Habitat shifts in rainbow trout: competitive influences of brown trout

Summary We compared habitat use by rainbow trout sympatric (three streams) and allopatric (two streams) with brown trout to determine whether competition occurred between these two species in the southern Appalachian Mountains. We measured water depth, water velocity, substrate, distance to overhead vegetation, sunlight, and surface turbulence both where we collected trout and for the streams in general. This enabled us to separate the effects of habitat availability from possible competitive effects. The results provided strong evidence for asymmetrical interspecific competition. Habitat use varied significantly between allopatric and sympatric rainbow trout in 68% of the comparisons made. Portions of some differences refelected differences in habitats available in the several streams. However, for all habitat variables measured except sunlight, rainbow trout used their preferred habitats less in sympatry with brown trout than in allopatry if brown trout also preferred the same habitats. Multivariate analysis indicated that water velocity and its correlates (substrate particle size and surface turbulence) were the most critical habitat variables in the interaction between the species, cover in the form of shade and close overhead vegetation was second most important, and water depth was least important.     

Brown trout spawning habitat selection preferences and redd characteristics in the Credit River, Ontario

Brown trout Salmo trutta redds in the Credit River, Ontario, were enumerated and assessed for physical, location and cover feature characteristics during the 2002 spawning season. Hydraulic habitats were classified on the basis of channel morphology and availability recorded. Combined use and availability data were used to assess habitat selection preferences and test for significant differences. Significant preferences for upstream pool and riffle habitat were found, but all available habitat types were used to some degree. Non‐preferred habitat redds were significantly larger than those located in preferred habitats and more likely to be found in association with woody debris cover. Larger redds were interpreted as indicative of larger fish. The use of non‐preferred downstream habitats by larger fish was further interpreted in the context of overwinter habitat requirements to reflect possible trade‐offs between reproductive and adult over‐winter survival requirements resulting in the selection of habitat for multiple life‐history functions. Results suggest that redd density alone should not be used to infer critical autumn habitat requirements for brown trout.     

Winter ecology of Arctic charr (<Emphasis Type="Italic">Salvelinus alpinus</Emphasis>) and brown trout (<Emphasis Type="Italic">Salmo trutta</Emphasis>) in a subarctic lake,Norway

We studied habitat choice, diet, food consumption and somatic growth of Arctic charr (Salvelinus alpinus) and brown trout (Salmo trutta) during the ice-covered winter period of a subarctic lake in northern Norway. Both Arctic charr and brown trout predominantly used the littoral zone during winter time. Despite very cold winter conditions (water temperature <1°C) and poor light conditions, both fish species fed continuously during the ice-covered period, although at a much lower rate than during the summer season. No somatic growth could be detected during the ice-covered winter period and the condition factor of both species significantly declined, suggesting that the winter feeding rates were similar to or below the maintenance requirements. Also, the species richness and diversity of ingested prey largely decreased from summer to winter for both fish species. The winter diet of Arctic charr <20 cm was dominated by benthic insect larvae, chironomids in particular, and Gammarus lacustris, but zooplankton was also important in December. G. lacustris was the dominant prey of charr >20 cm. The winter diet of brown trout <20 cm was dominated by insect larvae, whereas large-sized trout mainly was piscivorous, feeding on juvenile Arctic charr. Piscivorous feeding behaviour of trout was in contrast rarely seen during the summer months when their encounter with potential fish prey was rare as the small-sized charr mainly inhabited the profundal. The study demonstrated large differences in the ecology and interactions of Arctic charr and brown trout between the winter and summer seasons.     

Comparison between PIT and radio telemetry to evaluate winter habitat use and activity patterns of juvenile Atlantic salmon and brown trout

Winter habitat use and activity patterns of juvenile Atlantic salmon and brown trout were analysed in a comparative study between Passive Integrated Transponder (PIT) technology, radio telemetry and underwater observation by snorkelling. Two study periods were conducted in Stoney River, Newfoundland, Canada. During Study period I, 49 juvenile Atlantic salmon (fork length: 11.0–18.0 cm) and 7 brown trout (11.0–17.3 cm) were tagged with PIT tags and/or radio transmitters in late winter of 2004. During Study period II, 18 juvenile Atlantic salmon (fork length: 12.0–18.4 cm) and 23 brown trout (10.9–20.8 cm) were tagged and tracked twice a day at 10:00 h and 22:00 h on five consecutive days in late winter of 2005. From the 56 fish released during Study period I, on average 19.6 ± 6.0% of the PIT tagged fish and 99.3 ± 2.2% of the radio tagged fish were relocated during any given survey. Over the Study period II, 39% of fish emigrated from the study site. PIT technology had an efficiency of 39.2 ± 14.1% to detect the remaining fish. In contrast, radio telemetry relocated on average 96.9 ± 6.5% of the tagged fish whereas by snorkelling on average only 4.1 ± 5.6% of the tagged fish were observed. PIT telemetry may however be more efficient in smaller, less heterogeneous streams. The advantage of PIT technology over radio telemetry is clearly that it is relatively less costly permitting higher numbers of individuals to be tagged and there is no limit in the operational life of the transponder. In winter, juvenile salmonids preferred low flow velocity and no preferences were observed for any specific water depth over the range of available water depths. Fish selected preferentially boulder habitat over other substrates in the environment. Habitat utilisation did not differ between day and night. The use of winter preference indices may be important for future habitat modelling.     

The importance of seasonal macrophyte cover for the behaviour and performance of brown trout (Salmo trutta) in a groundwater‐fed river

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Habitat use by lemmings near Barrow, Alaska

Brown lemmings reach much higher densities than collared lemmings near Barrow, Alaska, and captures from 19 summers of snap trapping confirm previous reports that brown lemmings prefer lower, wetter habitats than do collared lemmings. Data also support the hypothesis that brown lemmings concentrate in higher habitats during early summer when melt water floods lower habitats, then shift to lower habitats where preferred food is more available as the waters recede. This pattern appears similar to seasonal shifts in habitat use reported for Norwegian lemmings. Two hypotheses were not supported by our data: (1) Unlike Norwegian lemmings, brown lemmings did not expand their use of suboptimal habitats at higher population densities. Rather, absolute densities changed in concert so that the relative densities among habitats remained unchanged. (2) Preferential use of polygon troughs during winter, as indicated by patterns of winter grazing, was not simply a function of snow depth. Instead it appeared to be related to shoot density of preferred foods. Nearly all patterns of habitat use seemed to be linked to food availability. Other factors, such as protection from predators by vegetative cover in summer and increased insulation from deeper snow in winter, did not appear to influence the distribution of lemmings as strongly.     

Intercohort competition causes spatial segregation in brown trout in artificial streams

When together in a flume, 13-cm L _s (age-1) trout appeared to increase their activity and behaved more aggressively, whereas 10-cm L _s (age-0) trout tended to move less and were rarely aggressive in the presence of the larger size class. Both size classes were less mobile and preferred lower water velocities in winter than in summer, and increased their use of instream cover in winter. When both size classes were present, only small trout decreased their use of low water velocities and cover. The results indicate that intercohort competition may cause spatial segregation among size groups of brown trout, especially in winter when trout attempt to minimize their maintenance costs.     

Static habitat partitioning and dynamic selection by sympatric young Atlantic salmon and brown trout in south-west England streams

Direct underwater observation of micro‐habitat use by 1838 young Atlantic salmon Salmo salar [mean L_T 7·9 ± 3.1(s.d.) cm, range 3·19] and 1227 brown trout Salmo trutta (L_T 10·9 ± 5·0 cm, range 3·56) showed both species were selective in habitat use, with differences between species and fish size. Atlantic salmon and brown trout selected relatively narrow ranges for the two micro‐habitat variables snout water velocity and height above bottom, but with differences between size‐classes. The smaller fishes <7 cm held positions in slower water closer to the bottom. On a larger scale, the Atlantic salmon more often used shallower stream areas, compared with brown trout. The larger parr preferred the deeper stream areas. Atlantic salmon used higher and slightly more variable mean water velocities than brown trout. Substrata used by the two species were similar. Finer substrata, although variable, were selected at the snout position, and differences were pronounced between size‐classes. On a meso‐habitat scale, brown trout were more frequently observed in slow pool‐glide habitats, while young Atlantic salmon favoured the faster high‐gradient meso‐habitats. Small juveniles <7 cm of both species were observed most frequently in riffle‐chute habitats. Atlantic salmon and brown trout segregated with respect to use of habitat, but considerable niche overlap between species indicated competitive interactions. In particular, for small fishes <7 cm of the two species, there was almost complete niche overlap for use of water depth, while they segregated with respect to water velocity. Habitat suitability indices developed for both species for mean water velocity and water depth, tended to have their optimum at lower values compared with previous studies in larger streams, with Atlantic salmon parr in the small streams occupying the same habitat as favoured by brown trout in larger streams. The data indicate both species may be flexible in their habitat selection depending on habitat availability. Species‐specific habitat overlap between streams may be complete. However, between‐species habitat partitioning remains similar.     

Assessing the suitable habitat for reintroduction of brown trout (Salmo trutta forma fario) in a lowland river: A modeling approach

Huge efforts have been made during the past decades to improve the water quality and to restore the physical habitat of rivers and streams in western Europe. This has led to an improvement in biological water quality and an increase in fish stocks in many countries. However, several rheophilic fish species such as brown trout are still categorized as vulnerable in lowland streams in Flanders (Belgium). In order to support cost‐efficient restoration programs, habitat suitability modeling can be used. In this study, we developed an ensemble of habitat suitability models using metaheuristic algorithms to explore the importance of a large number of environmental variables, including chemical, physical, and hydromorphological characteristics to determine the suitable habitat for reintroduction of brown trout in the Zwalm River basin (Flanders, Belgium), which is included in the Habitats Directive. Mean stream velocity, water temperature, hiding opportunities, and presence of pools or riffles were identified as the most important variables determining the habitat suitability. Brown trout mainly preferred streams with a relatively high mean reach stream velocity (0.2–1 m/s), a low water temperature (7–15°C), and the presence of pools. The ensemble of models indicated that most of the tributaries and headwaters were suitable for the species. Synthesis and applications. Our results indicate that this modeling approach can be used to support river management, not only for brown trout but also for other species in similar geographical regions. Specifically for the Zwalm River basin, future restoration of the physical habitat, removal of the remaining migration barriers and the development of suitable spawning grounds could promote the successful restoration of brown trout.     

Predation Threat Affects Behaviour and Habitat Use by Hatchery Brown Trout (Salmo Trutta L.) Juveniles

The effect of predation threat on behaviour and habitat use by brown trout (Salmo trutta L.) juveniles were studied in four indoor and six outdoor experimental channels. Northern pike (Esox lucius L.) was used as a predator. Brown trout were more active during the night compared to day, but in the presence of pike the increase in activity was less than in treatments without predator. Aggressive interactions between trout were higher during the day when pike was absent. In the presence of pike trout tended to use more coarse substrate classes as subdominant habitat suggesting increased use of cover and also higher or lower water depths. Our results increase knowledge of the effects of predation threat on habitat selection and behaviour by brown trout.     

Behavioural and physiological responses of brook trout Salvelinus fontinalis to midwinter flow reduction in a small ice-free mountain stream

This study presents an experimental analysis of the effects of midwinter flow reduction (50-75%, reduction in discharge in 4 h daily pulses) on the physical habitat and on behaviour and physiology of overwintering brook trout Salvelinus fontinalis in a small mountain stream. Flow reduction did not result in significant lowering of temperature or formation of surface or subsurface ice. The main findings were (1) daily movement by S. fontinalis increased (c. 2·5-fold) during flow reduction, but was limited to small-scale relocations (<10 m). (2) Undercut banks were the preferred habitat and availability of these habitats was reduced during flow reduction. (3) Although both experimental and reference fish did lose mass and condition during the experiment, no effects of flow reduction on stress indicators (blood cortisol or glucose) or bioenergetics (total body fat, water content or mass loss) were detected, probably because access to the preferred type of cover remained available. Like other salmonids, S. fontinalis moves little and seeks physical cover during winter. Unlike many of the more studied salmonids, however, this species overwinters successfully in small groundwater-rich streams that often remain ice-free, and this study identifies undercut banks as the critical winter habitat rather than substratum cover.     

Interaction for food and space between experimental populations of juvenile coho salmon (Oncorhynchus kisutch) and coastal cutthroat trout (Salmo clarki) in a laboratory stream

Populations of juvenile coho salmon and coastal cutthroat trout frequently cohabit small coastal streams in western North America. The pattern and mechanism of interactions for food and space between experimental populations of underyearlings of these two salmonids were examined at 3, 5 and 13 °C in a laboratory riffe/pool environment simulating winter and summer conditions. When tested separately at 13 °C, their habitat demands were similar and approximately 60–75% of either species occurred in pools. When tested together they segregated, with approximately 75% of coho in pools and up to 63% of cutthroat trout in riffles. In winter, at 3 °C, both species preferred pools and overhead cover, whether tested separately or together. At 5 °C, they partially segregated in a pattern similar to, but far less pronounced than, that in summer. Both species used similar forms of aggressive behaviour, although aggressive displays were more frequently used by coho, while nipping was more frequently used by cutthroat trout. Both salmonids were most aggressive when food was presented, irrespective of season, although coho responded with greater rapidity and intensity to feeding than did cutthroat trout. When tested together in summer, aggressiveness was high for both species, and agonistic interference by coho in pools and cutthroat trout in riffles appeared to largely account for their segregation. At 3 °C, aggression was low and both species weakly defended pools. At 5 °C, their aggression rose considerably and they partially segregated in a pattern resembling that at 13 °C. The mechanism of segregation between these two salmonids is clearly that of Nilsson's interactive type, which presumably functions to attenuate competition when streams are most likely to be resource limiting; typically, that is the late summer period of low flows and relatively high fish population densities.Based on a dissertation in partial fulfillment of the requirements for the degree of Doctor of Philosophy, Institute of Animal Resource Ecology, University of British Columbia, Vancouver, B.C.Ministry of Agriculture and Fisheries, Fisheries Research Division, P.O. Box 8324, Riccarton Christchurch, New Zealand     

The feeding and energetics of stream-resident trout in winter*

The feeding of brook and brown trout in a temperate Canadian stream was studied over two winters. Stomach analyses indicate that feeding continues throughout the winter. Both species had similar diets. Despite continuous feeding, condition factors declined significantly in early winter and remained low until the spring, suggesting an early-winter depletion of lipid reserves. Energy (calorific) intake was similar in winter to that found in late summer or spring and therefore was not the cause of low winter condition. The derived energy from feeding was insufficient to offset the costs of maintenance metabolism in the early winter period, for both immature and recently spawned fish. It is suggested that stream-dwelling trout suffer a metabolic deficit during acclimation to rapidly declining water temperatures in November/December. Such a phenomenon has important physiological implications for overwinter survival.     

Diet partitioning in sympatric Atlantic salmon and brown trout in streams with contrasting riparian vegetation

Prey intake by Atlantic salmon Salmo salar and brown trout Salmo trutta was measured across different riparian vegetation types: grassland, open canopy deciduous and closed canopy deciduous, in upland streams in County Mayo, Western Ireland. Fishes were collected by electrofishing while invertebrates were sampled from the benthos using a Surber sampler and drifting invertebrates collected in drift traps. Aquatic invertebrates dominated prey numbers in the diets of 0+ year Atlantic salmon and brown trout and 1+ year Atlantic salmon, whereas terrestrial invertebrates were of greater importance for diets of 1+ and 2+ year brown trout. Terrestrial prey biomass was generally greater than aquatic prey for 1+ and 2+ year brown trout across seasons and riparian types. Prey intake was greatest in spring and summer and least in autumn apart from 2+ year brown trout that sustained feeding into autumn. Total prey numbers captured tended to be greater for all age classes in streams with deciduous riparian canopy. Atlantic salmon consumed more aquatic prey and brown trout more terrestrial prey with an ontogenetic increase in prey species richness and diversity. Atlantic salmon and brown trout diets were most similar in summer. Terrestrial invertebrates provided an important energy subsidy particularly for brown trout. In grassland streams, each fish age class was strongly associated with aquatic, mainly benthic invertebrates. In streams with deciduous riparian canopy cover, diet composition partitioned between conspecifics with older brown trout associated with surface drifting terrestrial invertebrates and older Atlantic salmon associated with aquatic invertebrates with a high drift propensity in the water column and 0+ year fish feeding on benthic aquatic invertebrates. Deciduous riparian canopy cover may therefore facilitate vertical partitioning of feeding position within the water column between sympatric Atlantic salmon and brown trout. Implications for riparian management are discussed.     

Seasonality of macrophytes and interaction with flow in a New Zealand lowland stream

Introduced submerged macrophytes have come to dominate many shallow water bodies in New Zealand, and are a common component of many lowland streams. We investigated the seasonal variation of macrophyte abundance, its influence on flow and channel volume, and the implications of this on stream habitat and functioning in Whakapipi Stream, a typical lowland stream draining a predominantly agricultural catchment.Abundance of macrophytes over the summer was primarily controlled by the phenological cycles of the two dominant species. Mean minimum total macrophyte biomass (36 g m^–2) and cover (7%) occurred in winter (June and August, respectively), and mean maximum biomass (324 g m^–2), and cover (79%) occurred in late summer (March and February respectively). Egeria densa comprised the majority of both cover and biomass during the study period, except early summer (December) when Potamogeton crispus was prevalent in the shallow stream reaches.Macrophyte beds had a major impact on summer stream velocities, reducing average velocities by an estimated 41%. Stream cross-sectional area was maintained at relatively stable levels similar to that recorded over winter, when stream discharge was in the order of seven times greater. The mean velocity distribution coefficient (), and Manning's roughness coefficient (n) were dependent on and displayed a positive linear relationship with macrophyte abundance. The velocity distribution coefficient is recommended as a better indicator of macrophyte effects on velocity in natural streams, as it does not assume uniform velocity, channel depth and slope within the stream reach.Our study shows that submerged macrophytes play an important structuring role within the stream during the summer period, where macrophyte beds act as semi-permeable dams, retarding flow velocities and increasing stream depth and cross-sectional area. This promotes habitat heterogeneity by creating a greater range of flow velocity variation, and also provides large stable low-flow areas. Other likely ecosystem effects resulting from macrophyte/velocity interactions include increased sedimentation, potential for nutrient processing and increased primary production, both by macrophytes and attached epiphyton. The complex architecture of submerged macrophytes and their influence on stream flow may also provide an increased diversity of habitat for other aquatic biota. We propose that management of degraded lowland streams such as the Whakapipi Stream to maintain stretches with moderate quantities of submerged macrophytes interspersed with shaded areas would optimise stream health during low summer flows.     

Diel Habitat Partitioning by Bull Charr and Cutthroat Trout During Fall and Winter in Rocky Mountain Streams

We used underwater observation to determine diel habitat partitioning between bull charr, Salvelinus confluentus, and cutthroat trout, Oncorhynchus clarki, during fall and winter (0.1–8.3°C) in two Rocky Mountain streams that differed in habitat availability. The majority (>70%) of both species emerged from concealment cover at night, though bull charr exhibited a greater tendency for nocturnal behavior than cutthroat trout. Differences in day and night counts were most pronounced at temperatures <3°C, when very few fish of either species were observed in the water column during the day, but both species were common at night. Both species used concealment cover of large woody debris and boulder substrate crevices in deep pools during the day. At night, fish emerged from cover and habitat use shifted to shallow water with low cover. Microhabitat partitioning among species and size classes occurred at night, cutthroat trout moving into shallower, faster water that was farther from cover compared to bull charr. Smaller fish of both species occupied focal positions in slower, shallower water closer to the substrate than larger fish. Large, mixed-species aggregations also were common in beaver ponds both day and night. High variation in diel and site-specific winter habitat use suggests the need for caution in developing habitat suitability criteria for salmonids based solely on daytime observations or on observations from a few sites. Our results support the need to incorporate nocturnal habitat use and partitioning in studies of salmonid ecology.     

A mechanistic assessment of seasonal microhabitat selection by drift-feeding rainbow trout <Emphasis Type="Italic">Oncorhynchus mykiss</Emphasis> in a Southwestern headwater stream

The positioning of fishes within a riverscape is dependent on the proximity of complementary habitats. In this study, foraging and non-foraging habitat were quantified monthly over an entire year for a rainbow trout (Oncorhynchus mykiss) population in an isolated, headwater stream in southcentral New Mexico. The stream follows a seasonal thermal and hydrologic pattern typical for a Southwestern stream and was deemed suitable for re-introduction of the native and close relative, Rio Grande cutthroat trout (O. clarkii virginalis). However, uncertainty associated with limited habitat needed to be resolved if repatriation of the native fish was to be successful. Habitat was evaluated using resource selection functions with a mechanistic drift-foraging model to explain trout distributions. Macroinvertebrate drift was strongly season- and temperature-dependent (lower in winter and spring, higher in summer and fall). Models identified stream depth as the most limiting factor for habitat selection across seasons and size-classes. Additionally, positions closer to cover were selected during the winter by smaller size-classes (0, 1, 2), while net energy intake was important during the spring for most size-classes (0, 1, 2, 3). Drift-foraging models identified that 81% of observed trout selected positions that could meet maintenance levels throughout the year. Moreover, 40% of selected habitats could sustain maximum growth. Stream positions occupied by rainbow trout were more energetically profitable than random sites regardless of season or size-class. Larger size-classes (3, 4+) were energetically more limited throughout the year than were smaller size-classes. This research suggests that habitat in the form of deep pools is of paramount importance for rainbow trout or native cutthroat trout.     

Seasonal dietary shifts enhance parasite transmission to lake salmonids during ice cover

Changes in abiotic and biotic factors between seasons in subarctic lake systems are often profound, potentially affecting the community structure and population dynamics of parasites over the annual cycle. However, few winter studies exist and interactions between fish hosts and their parasites are typically confined to snapshot studies restricted to the summer season whereas host‐parasite dynamics during the ice‐covered period rarely have been explored. The present study addresses seasonal patterns in the infections of intestinal parasites and their association with the diet of sympatric living Arctic charr (Salvelinus alpinus) and brown trout (Salmo trutta) in Lake Takvatn, a subarctic lake in northern Norway. In total, 354 Arctic charr and 203 brown trout were sampled from the littoral habitat between June 2017 and May 2018. Six trophically transmitted intestinal parasite taxa were identified and quantified, and their seasonal variations were contrasted with dietary information from both stomachs and intestines of the fish. The winter period proved to be an important transmission window for parasites, with increased prevalence and intensity of amphipod‐transmitted parasites in Arctic charr and parasites transmitted through fish prey in brown trout. In Arctic charr, seasonal patterns in parasite infections resulted mainly from temporal changes in diet toward amphipods, whereas host body size and the utilization of fish prey were the main drivers in brown trout. The overall dynamics in the community structure of parasites chiefly mirrored the seasonal dietary shifts of their fish hosts.     

Influence of hydrologic attributes on brown trout recruitment in low-latitude range margins

Factors controlling brown trout Salmo trutta recruitment in Mediterranean areas are largely unknown, despite the relevance this may have for fisheries management. The effect of hydrological variability on survival of young brown trout was studied during seven consecutive years in five resident populations from the southern range of the species distribution. Recruit density at the end of summer varied markedly among year-classes and rivers during the study period. Previous work showed that egg density the previous fall did not account for more than 50% of the observed variation in recruitment density. Thus, we expected that climatic patterns, as determinants of discharge and water temperature, would play a role in the control of young trout abundance. We tested this by analyzing the effects of flow variation and predictability on young trout survival during the spawning to emergence and the summer drought periods. Both hatching and emergence times and length of hatching and emergence periods were similar between years within each river but varied considerably among populations, due to differences in water temperature. Interannual variation in flow attributes during spawning to emergence and summer drought affected juvenile survival in all populations, once the effect of endogenous factors was removed. Survival rate was significantly related to the timing, magnitude and duration of extreme water conditions, and to the rate of change in discharge during hatching and emergence times in most rivers. The magnitude and duration of low flows during summer drought appeared to be a critical factor for survival of young trout. Our findings suggest that density-independent factors, i.e., hydrological variability, play a central role in the population dynamics of brown trout in populations from low-latitude range margins. Reported effects of hydrologic attributes on trout survival are likely to be increasingly important if, as predicted, climate change leads to greater extremes and variability of flow regimes.