Most empirical and theoretical studies of resource use and population dynamics treat conspecific individuals as ecologically equivalent. This simplification is only justified if interindividual niche variation is rare, weak, or has a trivial effect on ecological processes. This article reviews the incidence, degree, causes, and implications of individual-level niche variation to challenge these simplifications. Evidence for individual specialization is available for 93 species dis- tributed across a broad range of taxonomic groups. Although few studies have quantified the degree to which individuals are specialized relative to their population, between-individual variation can some- times comprise the majority of the population's niche width. The degree of individual specialization varies widely among species and among populations, reflecting a diverse array of physiological, be- havioral, and ecological mechanisms that can generate intrapopu- lation variation. Finally, individual specialization has potentially im- portant ecological, evolutionary, and conservation implications. Theory suggests that niche variation facilitates frequency-dependent interactions that can profoundly affect the population's stability, the amount of intraspecific competition, fitness-function shapes, and the population's capacity to diversify and speciate rapidly. Our collection of case studies suggests that individual specialization is a widespread but underappreciated phenomenon that poses many important but unanswered questions.

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The Ecology of Individuals: Incidence and Implications of Individual Specialization

Summary
1. The thermal regime of rivers plays an important role in the overall health of aquatic ecosystems, including water quality issues and the distribution of aquatic species within the river environment. Consequently, for conducting environmental impact assessments as well as for effective fisheries management, it is important to understand the thermal behaviour of rivers and related heat exchange processes.

2. This study reviews the different river thermal processes responsible for water temperature variability on both the temporal (e.g. diel, daily, seasonal) and spatial scales, as well as providing information related to different water temperature models currently found in the literature.

3. Water temperature models are generally classified into three groups: regression, stochastic and deterministic models. Deterministic models employ an energy budget approach to predict river water temperature, whereas regression and stochastic models generally rely on air to water temperature relationships.

4. Water temperature variability can occur naturally or as a result of anthropogenic perturbations, such as thermal pollution, deforestation, flow modification and climate change. Literature information is provided on the thermal regime of rivers in relation to anthropogenic impacts and such information will contribute to the better protection of fish habitat and more efficient fisheries management.

https://onlinelibrary.wiley.com/doi/pdf/10.1111/j.1365-2427.2006.01597.x

The thermal regime of rivers : a review

 – 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

Publisher Summary This chapter reviews the present empirical and theoretical work on antipredator decision making. The ways in which predators influence the behavioral decisions made by their prey is now the subject of a large and growing literature. Some notable recent advances include clear demonstrations that antipredatory decision making (1) may influence many aspects of reproductive behavior, (2) has demonstrable long-term consequences for individual fitness, and (3) may influence the nature of ecological systems themselves. There have also been many advances in the theory of antipredator behavior, which should provide a sound conceptual basis for further progress. Attention is needed for further work on the effects that predator and prey have on the other's behavioral decisions. The range of reproductive behaviors influenced by the risk of predation also requires much more investigation. Work on the long-term costs of antipredator decision making needs more empirical documentation and greater taxonomic diversity. Work on the ecological implications of antipredatory decision making has only scratched the surface, especially with regard to population-level effects and species interactions. Theoretical investigations should also play a prominent role in future work.

Stress and Decision Making under the Risk of Predation: Recent Developments from Behavioral, Reproductive, and Ecological Perspectives

The accuracy of gene flow estimates is unknown in most natural populations because direct estimates of dispersal are often not possible. These estimates can be highly imprecise or even biased because population genetic structure reflects more than a simple balance between genetic drift and gene flow. Most of the models used to estimate gene flow also assume very simple patterns of movement. As a result, multiple interpretations of population structure involving contemporary gene flow, departures from equilibrium, and other factors are almost always possible. One way to isolate the relative contribution of gene flow to population genetic differentiation is to utilize comparative methods. Population genetic statistics such as FST, heterozygosity and Nei's D can be compared between species with differing dispersal abilities if these species are otherwise phylogenetically, geographically and demographically comparable. Accordingly, the available literature was searched for all groups that meet these criteria ...

Dispersal, gene flow, and population structure

Many fish populations have both resident and migratory individuals. Migrants usually grow larger and have higher reproductive potential but lower survival than resident conspecifics. The ‘decision’ about migration versus residence probably depends on the individual growth rate, or a physiological process like metabolic rate which is correlated with growth rate. Fish usually mature as their somatic growth levels off, where energetic costs of maintenance approach energetic intake. After maturation, growth also stagnates because of resource allocation to reproduction. Instead of maturation, however, fish may move to an alternative feeding habitat and their fitness may thereby be increased. When doing so, maturity is usually delayed, either to the new asymptotic length, or sooner, if that gives higher expected fitness. Females often dominate among migrants and males among residents. The reason is probably that females maximize their fitness by growing larger, because their reproductive success generally increases exponentially with body size. Males, on the other hand, may maximize fitness by alternative life histories, e.g. fighting versus sneaking, as in many salmonid species where small residents are the sneakers and large migrants the fighters. Partial migration appears to be partly developmental, depending on environmental conditions, and partly genetic, inherited as a quantitative trait influenced by a number of genes.

Partial migration: niche shift versus sexual maturation in fishes

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.

Farm Atlantic salmon escape and invade rivers throughout the North Atlantic annually, which has generated growing concern about their impacts on native salmon populations. A large–scale experiment was therefore undertaken in order to quantify the lifetime success and interactions of farm salmon invading a Norwegian river. Sexually mature farm and native salmon were genetically screened, radio tagged and released into the River Imsa where no other salmon had been allowed to ascend. The farm fishes were competitively and reproductively inferior, achieving less than one–third the breeding success of the native fishes. Moreover, this inferiority was sex biased, being more pronounced in farm males than females, resulting in the principal route of gene flow involving native males mating with farm females. There were also indications of selection against farm genotypes during early survival but not thereafter. However, evidence of resource competition and competitive displacement existed as the productivity of the native population was depressed by more than 30%. Ultimately, the lifetime reproductive success (adult to adult) of the farm fishes was 16% that of the native salmon. Our results indicate that such annual invasions have the potential for impacting on population productivity, disrupting local adaptations and reducing the genetic diversity of wild salmon populations.

Lifetime success and interactions of farm salmon invading a native population.

The Arctic charr Salvelinus alpinus exhibits 1–4 sympatric morphs in postglacial lakes, of which one or two are epibenthic zoobenthos feeders, one is a limnetic planktivore and one is a piscivorous form. In addition, northern rivers support partly migratory populations with anadromous and freshwater resident fish. The morphs vary in their coloration, morphology, life history, behaviour and genetic characteristics. The morphs usually differentiate according to their ontogenetic stage at maturity, which parallels paedomorphism in amphibians. The young usually start as epibenthic zoobenthivores, but may become pelagic at a certain size according to the predation risk experienced at that time. From a length of > 20–25 cm, charr start to become piscivorous. The proportion of piscivorous fish increases with increasing body size. In partly anadromous populations, fish that mature before smolting become freshwater resident, the others anadromous. In some rivers, the morphs occupy separate niches (epibenthic and limnetic), from emergence onwards. The morphs exhibit different degrees of reproductive isolation that vary from a high degree of interbreeding to complete isolation. Usually, they spawn within morph (assortative mating), but alternative male mating behaviour (sneaking, fighting) may occur in stream-spawning populations and at great depths in lakes. Morphologically specialized morphs appear to feed more effectively than intermediate forms, and selection according to feeding mode, site fidelity and associated assortative mating are prerequisites for the evolution of the different morphs. Charr morphs develop into stable feeding niches under conditions of intense intraspecific competition when there is little competition with other species. Sympatric morphs exhibit different degrees of speciation, but similar morphs in different systems are not individual species because of (1) their polyphyletic origin, (2) the supporting systems are often young, transient environments making the future situation for the populations uncertain, and (3) the genetic differentiation among morphs is low. Sympatric morphs may interbreed and produce fertile hybrids. Nevertheless, sympatric charr morphs should be managed as separate species. Changes in the natural conditions or human impacts to which the morphs are adapted will have a strong influence on the persistence and survival of each different morph.

Bror Jonsson

Papers

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

Partial migration: niche shift versus sexual maturation in fishes

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.

Lifetime success and interactions of farm salmon invading a native population.

Polymorphism and speciation in Arctic charr