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

Abstract – 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.     

Parentally acquired differences in resource acquisition ability between brown trout from alternative life history parentage

Dominance hierarchies, where they exist, affect individual food acquisition ability and fitness, both of which have the potential to influence life history pathways. Juvenile salmonids exhibit clear dominance hierarchies in early life. As one of the drivers for the adoption of alternative life histories in salmonids is the relative rate of resource acquisition, there is potential for juvenile behaviour to influence the subsequent life history strategy of the individual. Lacustrine brown trout, Salmo trutta, exhibit a multitude of life histories which includes among others the piscivorous (ferox) life history where individuals grow to large size and have delayed maturity and benthivorous and pelagic life histories where individuals grow to much smaller sizes, however mature earlier. Using a number of observable characteristics of dominance, this study compared differences in behaviour between size‐matched pairs of progeny, reared under common garden conditions which are derived from alternative, co‐existing life history strategy parents. We found that first‐generation progeny of ferox trout were more aggressive, acquired more food, had lighter skin pigmentation and held more desirable positions than the progeny of benthivorous brown trout in an experimental stream system. Ferox trout progeny were dominant over benthivorous brown trout progeny in 90% of trials in dyadic contests. Given such clear differences in dominance, this study indicates that parentally acquired dominance‐related differences, passed through either, or both, of genetic and nongenetic (e.g. maternal effects) means, are likely a contributing factor to the continued maintenance of distinct life history strategies of brown trout.     

Alternative routes to piscivory: Contrasting growth trajectories in brown trout (Salmo trutta) ecotypes exhibiting contrasting life history strategies

Large and long‐lived piscivorous brown trout, Salmo trutta, colloquially known as ferox trout, have been described from a number of oligotrophic lakes in Britain and Ireland. The “ferox” life history strategy is associated with accelerated growth following an ontogenetic switch to piscivory and extended longevity (up to 23 years in the UK). Thus, ferox trout often reach much larger sizes and older ages than sympatric lacustrine invertebrate‐feeding trout. Conventional models suggest that S. trutta adopting this life history strategy grow slowly before a size threshold is reached, after which, this gape‐limited predator undergoes a diet switch to a highly nutritional prey source (fish) resulting in a measurable growth acceleration. This conventional model of ferox trout growth was tested by comparing growth trajectories and age structures of ferox trout and sympatric invertebrate‐feeding trout in multiple lake systems in Scotland. In two of the three lakes examined, fish displaying alternative life history strategies, but living in sympatry, exhibited distinctly different growth trajectories. In the third lake, a similar pattern of growth was observed between trophic groups. Piscivorous trout were significantly older than sympatric invertebrate‐feeding trout at all sites, but ultimate body size was greater in only two of three sites. This study demonstrates that there are multiple ontogenetic growth pathways to achieving piscivory in S. trutta and that the adoption of a piscivorous diet may be a factor contributing to the extension of lifespan.     

Total population density during the first year of life as a major determinant of lifetime body‐length trajectory in marble trout

Abstract – The conditions experienced early in life can strongly influence life‐history trajectories in a variety of animal species. Here, we use data from four isolated populations of the endangered stream‐dwelling salmonid marble trout (Salmo marmoratus Cuvier 1817) living in the Soca and Idrijca river basins (Slovenia) to explore the influence of the total density experienced during and after the first year of life by marble trout year‐classes on body length of marble trout through the lifetime. Analyses were performed by pooling together the stream‐specific datasets to cover a wider range of densities. Mean body length of marble trout year‐classes through the lifetime (from age 1+ to 5+) was negatively related to total density of marble trout during the first year of life. The relationship between density during the first growth period and body length through the lifetime was well described by negative power curves. Total population density after the first year of life was not correlated with body length, thus suggesting that body growth trajectories are heavily determined early in life. Given size‐dependent sexual maturity and egg production in marble trout, the relationship between density early in life and lifetime individual growth may have strong implications in terms of population dynamics and regulation of population size.     

Fish life history,angler behaviour and optimal management of recreational fisheries

To predict recreational‐fishing impacts on freshwater fish species, it is important to understand the interplay between fish populations, anglers and management actions. We use an integrated bioeconomic model to study the importance of fish life‐history type (LHT) for determining (i) vulnerability to over‐exploitation by diverse angler types (generic, consumptive and trophy anglers), who respond dynamically to fishing‐quality changes; (ii) regulations [i.e., minimum‐size limits (MSLs) and licence densities] that maximize the social welfare of angler populations; and (iii) biological and social conditions resulting under such socially optimal regulations. We examine five prototypical freshwater species: European perch (Perca fluviatilis), brown trout (Salmo trutta), pikeperch (Sander lucioperca), pike (Esox lucius) and bull trout (Salvelinus confluentus). We find that LHT is important for determining the vulnerability of fish populations to overfishing, with pike, pikeperch, and bull trout being more vulnerable than perch and brown trout. Angler type influences the magnitude of fishing impacts, because of differences in fishing practices and angler‐type‐specific effects of LHT on angling effort. Our results indicate that angler types are systematically attracted to particular LHTs. Socially optimal minimum‐size limits generally increase with LHT vulnerability, whereas optimal licence densities are similar across LHTs. Yet, both regulations vary among angler types. Despite this variation, we find that biological sustainability occurs under socially optimal regulations, with one exception. Our results highlight the importance of jointly considering fish diversity, angler diversity and regulations when predicting sustainable management strategies for recreational fisheries. Failure to do so could result in socially suboptimal management and/or fishery collapse.     

Differences in early developmental rate and yolk conversion efficiency in offspring of trout with alternative life histories

Partial migration, in which some individuals of a population migrate while other individuals remain resident, is generally associated with ontogenetic shifts to better feeding areas or as a response to environmental instability, but its underlying mechanisms remain relatively unknown. Brown trout (Salmo trutta) exhibit partial migration, with some individuals remaining in freshwater (freshwater resident) while others undertake an anadromous migration, where they spend time at sea before returning to breed in freshwater (migrant). We reared full‐sibling groups of offspring from freshwater‐resident and anadromous brown trout from the same catchment in the laboratory under common garden conditions to examine potential differences in their early development. Freshwater‐resident parents produced eggs that were slower to hatch than those of anadromous parents, but freshwater‐resident offspring were quicker to absorb their yolk and reach the stage of exogenous feeding. Their offspring also had a higher conversion efficiency from the egg stage to the start of exogenous feeding (so were larger by the start of the fry stage) than did offspring from anadromous parents despite no difference in standard metabolic rate, maximal metabolic rate or aerobic scope. Given these differences in early development, we discuss how the migration history of the parents might influence the migration probability of the offspring.     

A common environment experiment reveals plastic and genetic contributions to the fast life‐history strategy of an invasive fish

Invasive species often exhibit a suite of life‐history traits that promote rapid population growth, including early age and small size at maturation, and high reproductive investment. The common expression of these “fast” life‐history traits in invasive populations could be the result of plastic and/or genetic responses to the non‐native environment, or in response to the process of range expansion. To determine the relative importance of plastic and genetic contributions to the expression of life‐history traits, we reared two native Canadian and two invasive Spanish populations of Pumpkinseed sunfish (Lepomis gibbosus) in a common environment in central Ontario, Canada. In the wild, European Pumpkinseed tend to exhibit faster juvenile growth rates, younger age and smaller size at maturity, and higher reproductive investment than native North American populations. When reared in a common environment, both native and invasive populations exhibited similar juvenile growth rates, and similar age and size at maturity, suggesting that the differences seen among wild populations are a plastic response to the warmer non‐native environment. However, reproductive investment was consistently higher in the Spanish populations regardless of rearing environment, suggesting a genetic difference in reproductive investment between native and invasive populations. Selection for greater reproductive investment in non‐native Pumpkinseed may have contributed to their widespread success in Europe.     

Marine migrations in anadromous brown trout (Salmo trutta). Fjord residency as a possible alternative in the continuum of migration to the open sea

Partial migration is a common phenomenon in many fish species. Trout (Salmo trutta) is a partially migratory species where some part of the population migrate to the marine environment, while another remains in freshwater. In the years 2008 and 2009, a total of 159 wild sea trout smolts were tagged with acoustic and PIT‐tags in the river Villestrup (Denmark) to study the initial postsmolt marine behaviour within a fjord system. We found that the strategies of the sea migrants vary: some stay in the fjord, while others migrate to the sea, suggesting that partial migration occurs even in the marine environments. Overall, a total of 53% of the tagged smolts migrated from the fjord to the sea, and 47% stayed (or potentially died) in the fjord. The ratios of fjord‐resident versus sea‐migrating postsmolts were consistent at the study times, and no differences between the early and late migration periods of the smolts were observed. The individual's size or body condition at the time of tagging did not affect survival or the migratory decisions in the fjord. High overall initial survival (74%) was found 30 days after the fjord entry. We suggest that within a continuum of migration to sea, there is a migratory decision point when sea trout postsmolts encounter a fjord system. At this point, postsmolts will assess the possibility of migration versus the alternative of fjord residency.     

Parr dispersal between streams via a marine environment: A novel mechanism behind straying for anadromous brown trout?

Otolith Sr:Ca profiles demonstrated that juvenile anadromous brown trout (ABT) Salmo trutta may descend to the brackish waters of the Baltic Sea from their natal streams as parr or fry and then migrate into non‐natal streams before transforming into smolt. To our knowledge, no such published documentation exists for ABT. The ecological significance of this life‐history strategy is presently not clear. However, stream shifting through the marine environment should to be considered with regard to stock management and the assessment and restoration of salmonid populations and their potential habitats.     

Suitability of different salmonid strains as hosts for the endangered freshwater pearl mussel (Margaritifera margaritifera L.)

• 1. The complex life cycle of endangered European freshwater pearl mussel Margaritifera margaritifera L. involves an obligatory parasitic phase on a host fish. Knowledge on the host–parasite interaction and on the suitability of different host fish species and strains is required both for the management of wild fish and mussel populations as well as for improving the efficiency of captive breeding methods.
• 2. In this study, the suitability of different salmonid strains for hosting glochidia was tested, including Danube salmon (Hucho hucho L.) and three brown trout (Salmo trutta L.) strains from inside and outside the freshwater pearl mussel distribution range. All brown trout strains as well as Danube salmon were successfully infected with freshwater pearl mussel glochidia and encystment of mussel larvae was detected.
• 3. One brown trout strain originating from the natural pearl mussel distribution range was identified as the most suitable host, revealing the highest fish‐weight‐normalized infection rates and highest glochidial growth rates, whereas endemic Danube salmon was least suitable. Under natural conditions, the role of Danube salmon may be attributed to the long‐distance dispersal of glochidia in the Danube system, whereas sedentary brown trout appear to be the most important hosts at a local scale.
• 4. Successful infection of suitable hosts and the maintenance of these host–parasite systems in calcareous water were demonstrated in this study. These results indicate that neither the infection process nor the encystment phase of freshwater pearl mussels is dependent on low lime concentrations.
• 5. The results of this study suggest that careful selection and management of appropriate host fish strains is mandatory for sustainable conservation management of freshwater pearl mussel populations. Copyright © 2010 John Wiley & Sons, Ltd.

     

The effects of stream substratum composition on the emergence of salmonid fry

Sternecker K, Geist J. The effects of stream substratum composition on the emergence of salmonid fry.
Ecology of Freshwater Fish 2010: 19: 537–544. © 2010 John Wiley & Sons A/S Abstract – Salmonid fishes are target species for the conservation of freshwater habitats, but their natural reproduction is often insufficient. The emergence of fry is a crucial phase in the life cycle of salmonids and the stream substratum is the key habitat which regulates the emergence success. In this study, brown trout (Salmo trutta) and Danube salmon (Hucho hucho) eggs were exposed to different sediment textures and the emergence and the postemergence survival and growth were observed under constant water chemical conditions in the laboratory. In both species, textural effects on emergence rate, chronology of emergence, survival rate after emergence and growth after emergence were detected. Fine‐textured substratum (5–8 mm) formed a physical barrier to the posthatch migration of salmonids from the interstitial zone to the open water. The time period between the first and the last emerged fish was shorter in treatments with fine texture compared with coarse substratum. The survival rate was higher in treatments of coarser sediment. The effects of different textures on the growth of fry after emergence differed between brown trout and Danube salmon, which can be explained by different life history strategies. These results suggest that physical characteristics of substratum texture can have strong effects on salmonid emergence, and ultimately on the persistence of salmonid populations. They also suggest that biodiversity conservation in stream ecosystems can greatly benefit from an inclusion of the physical characteristics of the stream bed into catchment‐based management plans.     

A survey of salmon gill poxvirus (SGPV) in wild salmonids in Norway

In 2016, the Norwegian health monitoring programme for wild salmonids conducted a real‐time PCR‐based screening for salmon gill poxvirus (SGPV) in anadromous Arctic char (Salvelinus alpinus L.), anadromous and non‐anadromous Atlantic salmon (Salmo salar L.) and trout (Salmo trutta L.). SGPV was widely distributed in wild Atlantic salmon returning from marine migration. In addition, characteristic gill lesions, including apoptosis, were detected in this species. A low amount of SGPV DNA, as indicated by high Ct‐values, was detected in anadromous trout, but only in fish cohabiting with SGPV‐positive salmon. SGPV was not detected in trout and salmon from non‐anadromous water courses, and thus seems to be primarily linked to the marine environment. This could indicate that trout are not a natural host for the virus. SGPV was not detected in Arctic char but, due to a low sample size, these results are inconclusive. The use of freshwater from anadromous water sources may constitute a risk of introducing SGPV to aquaculture facilities. Moreover, SGPV‐infected Atlantic salmon farms will hold considerable potential for virus propagation and spillback to wild populations. This interaction should therefore be further investigated.     

Red operculum spots,body size,maturation and evidence for a satellite male phenotype in non‐native European populations of pumpkinseed Lepomis gibbosus

Carotenoid‐based pigmentation is a striking feature of many taxa, yet the function, if any, of colour traits is often unclear. Pumpkinseed Lepomis gibbosus, a widely introduced freshwater sunfish that exhibits alternative male mating strategies, express a striking, red operculum spot. To investigate the potential function of this red spot as a signal in this species' mating system, we determined the presence and measured the size of red operculum spots in fish collected from 12 populations in five European countries in which pumpkinseed is an established non‐native species. We subsequently related the presence and size of the red spot to body size and mating strategy, based on an analysis of relative gonad size (gonado‐somatic index, GSI), using a mixed modelling approach. The study demonstrated that the presence of a red operculum spot in pumpkinseed is associated with sexual maturation, with GSI frequency distributions suggesting that cuckolders in some non‐native populations comprised both sneaker and satellite males, the latter not having previously been reported for this species. Further, the size of red spot correlated strongly with body size in parental and cuckolder males, although there was no difference in the presence or size of the red operculum spot between male mating strategies. The function of a red operculum spot in females is not clear but may be partly mediated by pleiotropic genetic mechanisms. Red operculum spots appear to function as signals of male maturation and body size in pumpkinseed, irrespective of mating strategy.     

Intrinsic and extrinsic drivers of life‐history variability for a south‐western cutthroat trout

The impacts of climate change on cold‐water fishes will likely negatively manifest in populations at the trailing edge of their distributions. Rio Grande cutthroat trout (Oncorhynchus clarkii virginalis, RGCT) occupy arid south‐western U.S. streams at the southern‐most edge of all cutthroat trout distributions, making RGCT particularly vulnerable to the anticipated warming and drying in this region. We hypothesised that RGCT possess a portfolio of life‐history traits that aid in their persistence within streams of varying temperature and stream drying conditions. We used otolith and multistate capture–mark–recapture data to determine how these environmental constraints influence life‐history trait expression (length‐ and age‐at‐maturity) and demography in RGCT populations from northern New Mexico, United States. We found evidence that RGCT reached maturity fastest at sites with warm stream temperatures and low densities. We did not find a strong relationship between discharge and any demographic rate, although apparent survival of mature RGCT decreased as stream temperature increased. Our study suggests plasticity in trait expression may be a life‐history characteristic which can assist trailing edge populations like RGCT persist in a changing climate.     

Anadromy and the dispersal of an invasive fish species (Oncorhynchus mykiss) in Eastern Quebec,as revealed by otolith microchemistry

Thibault I, Hedger RD, Dodson JJ, Shiao J‐C, Iizuka Y, Tzeng W‐N. Anadromy and the dispersal of an invasive fish species (Oncorhynchus mykiss) in Eastern Quebec, as revealed by otolith microchemistry. Ecology of Freshwater Fish 2010: 19: 348–360. © 2010 John Wiley & Sons A/S Abstract – Rainbow trout (Oncorhynchus mykiss) is invading rivers bordering the St. Lawrence Estuary (Quebec, Canada). Some rivers in Eastern Quebec support self‐sustaining populations while adult vagrants are frequently captured in rivers where no reproduction has been confirmed. We hypothesised that the development of anadromy has promoted the species dispersal. Otolith Sr:Ca analyses revealed that although all fish captured in the upstream stocking region were freshwater residents, both anadromous and freshwater resident phenotypes were found downstream in Eastern Quebec. The proportion of fish exhibiting the anadromous life cycle increased with the distance from the stocking zone. Eastern Quebec steelhead migrated to sea at the same age but at a larger size than steelhead within their native range. Age at first reproduction was similar to that observed in native populations. The development of the anadromous life cycle enables this species to colonise new rivers following long‐distance migrations along the St. Lawrence Estuary corridor.     

Competitive release modifies the impacts of hydrologic alteration for a partially migratory stream predator

Understanding ecological effects of altered stream flows is an essential objective. In a comparative field study of juvenile steelhead trout (Oncorhynchus mykiss) that compared natural‐ and reduced‐flow sites, we used mark–recapture modelling and information theory to quantify spatially and temporally explicit patterns of density, specific growth rate, survival and outmigration; and test predictions for biotic and abiotic drivers. Densities were lower in water withdrawal treatments, resulting in lower intraspecific competition and, higher specific growth rate and survival. We observed yearly differences in density and intraspecific competition, with a negative relationship between density and specific growth rate over a wide range of densities, but reductions in survival only at the highest densities. Moreover, individual variability within sites was important. At high density (sites and years), survival related negatively to body size. In contrast, when overall density was lower, specific growth rate was negatively related to body size. Lastly, individuals were more likely to outmigrate when they had larger body size, lower survival or reared in habitats with reduced flows, and these patterns appeared mediated by the intensity of intraspecific competition. Our results underscore the harsh bioenergetic conditions induced by higher temperatures and densities during summer baseflow (relative to other seasons), particularly for larger fish demanding more resources, and suggest a density‐dependent mechanism for why this period is important for regulating salmonid populations. We found that a complex combination of natural (e.g., density) and anthropogenic (e.g., withdrawal) factors affected juvenile salmon populations and life history expression in the face of altered flows.     

Association of brook trout and Oncorhynchus spp. with large wood jams in a Lake Superior tributary in a northern old‐growth watershed

Wood in streams functions as fish habitat, but relationships between fish abundance (or size) and large wood in streams are not consistent. One possible reason for variable relationships between fish and wood in streams is that the association of fish with wood habitat may depend on ecological context such as large‐scale geomorphology. We studied the relationship between salmonid assemblages and large wood jams (LWJ) in four settings that differed geomorphically at the scale of the stream corridor along a tributary to Lake Superior in old‐growth conifer–hardwood forest in northern Michigan. The focal fish species of this study were brook trout (Salvelinus fontinalis), which were wild in the stream. Relocation efforts for coaster brook trout (an adfluvial life history variant of brook trout) were ongoing in the study stream. We measured fish abundance and length in pairs of pools of similar size and substrate, but varying in the presence of LWJ; this allowed us to evaluate associations of fish simply with the presence of LWJ rather than with other channel or flow‐shaping functions of LWJ. The length of Oncorhynchus spp. and young introduced brook trout was not strongly correlated with LWJ presence; however, the presence of LWJ in pools was positively correlated with larger wild brook trout. We also found that the correspondence of LWJ with the abundance of salmonids appears to be moderated by the presence of alternative habitat in this relatively natural, old‐growth forest stream.     

Detection of density‐dependent growth at two spatial scales in marble trout (Salmo marmoratus) populations

Vincenzi S, Crivelli AJ, Jesensek D, De Leo GA. Detection of density‐dependent growth at two spatial scales in marble trout (Salmo marmoratus) populations.
Ecology of Freshwater Fish 2010: 19: 338–347. © 2010 John Wiley & Sons A/S Abstract – Density‐dependent body growth has often been observed in freshwater salmonid populations. Several studies suggest this compensatory pattern as a potential mechanism of population regulation. The choice of the spatial scale is important for the detection of density‐dependent growth, as study areas need to be of the appropriate size to capture the density of conspecifics actually experienced by individuals over the preceding growth period. Here, we used four marble trout (Salmo marmoratus) populations (Gatsnik, Gorska, Huda and Zakojska) living in Slovenian stream to study the relationships between early density of marble trout and subsequent body growth. As streams are divided in sectors delimited by natural barriers that prevent or strongly limit movement of individuals, we tested the relationship between early density and body size through the lifetime at two spatial scales, that is, sector level (for Gatsnik and Zakojska) and whole stream level (the four populations were pooled). Sector length in Gatsnik and Zakojska ranged from 113 to 516 m. At both sector and whole stream level, temporal data were pooled. Growth declined significantly with increasing density both at the sector and whole stream levels, and the density‐dependent relationship was described by negative power curves. However, at the sector level the density‐dependent pattern was stronger in Gatsnik, a stream in which fish could move across sectors, than in Zakojska, where upstream movement across sectors is prevented by waterfalls.     

Factors influencing growth of individual brown trout in three streams of the upper Midwestern United States

Growth rate variation of three age groups of brown trout, Salmo trutta L. (age‐0, 1 and 2, and 3+), was quantified from recaptured, individually tagged brown trout and related to season, stream reach, relative abundance, initial length and movement to examine factors influencing growth in length in three streams in the Midwestern United States. Total variation in growth was almost five times greater for age‐0 than for age‐3+ trout. Individual trout accounted for about 13% of total variation in age‐0 growth, season about 57%, and trout initial length and relative abundance combined another 2%. The 2006 age‐0 cohort had the fastest growth rates in their second spring and summer (2007) and slowest growth in their first winter (2006–2007). About 53% of total growth variation of age‐1 and age‐2 trout was accounted for by individual trout, season, initial length and stream reach. Predicted growth rates indicated strong effects of season and initial length. A significant interaction between these two factors indicated that, although smaller trout grew faster than larger trout, this length effect was most pronounced in spring and summer. About 35% of total growth variation of age‐3+ trout was accounted for by individual trout and season. Together, season and individual trout characteristics were identified as the most important factors influencing brown trout growth in these streams.     

Reliance on lakes by salmon,trout and charr (Oncorhynchus,Salmo and Salvelinus): An evaluation of spawning habitats,rearing strategies and trophic polymorphisms

Salmonid fishes may reside within or migrate between stream and lake habitats, or undergo anadromous migrations between freshwater and the ocean. While the degree of anadromy of salmonids has been thoroughly compared, no analogous review has examined the degree of lake use. To assess the extent of reliance on lake habitat in this family, we considered 16 species of salmon, trout and charr from the genera Oncorhynchus, Salmo and Salvelinus, comparing their (a) use of lakes as spawning habitat, (b) rearing strategies in lakes, and (c) occurrence and diversity of lacustrine trophic polymorphism. In identifying the primary life‐history patterns of each species and exploring the lesser‐known lacustrine behaviours, we found that the extent of reliance on lakes exhibits a negative association with the degree of anadromy. Oncorhynchus rely least on lakes, Salmo to an intermediate level and Salvelinus the most, opposite of the general prevalence of anadromy among these genera. Lakes are critical to adfluvial and lake‐resident salmonids, but they also support anadromous and fluvial life histories by providing spawning, rearing, overwintering and/or summer refuge habitat. Adfluviality, although a non‐anadromous life history, consists of similar migration‐related traits and behaviours as anadromy, including the parr–smolt transformation, sex‐biased patterns of migration and residency, and the presence of precocious males. Lakes support life‐history variants, reproductive ecotypes and trophic morphs unique to lacustrine habitat. Therefore, conservation of salmonids is dependent on maintaining the diversity and quality of their habitats, including lakes.