 – Among the species in the family Salmonidae, those represented by the genera Salmo, Salvelinus, and Oncorhynchus (subfamily Salmoninae) are the most studied. Here, various aspects of phenotypic and life-history variation of Atlantic salmon Salmo salar L., brown trout Salmo trutta L., and Arctic charr Salvelinus alpinus (L.) are reviewed. While many strategies and tactics are commonly used by these species, there are also differences in their ecology and population dynamics that result in a variety of interesting and diverse topics that are challenging for future research. Atlantic salmon display considerable phenotypic plasticity and variability in life-history characters ranging from fully freshwater resident forms, where females can mature at approximately 10 cm in length, to anadromous populations characterised by 3–5 sea-winter (5SW) salmon. Even within simple 1SW populations, 20 or more spawning life-history types can be identified. Juveniles in freshwater can use both fluvial and lacustrine habitats for rearing, and while most smolts migrate to sea during the spring, fall migrations occur in some populations. At sea, some salmon undertake extensive oceanic migrations while other populations stay within the geographical confines of areas such as the Baltic Sea. At the other extreme are those that reside in estuaries and return to freshwater to spawn after spending only a few months at sea. The review of information on the diversity of life-history forms is related to conservation aspects associated with Atlantic salmon populations and current trends in abundance and survival. Brown trout is indigenous to Europe, North Africa and western Asia, but was introduced into at least 24 countries outside Europe and now has a world-wide distribution. It exploits both fresh and salt waters for feeding and spawning (brackish), and populations are often partially migratory. One part of the population leaves and feeds elsewhere, while another part stays as residents. In large, complex systems, the species is polymorphic with different size morphs in the various parts of the habitat. Brown trout feed close to the surface and near shore, but large individuals may move far offshore. The species exhibits ontogenetic niche shifts partly related to size and partly to developmental rate. They switch when the amount of surplus energy available for growth becomes small with fast growers being younger and smaller fish than slow growers. Brown trout is an opportunistic carnivore, but individuals specialise at least temporarily on particular food items; insect larvae are important for the young in streams, while littoral epibenthos in lakes and fish are most important for large trout. The sexes differ in resource use and size. Females are more inclined than males to become migratory and feed in pelagic waters. Males exploit running water, near-shore and surface waters more than females. Therefore, females feed more on zooplankton and exhibit a more uniform phenotype than males. The Arctic charr is the northernmost freshwater fish on earth, with a circumpolar distribution in the Holarctic that matches the last glaciation. Recent mtDNA studies indicate that there are five phylogeographic lineages (Atlantic, Arctic, Bering, Siberian and Acadian) that may be of Pleistocene origin. Phenotypic expression and ecology are more variable in charr than in most fish. Weights at maturation range from 3 g to 12 kg. Population differences in morphology and coloration are large and can have some genetic basis. Charr live in streams, at sea and in all habitats of oligotrophic lakes, including very deep areas. Ontogenetic habitat shifts between lacustrine habitats are common. The charr feed on all major prey types of streams, lakes and near-shore marine habitats, but has high niche flexibility in competition. Cannibalism is expressed in several cases, and can be important for developing and maintaining bimodal size distributions. Anadromy is found in the northern part of its range and involves about 40, but sometimes more days in the sea. All charr overwinter in freshwater. Partial migration is common, but the degree of anadromy varies greatly among populations. The food at sea includes zooplankton and pelagic fish, but also epibenthos. Polymorphism and sympatric morphs are much studied. As a prominent fish of glaciated lakes, charr is an important species for studying ecological speciation by the combination of field studies and experiments, particularly in the fields of morphometric heterochrony and comparative behaviour.

https://onlinelibrary.wiley.com/doi/pdf/10.1034/j.1600-0633.2003.00010.x

Atlantic salmon Salmo salar L., brown trout Salmo trutta L. and Arctic charr Salvelinus alpinus (L.): a review of aspects of their life histories

Habitat requirements of Atlantic salmon and brown trout in rivers and streams

The present paper reviews the effects of water temperature and flow on migrations, embryonic development, hatching, emergence, growth and life-history traits in light of the ongoing climate change with emphasis on anadromous Atlantic salmon Salmo salar and brown trout Salmo trutta. The expected climate change in the Atlantic is for milder and wetter winters, with more precipitation falling as rain and less as snow, decrease in ice-covered periods and frequent periods with extreme weather. Overall, thermal limits for salmonids are species specific. Scope for activity and growth and optimal temperature for growth increase with temperature to an optimal point before constrain by the oxygen content of the water. The optimal temperature for growth decreases with increasing fish size and varies little among populations within species, whereas the growth efficiency may be locally adapted to the temperature conditions of the home stream during the growth season. Indirectly, temperature influences age and size at smolting through its effect on growth. Time of spawning, egg hatching and emergence of the larvae vary with temperature and selective effects on time of first feeding. Traits such as age at first maturity, longevity and fecundity decrease with increasing temperature whilst egg size increases with temperature. Water flow influences the accessibility of rivers for returning adults and speed of both upstream and downstream migration. Extremes in water flow and temperature can decrease recruitment and survival. There is reason to expect a northward movement of the thermal niche of anadromous salmonids with decreased production and population extinction in the southern part of the distribution areas, migrations earlier in the season, later spawning, younger age at smolting and sexual maturity and increased disease susceptibility and mortality. Future research challenges are summarized at the end of the paper.

A review of the likely effects of climate change on anadromous Atlantic salmon Salmo salar and brown trout Salmo trutta, with particular reference to water temperature and flow.

Here we critically review the scale and extent of adaptive genetic variation in Atlantic salmon (Salmo salar L.), an important model system in evolutionary and conservation biology that provides fundamental insights into population persistence, adaptive response and the effects of anthropogenic change. We consider the process of adaptation as the end product of natural selection, one that can best be viewed as the degree of matching between phenotype and environment. We recognise three potential sources of adaptive variation: heritable variation in phenotypic traits related to fitness, variation at the molecular level in genes influenced by selection, and variation in the way genes interact with the environment to produce phenotypes of varying plasticity. Of all phenotypic traits examined, variation in body size (or in correlated characters such as growth rates, age of seaward migration or age at sexual maturity) generally shows the highest heritability, as well as a strong effect on fitness. Thus, body size in Atlantic salmon tends to be positively correlated with freshwater and marine survival, as well as with fecundity, egg size, reproductive success, and offspring survival. By contrast, the fitness implications of variation in behavioural traits such as aggression, sheltering behaviour, or timing of migration are largely unknown. The adaptive significance of molecular variation in salmonids is also scant and largely circumstantial, despite extensive molecular screening on these species. Adaptive variation can result in local adaptations (LA) when, among other necessary conditions, populations live in patchy environments, exchange few or no migrants, and are subjected to differential selective pressures. Evidence for LA in Atlantic salmon is indirect and comes mostly from ecological correlates in fitness-related traits, the failure of many translocations, the poor performance of domesticated stocks, results of a few common-garden experiments (where different populations were raised in a common environment in an attempt to dissociate heritable from environmentally induced phenotypic variation), and the pattern of inherited resistance to some parasites and diseases. Genotype x environment interactions occurr for many fitness traits, suggesting that LA might be important. However, the scale and extent of adaptive variation remains poorly understood and probably varies, depending on habitat heterogeneity, environmental stability and the relative roles of selection and drift. As maladaptation often results from phenotype-environment mismatch, we argue that acting as if populations are not locally adapted carries a much greater risk of mismanagement than acting under the assumption for local adaptations when there are none. As such, an evolutionary approach to salmon conservation is required, aimed at maintaining the conditions necessary for natural selection to operate most efficiently and unhindered. This may require minimising alterations to native genotypes and habitats to which populations have likely become adapted, but also allowing for population size to reach or extend beyond carrying capacity to encourage competition and other sources of natural mortality.

A critical review of adaptive genetic variation in Atlantic salmon: implications for conservation

Aquatic Ecosystem Health and Management

– Habitat is important in determining stream carrying capacity and population density in young Atlantic salmon and brown trout. We review stream habitat selection studies and relate results to variable and interacting abiotic and biotic factors. The importance of spatial and temporal scales are often overlooked. Different physical variables may influence fish position choice at different spatial scales. Temporally variable water flows and temperatures are pervasive environmental factors in streams that affect behavior and habitat selection. The more frequently measured abiotic variables are water depth, water velocity (or stream gradient), substrate particle size, and cover. Summer daytime, feeding habitats of Atlantic salmon are size structured. Larger parr (>7 cm) have a wider spatial niche than small parr. Selected snout water velocities are consistently low (3–25 cm. s−1). Mean (or surface) water velocities are in the preferred range of 30–50 cm. s−1, and usually in combination with coarse substratum (16–256 mm). However, salmon parr demonstrate flexibility with respect to preferred water velocity, depending on fish size, intra- and interspecific competition, and predation risk. Water depth is less important, except in small streams. In large rivers and lakes a variety of water depths are used by salmon parr. Summer daytime, feeding habitat of brown trout is also characterized by a narrow selection of low snout water velocities. Habitat use is size-structured, which appears to be mainly a result of intraspecific competition. The small trout parr (<7 cm) are abundant in the shallow swift stream areas (<20–30 cm depths, 10–50 cm. s−1 water velocities) with cobble substrates. The larger trout have increasingly strong preferences for deep-slow stream areas, in particular pools. Water depth is considered the most important habitat variable for brown trout. Spatial niche overlap is considerable where the two species are sympatric, although young Atlantic salmon tend to be distributed more in the faster flowing and shallow habitats compared with trout. Habitat use by salmon is restricted through interspecific competition with the more aggressive brown trout (interactive segregation). However, subtle innate differences in behavior at an early stage also indicate selective segregation. Seasonal changes in habitat use related to water temperatures occur in both species. In winter, they have a stronger preference for cover and shelter, and may seek shelter in the streambed and/or deeper water. At low temperatures (higher latitudes), there are also marked shifts in habitat use during day and night as the fish become nocturnal. Passive sheltering in the substrate or aggregating in deep-slow stream areas is the typical daytime behavior. While active at night, the fish move to more exposed holding positions primarily on but also above the substrate. Diurnal changes in habitat use take place also in summer; brown trout may utilize a wider spatial niche at night with more fish occupying the shallow-slow stream areas. Brown trout and young Atlantic salmon also exhibit a flexible response to variability in streamflows, wherein habitat selection may change considerably. Important topics in need of further research include: influence of spatial measurement scale, effects of temporal and spatial variability in habitat conditions on habitat selection, effects of interactive competition and trophic interactions (predation risk) on habitat selection, influence of extreme natural events on habitat selection use or suitability (floods, ice formation and jams, droughts), and individual variation in habitat use or behavior.

J. Heggenes

Papers

Spatial niche variability for young Atlantic salmon (Salmo salar) and brown trout (S. trutta) in heterogeneous streams