies, but little is known about how differences in the availability of n-3 LC-PUFA in natural prey influence brain development of wild consumers. Consumers foraging at the interface of aquatic and terrestrial food webs can differ substantially in their intake of n-3 LC-PUFA, which may lead to differences in brain development, yet this hypothesis remains to be tested. Here we use the previously demonstrated shift towards higher reliance on n-3 LC-PUFA deprived terrestrial prey of native brown trout Salmo trutta living in sympatry with invasive brook trout Salvelinus fontinalis to explore this hypothesis. We found that the content of n-3 LC-PUFA in muscle tissues of brown trout decreased with increasing consumption of n-3 LC-PUFA deprived terrestrial prey. Brain volume was positively related to the content of the n-3 LC-PUFA, docosahexaenoic acid, in muscle tissues of brown trout. Our study thus suggests that increased reliance on diets low in n-3 LC-PUFA, such as terrestrial subsidies, can have a significant negative impact on brain development of wild trout. Our findings provide the first evidence of how brains of wild vertebrate consumers response to scarcity of n-3 LC-PUFA content in natural prey.

Asymmetric competition over space use and territory between native brown trout (Salmo trutta) and invasive brook trout (Salvelinus fontinalis).

Interference competition over food and territory can shape population structure and habitat use within and between species. The introduction of invasive species often leads to novel competitive interactions over shared resources and invaders can eventually exclude the native species from preferred habitats. Invasive brook trout (Salvelinus fontinalis) introduced to northern Europe have excluded native brown trout (Salmo trutta) from numerous headwater streams. The fact that invasive brook trout can displace the more aggressive brown trout is puzzling. However, the earlier spawning and hatching of brook trout, compared to brown trout, may lead to unequal competition due to size advantage and prior resident status of brook trout at the fry stage. In this study, we examine the effect of competition between brown trout and brook trout using the natural size distribution of the two species. In two consecutive experiments, we first measured space use and feeding of a fry (age 0+) in the presence of a juvenile (age 1+). In experiment 2, we assessed territorial interactions between the species at the fry stage (age 0+) and if smaller brown trout could compensate the disadvantage by manipulating residence duration. Fry of brook trout feed sooner and spend more time close to the larger individual than brown trout fry. We also found that brook trout fry won most territorial contests against brown trout, and that increased residence duration led to longer and more aggressive interactions. The results suggest that smaller brown trout are displaced to suboptimal habitats in the presence of a larger brook trout. Therefore, the later emergence from gravel beds resulting in the naturally occurring size disadvantage of brown trout at the fry stage may lead to unequal territorial interactions that could explain why brown trout are displaced from preferred habitats in sympatry with brook trout.

Northern European Salmo trutta (L.) populations are genetically divergent across geographical regions and environmental gradients.

The salmonid fish Brown trout is iconic as a model for the application of conservation genetics to understand and manage local interspecific variation. However, there is still scant information about relationships between local and large-scale population structure, and to what extent geographical and environmental variables are associated with barriers to gene flow. We used information from 3,782 mapped SNPs developed for the present study and conducted outlier tests and gene-environment association (GEA) analyses in order to examine drivers of population structure. Analyses comprised >2,600 fish from 72 riverine populations spanning a central part of the species' distribution in northern Europe. We report hitherto unidentified genetic breaks in population structure, indicating strong barriers to gene flow. GEA loci were widely spread across genomic regions and showed correlations with climatic, abiotic and geographical parameters. In some cases, individual loci showed consistent GEA across the geographical regions Britain, Europe and Scandinavia. In other cases, correlations were observed only within a sub-set of regions, suggesting that locus-specific variation was associated with local processes. A paired-population sampling design allowed us to evaluate sampling effects on detection of outlier loci and GEA. Two widely applied methods for outlier detection (pcadapt and bayescan) showed low overlap in loci identified as statistical outliers across sub-sets of data. Two GEA analytical approaches (LFMM and RDA) showed good correspondence concerning loci associated with specific variables, but LFMM identified five times more statistically significant associations than RDA. Our results emphasize the importance of carefully considering the statistical methods applied for the hypotheses being tested in outlier analysis. Sampling design may have lower impact on results if the objective is to identify GEA loci and their population distribution. Our study provides new insights into trout populations, and results have direct management implications in serving as a tool for identification of conservation units.

Behavioral type, in interaction with body size, affects the recapture rate of brown trout Salmo trutta juveniles in their nursery stream.

Movement activity levels of wild animals often differ consistently among individuals, reflecting different behavioral types. Previous studies have shown that laboratory-scored activity can predict several ecologically relevant characteristics. In an experiment on wild brown trout Salmo trutta, spanning from June to October, we investigated how spring swimming activity, measured in a standardized laboratory test, related to relative recapture probability in autumn. Based on laboratory activity scores, individuals clustered into 2 groups, which showed contrasting patterns in the size-dependency of their recapture probability. Size had a slightly positive effect on recapture probability for passive fish but a clear negative effect on active fish. Our results show that the population structure in a cohort, in terms of relative proportions of behavioral types in different size classes, can vary over time. The results of this study could depend on either selective mortality or migration. However, selective disappearance of individuals with specific phenotypes, regardless of the mechanism, will have implications for trout population management, such as stocking efficiency of hatchery fish with high growth rates or maintenance of fishways past migration barriers.

Encystment of parasitic freshwater pearl mussel (Margaritifera margaritifera) larvae coincides with increased metabolic rate and haematocrit in juvenile brown trout (Salmo trutta).

Gill parasites on fish are likely to negatively influence their host by inhibiting respiration, oxygen transport capacity and overall fitness. The glochidia larvae of the endangered freshwater pearl mussel (FPM, Margaritifera margaritifera (Linnaeus, 1758)) are obligate parasites on the gills of juvenile salmonid fish. We investigated the effects of FPM glochidia encystment on the metabolism and haematology of brown trout (Salmo trutta Linnaeus, 1758). Specifically, we measured whole-animal oxygen uptake rates at rest and following an exhaustive exercise protocol using intermittent flow-through respirometry, as well as haematocrit, in infested and uninfested trout. Glochidia encystment significantly affected whole-animal metabolic rate, as infested trout exhibited higher standard and maximum metabolic rates. Furthermore, glochidia-infested trout also had elevated levels of haematocrit. The combination of an increased metabolism and haematocrit in infested fish indicates that glochidia encystment has a physiological effect on the trout, perhaps as a compensatory response to the potential respiratory stress caused by the glochidia. When relating glochidia load to metabolism and haematocrit, fish with low numbers of encysted glochidia were the ones with particularly elevated metabolism and haematocrit. Standard metabolic rate decreased with substantial glochidia loads towards levels similar to those of uninfested fish. This suggests that initial effects visible at low levels of encystment may be countered by additional physiological effects at high loads, e.g. potential changes in energy utilization, and also that high numbers of glochidia may restrict oxygen uptake by the gills.

Inactive trout come out at night: behavioral variation, circadian activity, and fitness in the wild.

Theory suggests that high activity levels in animals increase growth at the cost of increased mortality. This growth-mortality tradeoff has recently been incorporated into the wider framework of the pace-of-life syndrome (POLS) hypothesis. However, activity is often quantified only in the laboratory and on a diurnal basis, leaving open the possibility that animals manage predation risk and feeding efficiency in the wild by modulating their circadian activity rhythms. Here we investigate how laboratory activity in wild brown trout parr (Salmo trutta L.) associates with circadian activity, growth, and mortality in their natal stream. We found that individuals with high activity in the laboratory displayed high dispersal and cathemeral activity in their natal stream. In contrast, trout with low laboratory activity showed variation of activity in the wild, which was negatively related to the light intensity. Our results do not support the growth-mortality trade-off of the POLS hypothesis as highly active, fast-growing individuals showed higher survival than inactive conspecifics. These novel results show for the first time that active and inactive individuals, as scored in the lab, can show different circadian patterns of behavior in the wild driven by light intensity. This implies that studies conducted under a narrow range of light conditions can bias our understanding of individual behavioral variation and its fitness consequences in the wild.

Sight or smell? Behavioural and heart rate responses in subordinate rainbow trout exposed to cues from dominant fish.

Many animals, including fish, can utilize both vision and the chemical senses in intra-specific communication. However, the relative influence of these sensory modalities on behavioral and physiological responses in social interactions is not well understood. The aim of this study was therefore to investigate the relative effects of visual and chemical stimuli from dominant individuals on the behavioral and physiological responses of subordinate rainbow trout (Oncorhynchus mykiss). External electrodes were used to detect ECG signals from free-swimming fish. This method allowed the simultaneous recording of behavioral and physiological responses, and possible sex differences in these responses were also investigated. The results suggest that, in this context, visual cues are more important than chemical cues in settling the social hierarchy in rainbow trout because a combination of chemical and visual exposure generally yielded a response in focal fish that was similar to the response elicited by visual exposure alone. Both activity and physiological responses were most pronounced during the first ten seconds after exposure, with subordinate fish moving closer to the dominant, accompanied by a strong bradycardic response. Furthermore, females acted more boldly and moved closer to the dominant fish than males, but here the effect of the modes was additive, with a stronger effect of the combined visual and chemical exposure. Overall, the extra information furnished to the fish in the form of chemical cues did not change either the behavioral or the physiological response. This result suggests that visual cues are more important than chemically mediated ones for social communication and individual recognition in rainbow trout.

Habitat exclusion and reduced growth: a field experiment on the effects of inter-cohort competition in young-of-the-year brown trout.

Competition during the juvenile phase is a key process for regulating density in organisms with high fecundity. Juvenile density-dependent bottlenecks may become even more pronounced if several cohorts compete, but this has received relatively limited attention in previous literature. We performed a manipulation experiment in seven coastal streams to investigate the presence of inter-cohort competition, using habitat selection, body-size and density of newly emerged (age-0) brown trout (Salmo trutta) as response variables. The trout population (age ≥ 1 fish) was estimated using electro-fishing prior to the emergence of fry (April-May) and was either removed (manipulated sections) or maintained (control sections). Age-0 habitat selection was examined in June while density and body-size was evaluated in October (end of the growth season). We found that age-0 trout selected habitats that were located further from riffles (nursery habitats) in the absence of age ≥ 1 trout, suggesting a niche overlap between cohorts in the habitat dimension and, hence, that both inter-cohort competitive interactions and ontogenetic preference may influence habitat utilisation in the wild. Furthermore, we also found age-0 body-size to be significantly larger in manipulated sections and negatively related to its own density. We argue that competition from older cohorts influence the availability of age-0 feeding territories at the critical phase of emergence with secondary negative effects on age-0 growth. These results not only have implications for understanding the mechanisms of density dependence but can also provide valuable knowledge to the management of salmonid populations and their habitats in the wild.