Instream and catchment characteristics affecting the occurrence and population density of brown trout, Salmo trutta L., in forest brooks of a boreal river basin

The occurrence and density of ≥ 1+ brown trout, Salmo trutta L., and their relationship with prevailing instream and catchment characteristics were studied in 50 small forest streams, partially dredged for forest ditching. The occurrence of trout at a stream site was largely determined by the abundance of pools, size of upper catchment and water pH. Moreover, at sites where trout occurred, the abundance of pools was lower at dredged locations than at those in a natural state. In riffles in a natural state, there was a positive relationship between trout density and three instream variables: the abundance of stream pools, cascades and instream vegetation, while an inverse relationship was found with the abundance of substratum of 2–10 cm in diameter. Of the catchment variables, correspondingly, the proportion of forest in the upper catchment was positively related and the proportion of peatland negatively related to trout density. No significant regression model could be fitted for dredged riffles. The possibility of enhancing trout populations in dredged riffles is discussed.     

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

Abstract– 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.     

The age structure and distribution of Atlantic salmon parr, Salmo salar L., in small tributaries and main stems of the subarctic River Teno, northern Finland

Parr of the Atlantic salmon in subarctic River Teno, northern Finland (70°N, 28°E), are found to migrate to small tributaries that are not spawning areas for the adult salmon. The age distribution of the salmon differs significantly between these brooks and the natal rivers, the parr in the brooks being typically 2–4 years old, whereas those in the main rivers were mostly of age 0–2 years. The older fish were found in the uppermost regions of the brooks. The ratio of salmon to brown trout decreases towards the upper reaches of the brooks. At least some of the young salmon overwinter in the brooks. However, the number of fish and the area inhabited diminished towards the autumn in some occasions. It is suggested that this migratory behavior may be an adaptational phenomenon that could cause variation in survival among young salmon in the River Teno.     

Density-dependence of growth characteristics and maturation in stream-dwelling resident brown trout, Salmo trutta, in Serbia

Abstract  Breakpoints in both length and weight that represent the onset of first maturation, revealed that growth and maturation were correlated with density-related parameters in stream-dwelling resident brown trout, Salmo trutta L., from nine different rivers in Serbia. The determination of density dependence of growth is not simple, as both river width and depth, as single variables of habitat and relative biomass, and density were not significantly correlated with breakpoints. The significant relationship between breakpoints and the mean weight of brown trout ( x _p) suggests that differences in breakpoints were inversely related to brown trout density. The implications of density dependence for management of streams holding resident brown trout are discussed.     

Density of brown trout, Salmo trutta L., and Atlantic salmon, Salmo salar L., parr after point and scatter stocking of fry

Atlantic salmon, Salmo salar L., and brown trout, Salmo trutta L., fry were point and scatter stocked in the early part of June at densities of 63–263 fry 100 m⁻² per species in the River Viantienjoki, a small river in northern Finland, and their population densities were assessed in late summer. Both species were always stocked together in similar quantities. Point stocking was used in the first 2 years and scatter stocking in the following 2 years. In point stocking, there was no correlation between the distance from the stocking sites (maximum = 250 m) and parr density in census sites ( r = −0.013 and 0.019 for brown trout and Atlantic salmon, respectively). The stocking density of fry did not influence parr density in August by either method or by species. Stocking density explained only from 11% to 23% of the parr survival depending on the species or stocking method. The mean densities of Atlantic salmon and brown trout parr did not differ significantly from each other at any fishing site ( P > 0.05). Both point and scatter stocking appear to be suitable methods for use in small rivers. The parr densities depend more on the other factors (e.g. habitat quality) than the stocking method, and the choice between methods could be based on the time and labour available.     

Do instream habitat variables and the abundance of brown trout Salmo trutta (L.) affect the distribution and growth of stone loach,Barbatula barbatula (L.)?

Abstract –  Along a stream, we investigated whether the abundance of stone loach ( Barbatula barbatula , L.) was related to the presence of brown trout ( Salmo trutta , L.) and instream habitat variables. First, a field survey was carried out where different habitat variables and the densities of both species were quantified and subjected to principal components analysis. Then the abundance of stone loach was related to the scores of the retained axes (eigenvalues >1). The abundance of stone loach was positively correlated to substrate particle size, amount of shade, temperature, discharge and current velocity, but negatively correlated to brown trout abundance. Secondly, a month-long field enclosure experiment in a stream was performed to test for any negative effects of brown trout on stone loach growth. Four treatments were used: intraspecific competition (stone loach at double density), interspecific competition (stone loach + small trout), predation (stone loach + large trout) and a control (stone loach alone). The results showed that large trout tended to have negative effects on final stone loach biomass. The absence of a negative effect of large trout on resource density suggests that nonlethal effects rather than resource competition caused this trend.     

Relationships between trout habitat use and woody debris in two southern New England streams

Abstract – In-stream habitat was measured and trout density was estimated in Merrick Brook (105 habitat units) and the Tankerhoosen River (135 habitat units), Connecticut to determine relationships between habitat use of brook trout Salvelinus fontinalis and brown trout Salmo trutta and woody debris. In each habitat unit, woody debris was inventoried, and length, width, depth, area, width : depth ratio and undercut bank area were estimated. Trout abundance was estimated by snorkeling. Multiple regression was used to test relationships between trout density and principal components describing habitat unit variables. In Merrick Brook, habitat unit size and shape explained most of the variability in density of brook trout (<130 and ≥130 mm) and brown trout (<150 mm) among habitat units, although principle components describing large woody debris or fine woody debris contributed significantly to variations in density of brook trout (≥130 mm) and brown trout (<150 and ≥150 mm). In the Tankerhoosen River, fine woody debris explained most of the variability in density of brook trout (<130 and ≥130 mm), followed by habitat unit size and shape. Both large woody debris and fine woody debris contributed significantly to variations in density of brown trout (≥150 mm). These results suggest that woody debris is an important component of wild trout habitat above that provided by habitat unit shape and size alone.     

Lipid energy reserves influence life-history decision of Atlantic salmon (Salmo salar) and brown trout (S. trutta) in fresh water

Abstract –  Lipid density appears to influence life-history decisions in salmonid fishes. This study shows that parr and smolts of anadromous Atlantic salmon from a south Norwegian river have on average between 30 and 40% higher energy level than corresponding brown trout in spring and summer, which may explain differences in life-history traits between the two species. The higher energy density of young salmon was chiefly due to a 1.8 times higher lipid density in parr and 2.4 times higher lipid density in smolts. The difference was smaller among immature parr in the autumn, with only 1.4 times higher lipid density in salmon than trout. The reason for the decreased difference was probably that the more energy rich salmon parr had attained maturity at the time. Among mature male parr, the somatic energy density was approximately 10% higher in trout than salmon. However, the gonadal energy content was more than twice as high in salmon than in trout. The higher somatic energy allocation in parr of Atlantic salmon probably influences protein growth of the two species in fresh water, and increases the ability of salmon relative to trout to undertake long distance feeding migrations and make large investments in reproduction.     

Selective segregation in intraspecific competition between juvenile Atlantic salmon (Salmo salar) and brown trout (Salmo trutta)

Interactive segregation has been suggested as the ruling competition mechanism determining niche and niche segregation between juvenile Atlantic salmon (Salmo salar) and brown trout (Salmo trutta). Results from allopatry–sympatry observations of habitat use in both nature and in experiments were contrary to predictions derived from the interactive segregation hypothesis. Habitat use parameters under natural conditions such as distance to shore for Atlantic salmon parr were nearly identical in allopatric (mean ± SD; 3.2 ± 1.4 m) and sympatric (3.3 ± 1.4 m) situations. Occupied water depths largely reflected available water, but water depths <15 cm were avoided by salmon parr. Under experimental conditions, habitat use of allopatric salmon was density independent and salmon size had only minor effects, with smaller fish being more likely to occur in the shallow. Habitat use of salmon in sympatry with trout did not differ from allopatric salmon habitat use, and only salmon size had minor effects on depth choice – occurrence of trout or fish density had no effect. Allopatric trout was in general more frequent in the shallow habitat than salmon. Habitat use of sympatric trout was affected by the occurrence of salmon and trout size, resulting in a higher use of the shallow habitats for small trout. To conclude, selective segregation has a dominant role in salmon habitat use (not affected by trout occurrence), whereas a mixed situation occurs in trout habitat use with elements of interactive segregation when competing with Atlantic salmon (affected by salmon occurrence).     

Use of first‐order tributaries by brown trout (Salmo trutta) as nursery habitat in a cold water stream network

Many investigators have examined the importance of suitable in‐stream habitat and flow regime to salmonid fishes. However, there is much less known about the use of small (<5 l·s^?1 discharge) first‐order streams within a larger stream network by salmonids. The purpose of this study was to evaluate the use of small headwater streams by juvenile brown trout Salmo trutta in the Emmons Creek stream network in Wisconsin, USA, and to determine whether abundance was related to habitat variables in these streams. Fishes in eight spring‐fed first‐order streams were sampled during a 7‐month period using a backpack electroshocker and measured for total length. Habitat variables assessed included stream discharge, water velocity, sediment composition and the abundance of cover items (woody debris and macrophytes). Densities of YOY trout ranged from 0 to 1 per m² over the course of the study and differed among first‐order streams. Stepwise multiple regression revealed discharge to be negatively associated with trout density in spring but not in summer. All other habitat variables were not significantly related to trout density. Our results demonstrate the viability of small first‐order streams as nursery habitat for brown trout and support the inclusion of headwater streams in conservation and stream restoration efforts.     

Habitat selection by rainbow trout Oncorhynchus mykiss in a California stream: implications for the Instream Flow Incremental Methodology

We quantified microhabitat selection of rainbow trout (Oncorhynchus mykiss) at 2 flows (low= 1.13 m³. s^?1 and high =4.95 m³. s^?1) in the Pit River, California. Flows were controlled by an upstream dam and habitat availability was similar during 4 sampling periods at low flow and 2 periods at high flow. A principal components analysis reduced 6 microhabitat variables to 3 new variables that explained 80% of the observed variance. The 3 components loaded heavily on velocity variables, depth variables and substrate. Microhabitat selection generally differed among macrohabitats (i. e., pools, runs, and riffles). Rainbow trout selected different microhabitats at high flow relative to low flow in response to the availability of deeper, faster water. At low flow, depth and velocity selection were positively correlated with seasonal temperature change for adults but not juveniles. Rainbow trout apparently sought shelter in interstitial spaces in the substrate of runs and riffles during the day in early winter. Generally, large rainbow trout were observed in pools, intermediate-sized fish in runs, and small trout in riffles. The largest fish occurred in slow, deep areas of pools, where they moved slowly without orientation to flow and were not observed feeding, whereas small fish generally faced upstream and fed in all habitat types. Foraging forays were directed up in the water column at velocities similar to the mean water column velocities at holding positions. Rainbow trout were the most abundant species in 76% of the population survey stations. Other species that might have influenced microhabitat selection by rainbow trout were uncommon.     

Some behavioural and ecological factors affecting distribution,biomass and production of juvenile Atlantic salmon

Observations were made in an experimental stream tank (total area 14.7 m²) on juvenile Atlantic salmon, Salmo salar L (parr), relating experimental observations to field observations, including the reported diurnal fasting behaviour of juvenile salmon at water temperatures <10 °C. Densities in the tank ranged from five to twenty parr, at water temperatures ranging from 4.6 °C to 15.8 °C. The wide channel of the stream tank, with mean water velocity of 18.8 cm·s^?1, was the preferred section, where territorial behaviour was observed. Biomass was regulated in the wide channel by territorial mosaics or by dominance hierarchies. Dominance hierarchies were reflected in coloration of the fish. Dominant salmon were generally in the wide channel. Densities of salmon parr (of mean fork length 10.2 cm) in the channel ranged from 0.84 m^?2 to 1.73 m^?2, with an average biomass of 14.2 g·m^?2. Growth was least at the 5.9 °C temperature treatment. In experiments at temperatures below 10 °C, feeding, dominance hierarchy and territorial behaviour were observed in daylight hours, contrary to the published literature. Interactions with other species may affect behaviour. Some observations were made on a closely related species, brown trout (Salmo trutta L.), a commonly cohabiting species in many systems. Trout displaced salmon from their preferred locations in the tank and were more aggressive than the salmon, reducing agonistic behaviour by the salmon. The commonest agonistic act shown by salmon was ‘charge’ and that by the trout was ‘approach’. Some field observations affecting behaviour and production are discussed.     

The influence of water depths and inter-specific interactions on cover responses of juvenile Atlantic salmon

Abstract  – The responses of salmon parr, Salmo salar, to instream cover, related to several water depths, were tested in an ellipsoidal stream tank. Opaque plastic covers, most of which were 20 cm in length and 16 cm high, were randomly distributed through the tank, occupying about 3% of the bottom area. Six salmon parr were used for each experiment. In experiments with channel depths of 40 cm, the proportion of salmon under covers was 36.4%, at 30 cm 60.7%, and at 20 cm, 79.4%. Presence of similar-sized brown trout, Salmo trutta, increased the use of covers in channels and greater use of the pool area by salmon. The salmon were completely dominated by the trout, making about four times more more aggressive than salmon. Our findings show that water depth itself provides cover for young salmon.     

Effects of predation and intraspecific interactions on habitat use and foraging by brown trout in artificial streams

Abstract— We studied habitat use, foraging rates and behavior of 10 cm and 12 cm long brown trout, Salmo trutta , at two densities, 1.5 and 3.0 fish. m⁻², in artificial streams that contained either the amphipod, Gammarus pulex , alone or G. pulex together with the piscivore, northern pike, Esox lucius. Gammarus were stocked in and largely restricted to the pools at a density of 128 Gammurus. m⁻² . pool⁻¹ Large trout (12 cm) used pools more and riffles less when small trout (10 cm) were present than when small trout were absent. Small trout consumed fewer Gammarus when together with large trout than when alone, but showed no difference in habitat use in the presence and abscnce of large trout. Habitat use and number of Gammarus consumed per trout were not affected by trout density for either size-class when alone. For both size-classes of trout, use of pools and foraging rates were higher in the absence than in the presence of pike, and pike primarily resided in the pools. The number of aggressive interactions by both size-classes of trout decreased when pike was present. Our results indicate that for habitats that differ in food resources and predation risk, size structure may affect habitat use and foraging by brown trout.     

Influence of Physical Factors on Density of Stocked Brown Trout (Salmo trutta fario L.) in a Danish Lowland Stream

Physical factors and brown trout densities were studied in a small Danish lowland stream. The densities of brown trout larger than 15 cm were significantly correlated with gradient, mean depth, coefficient of variation in current velocity 7 cm above the bottom, the ratio between wetted perimeter and width, amount of overhanging banks and degree of macrophyte cover. Coefficient of variation in current velocity 7 cm above the bottom was the most important factor for brown trout density (r_s= 0.8364, 24 df, (P < 0.001)), which supports the idea of this value as a measure of stream complexity. A rather small relation between trout density and amount of overhanging bank cover (r_s= 0.4179, 24 df, (P < 0.050)), contrary to the closer relationships found in previous studies, is discussed as an effect of the self-shading capacity of this rather narrow and deep stream.     

Water level influences migratory patterns of anadromous brown trout in small streams

Habitats modify the population ecology of species. Here, we show how low water level influences abundance and size of adult anadromous brown trout (Salmo trutta) entering a small, South Norwegian stream for spawning. After smolting, the fish appear chiefly to feed within 10 km of the home stream. In the autumn, South Norwegian streams typically flood because of heavy rainfall, when the anadromous brown trout entered from the sea. Mean annual duration of the upstream migration period was 34 days and ended when the flooding ended and the water temperature dropped to below 4°C. During most of the migration period, on average two trout ascended the river per day. The sexes entered the spawning area concurrently, and the male:female ratio of the anadromous trout was 1.27. No fish entered when the water depth just downstream of the spawning area was below 5 cm, and mean number of fish increased with increasing water depth to ca. 30 cm, but not at higher flows when the ascent gradually decreased. Mean and maximum size of the entering spawners increased with water depth between 5 and 16 cm. Among those that had been to sea, most were recaptured in the home stream, 4% in other streams, but only two of the strays were caught close to spawning time. The present results illustrate that population traits of anadromous brown trout from a small stream differ from those in larger rivers, probably because of selection associated with water flow.     

Predation on wild and hatchery salmon by non‐native brown trout (Salmo trutta) in the Trinity River,California

Non‐native predators may interfere with conservation efforts for native species. For example, fisheries managers have recently become concerned that non‐native brown trout may impede efforts to restore native salmon and trout in California's Trinity River. However, the extent of brown trout predation on these species is unknown. We quantified brown trout predation on wild and hatchery‐produced salmon and trout in the Trinity River in 2015. We first estimated the total biomass of prey consumed annually by brown trout using a bioenergetics model and measurements of brown trout growth and abundance over a 64‐km study reach. Then, we used stable isotope analysis and gastric lavage to allocate total consumption to specific prey taxa. Although hatchery‐produced fish are primarily released in the spring, hatchery fish accounted for most of the annual consumption by large, piscivorous brown trout (>40 cm long). In all, the 1579 (95% CI 1,279–1,878) brown trout >20 cm long in the study reach ate 5,930 kg (95% CI 3,800–8,805 kg) of hatchery fish in 2015. Brown trout predation on hatchery fish was ca. 7% of the total biomass released from the hatchery. Brown trout only ate 924 kg (95% CI 60–3,526 kg) of wild fish in 2015, but this was potentially a large proportion of wild salmon production because wild fish were relatively small. As large brown trout rely heavily on hatchery‐produced fish, modifying hatchery practices to minimise predation may enhance survival of hatchery fish and potentially reduce the abundance of predatory brown trout.     

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.     

Determinants of parr size variations within a population of brown trout Salmo trutta L.

Abstract – The size of 2-month old trout Salmo trutta parr differed between sites and between years along the River Esva catchment (Asturias, northwestern Spain). Such variation was in a direction opposite to the variation observed in parental size. Parr were smaller in sites where parents grew faster, whereas larger parr occurred in sites where parents grew less. Parr size of six cohorts (1990–1996, except 1995) at 14 sites along the River Esva was inversely related to the growth rate and length of parents and positively related to canopy, egg size, and water temperature. The latter acted similarly on all parr independently of egg size and the site where the egg originated. Covariation patterns among parr size, parental traits, and canopy suggest that a canopy-regulated, growth-determined trade-off between egg size and number, previously described for the Esva trout, also extends to alevin size. In forested, shaded sites, adult trout grew less and spawned fewer larger eggs that resulted in larger alevins, whereas in fully insolated, production-rich sites, trout grew faster and spawned higher numbers of smaller eggs that produced smaller parr. I hypothesize that the phenotypic plasticity illustrated by the environmentally induced trade-off between egg size and number further extended to alevin size may be evolutionarily advantageous because it relates the size of trout alevins to food availability, as predicted by the growth previously experienced by parents. ^NOTE     

Spatial and temporal variation in brook trout and brown trout proportional stock densities in Wisconsin streams

Abstract  Proper interpretation of measures used to describe fish populations requires knowledge of the measure's inherent spatial and temporal variation. Proportional stock density (PSD), the ratio of 'quality-length' fish to 'stock-length' fish multiplied by 100, is commonly used as a measure of population size structure; PSD values range from 0 to 100. Spatial and temporal variation in brook trout, Salvelinus fontinalis (Mitchill), and brown trout Salmo trutta L., PSD scores in Wisconsin are described and tested to determine if variation differed by stream order and ecoregion. Neither stream order nor ecoregion significantly affected variation of PSD scores. The mean standard deviation of PSD scores over time at a site was 12.49 for brook trout populations and 12.95 for brown trout populations. The mean standard deviation of PSD scores between sites in the same stream was 15.07 for brook trout populations and 12.50 for brown trout populations. Sampling frequency required to characterise a PSD score of a single population of trout in Wisconsin streams with a degree of precision equal to the amount of observed temporal variation is approximately 14 sites for brook trout and 20 sites for brown trout.