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.     

Brown trout redd superimposition in relation to spawning habitat availability

Abstract – The relationship between redd superimposition and spawning habitat availability was investigated in the brown trout (Salmo trutta L.) population inhabiting the river Castril (Granada, Spain). Redd surveys were conducted in 24 river sections to estimate the rate of redd superimposition. Used and available microhabitat was evaluated to compute the suitable spawning habitat (SSH) for brown trout. After analysing the microhabitat characteristics positively selected by females, SSH was defined as an area that met all the following five requirements: water depth between 10 and 50 cm, mean water velocity between 30 and 60 cm s^?1, bottom water velocity between 15 and 60 cm s^?1, substrate size between 4 and 30 mm and no embeddedness. Simple regression analyses showed that redd superimposition was not correlated with redd numbers, SSH or redd density. A simulation‐based analysis was performed to estimate the superimposition rate if redds were randomly placed inside the SSH. This analysis revealed that the observed superimposition rate was higher than expected in 23 of 24 instances, this difference being significant (P < 0.05) in eight instances and right at the limit of statistical significance (P = 0.05) in another eight instances. Redd superimposition was high in sections with high redd density. High superimposition however was not exclusive to sections with high redd density and was found in moderate‐ and low‐redd‐density sections. This suggests that factors other than habitat availability are also responsible for redd superimposition. We argue that female preference for spawning over previously excavated redds may be the most likely explanation for high superimposition at lower densities.     

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.     

Differences in longitudinal distribution patterns along a Honshu stream of brown trout <Emphasis Type="Italic">Salmo trutta</Emphasis>, white-spotted charr <Emphasis Type="Italic">Salvelinus leucomaenis</Emphasis> and masu salmon <Emphasis Type="Italic">Oncorhynchus masou</Emphasis>

Brown trout Salmo trutta were first introduced into Japan in 1892, and they currently naturally reproduce in several rivers in Honshu and Hokkaido, Japan. Although negative impacts of brown trout introductions on native salmonid fishes have been documented in some Hokkaido rivers, studies of ecological interactions between brown trout and native salmonid fishes on Honshu are limited. In this study, we describe the longitudinal distribution patterns of introduced brown trout, white-spotted charr Salvelinus leucomaenis and masu salmon Oncorhynchus masou in a 4 km stretch of a stream in central Honshu. Underwater observations were conducted in all pools within upstream, middle and downstream sections (190–400 m in length) of this stretch in order to estimate the densities of these species. Only white-spotted charr was observed in the upstream section, while brown trout and masu salmon were observed in the middle and downstream sections. Masu salmon densities, however, were much lower than brown trout densities. In the downstream section, white-spotted charr was absent. These results are consistent with results from previous studies of Hokkaido rivers, where it was found that white-spotted charr in low-gradient areas tend to be displaced by brown trout.     

Modelling effects of climate change on Michigan brown trout and rainbow trout: Precipitation and groundwater as key predictors

Understanding how changes in stream temperature affect survival and growth of coldwater fishes, including brown trout (Salmo trutta) and rainbow trout (Oncorhynchus mykiss), is important for conserving coldwater stream fisheries in a changing climate. However, some contemporary stream temperature models assume spatially uniform (i.e. unrealistic) air–stream temperature relationships or demand hydrometeorological predictors (e.g. solar radiation and convection) that are expensive and often impractical for fisheries managers to measure. As such, we produced a relatively cost-effective, management-relevant modelling approach for predicting effects of changes in air temperature, precipitation and groundwater inputs on stream temperature and, consequently, the survival and growth of brown trout and rainbow trout in Michigan, USA. We found that precipitation- and groundwater-corrected stream temperature models (mean adjusted R² = .77, range = 0.65–0.88) performed better than linear air–stream temperature models (mean adjusted R² = .59, range = 0.21–0.80). Stream temperature was projected to increase by 0.07–3.88°C (1%–22%) with simulated changes in air temperature, precipitation and groundwater inputs. The greatest warming was predicted for surface runoff-dominated sites with limited groundwater-driven thermal buffering, where thermal habitat suitability for salmonid survival and growth declined 20%–40%. However, groundwater-dominated sites may not be immune to temperature warming, especially if groundwater temperature increases or groundwater inputs decline in a changing climate. Our modelling approach provides a reliable, cost-effective method for predicting effects of climate change on brown trout and rainbow trout survival and growth, allowing for strategic management actions to increase the thermal resilience and sustainability of salmonid populations (e.g. groundwater conservation and riparian/watershed rehabilitation).     

Diel microhabitat use of age-0 bull trout in Indian Creek, Washington

Abstract – We studied diel microhabitat use at the focal point of age‐0 bull trout, Salvelinus confluentus, in Indian Creek, Washington during mid‐summer and fall of 1997. Microhabitat variables included water depth and velocity, distance from the stream bottom, habitat and refuge use, substrate type, and substrate embeddedness. Age‐0 fish were located over fines and gravel substrates in shallow, low‐velocity water near stream margins, but were located in shallower water at night. Bull trout were highly associated with loose substrate, and used the substrate interstices for refuge cover. Diurnal bull trout counts decreased and no age‐0 fish were observed after 15 September at water temperatures below 6.1 °C. Nocturnal counts remained relatively constant throughout the study. Our results suggest that age‐0 bull trout surveys be conducted at night when summer water temperatures begin to decline.     

Comparing estimates of number of breeders Nb based on microsatellites and single nucleotide polymorphism of three groups of brown trout (Salmo trutta L.)

Microsatellites (10 MSs) and single nucleotide polymorphism (3871 SNPs) were analysed in three closely related, single‐cohort groups of brown trout, Salmo trutta L. Autumn (W/0+) and subsequent year (W/1+) samples of brown trout of the same stream were retrieved using electric fishing, and one sample of artificially reared brown trout (H/1+), bred from wild parents, was collected from a hatchery. The number of breeders, N_b, was estimated based on each total sample set and from subsets of loci by means of the software LDNe, and by means of Colony2, NeEstimator and ONeSAMP softwares. MS‐based estimates were consequently higher than those based on SNPs. ML‐Relate and Colony2 software was used to estimate putative full‐sib pairs, and there was considerable variation between methods. For monitoring species and populations, both methods are recommended, but to compare N_b across samples and years, the same markers of MSs or SNPs must be used as estimates varied across markers within sample.     

Predation on native sculpin by exotic brown trout exceeds that by native cutthroat trout within a mountain watershed (Logan,UT, USA)

We explored potential negative effects of exotic brown trout (Salmo trutta) on native sculpin (Cottus sp.) on the Logan River, Utah, USA by (i) examining factors most strongly correlated with sculpin abundance (e.g., abiotic conditions or piscivory?), (ii) contrasting the extent of brown trout predation on sculpin with that by native cutthroat trout (Oncorhynchus clarkii utah) and (iii) estimating the number of sculpin consumed by brown trout along an elevational gradient using bioenergetics. Abundance of sculpin across reaches showed a strong (r ≥ 0.40) and significant (P < 0.05) correlation with physical variables describing width (positive) and gradient (negative), but not with abundance of piscivorous brown trout or cutthroat trout. In mainstem reaches containing sculpin, we found fish in 0% of age‐1, 10% of age‐2 and 33% of age‐3 and older brown trout diets. Approximately 81% of fish consumed by brown trout were sculpin. Despite a similar length–gape relationship for native cutthroat trout, we found only two fish (one sculpin and one unknown) in the diets of native cutthroat trout similar in size to age‐3 brown trout. Based on bioenergetics, we estimate that an average large (> 260 mm) brown trout consumes as many as 34 sculpin per year. Nevertheless, results suggest that sculpin abundance in this system is controlled by abiotic factors and not brown trout predation. Additional research is needed to better understand how piscivory influences brown trout invasion success, including in‐stream experiments exploring trophic dynamics and interactions between brown trout and native prey under different environmental conditions.     

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.     

Experimental evaluation of rainbow trout Oncorhynchus mykiss predation on longnose dace Rhinichthys cataractae

Laboratory and in‐stream enclosure experiments were used to determine whether rainbow trout Oncorhynchus mykiss influence survival of longnose dace Rhinichthys cataractae. In the laboratory, adult rainbow trout preyed on longnose dace in 42% of trials and juvenile rainbow trout did not prey on longnose dace during the first 6 h after rainbow trout introduction. Survival of longnose dace did not differ in the presence of adult rainbow trout previously exposed to active prey and those not previously exposed to active prey ( = 0.28, P = 0.60). In field enclosures, the number of longnose dace decreased at a faster rate in the presence of rainbow trout relative to controls within the first 72 h, but did not differ between moderate and high densities of rainbow trout (F_2,258.9 = 3.73, P = 0.03). Additionally, longnose dace were found in 7% of rainbow trout stomachs after 72 h in enclosures. Rainbow trout acclimated to the stream for longer periods had a greater initial influence on the number of longnose dace remaining in enclosures relative to those acclimated for shorter periods regardless of rainbow trout density treatment (F_4,148.5 = 2.50, P = 0.04). More research is needed to determine how predation rates will change in natural environments, under differing amounts of habitat and food resources and in the context of whole assemblages. However, if rainbow trout are introduced into the habitat of longnose dace, some predation on longnose dace is expected, even when rainbow trout have no previous experience with active prey.     

Habitat use by Thymallus thymallus in a chalk stream and implications for habitat management

Abstract Results of a number of studies on the interactions between grayling, Thymallus thymallus L., and its habitat in tributaries of the River Itchen, a chalk stream in Hampshire, southern England are reported. These include an investigation into the effect of riparian shading on T. thymallus growth and population density in relation to the development of in‐stream macrophyte cover, and the use of a multi‐point decoder system to record micro‐habitat use and preference of individual T. thymallus. In all stream sections, T. thymallus recruitment fluctuated greatly. Densities were generally low often restricting meaningful comparisons. However, where large differences occurred, wooded sections, with less aquatic macrophyte cover, generally had higher densities of T. thymallus. All age groups of T. thymallus (0+, 1+ and 2+) tended to occupy hard gravel substratum, both by day and by night in the autumn and winter, and used deeper and faster water with increasing age. The 1+ and 2+ groups were generally found in water 40–70‐cm deep with a velocity between 0.3 and 0.5 m s^?1, whilst the 0+ group showed a preference for shallower water with reduced velocity at night, particularly in the winter. The results are discussed in relation to habitat management where T. thymallus occur with other salmonids.     

Country‐wide analysis of large wood as a driver of fish abundance in Swedish streams: Which species benefit and where?

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Genetic characterization of naturalized populations of brown trout Salmo trutta L. in southern Chile using allozyme and microsatellite markers

This study describes the genetic structure of five naturalized populations of brown trout in southern Chile using allozyme and microsatellite markers to establish levels of intra‐ and interpopulation genetic variability and divergence. Fourteen enzymatic systems were used comprising 20 loci and three microsatellite loci specific to brown trout. The genetic variability values (allozymes, P=20–35%, average=27%, H_O=0.118–0.160, average=0.141; microsatellites, P=33.3–100%, average=66.66%, H_O=0.202–0.274, average=0.229) are similar to values described in other naturalized populations of brown trout present in Chile, but higher than those observed in European populations of this species. Values of total genetic diversity (H_T) (allozymes=0.1216 and microsatellites=0.3504) and relative genetic divergence (G_ST) (allozymes=9.5% and microsatellites=15%) were also similar to the results obtained in previous studies of Chilean populations of brown trout. These values, when compared with those obtained in Europe, proved to be similar for H_T but lower for G_ST. The low interpopulational genetic differentiation was in accordance with the small genetic distance observed between the populations analysed (D Nei=0.004–0.025). On the other hand, the high frequency of one of the two alternative alleles of the phylogeographic marker locus LDH‐5* in the populations analysed (LDH‐5*90>0.84) would indicate a European origin, in particular Atlantic as opposed to Mediterranean, for the brown trout introduced into Chile. The high levels of genetic variability suggest a mixed origin for the naturalized brown trout in Chile, which could have originated either before or during the introduction process. Nevertheless, the low level of genetic differentiation between populations could reflect the short lapse of time in evolutionary terms, during which populations introduced into Chile have been exposed to different evolutionary forces, and which has not been sufficiently long to produce greater genetic differentiation between populations.     

Effect of fine wood on juvenile brown trout behaviour in experimental stream channels

In‐stream wood can increase shelter availability and prey abundance for stream‐living fish such as brown trout, Salmo trutta, but the input of wood to streams has decreased in recent years due to harvesting of riparian vegetation. During the last decades, fine wood (FW) has been increasingly used for biofuel, and the input of FW to streams may therefore decrease. Although effects of in‐stream FW have not been studied as extensively as those of large wood (LW), it is probably important as shelter for small‐sized trout. In a laboratory stream experiment, we tested the behavioural response of young‐of‐the‐year wild brown trout to three densities of FW, with trout tested alone and in groups of four. Video recordings were used to measure the proportion of time allocated to sheltering, cruising and foraging, as well as the number of aggressive interactions and prey attacks. Cruising activity increased with decreasing FW density and was higher in the four‐fish groups than when fish were alone. Foraging decreased and time spent sheltering in FW increased with increasing FW density. Our study shows that juvenile trout activity is higher in higher fish densities and that trout response to FW is related to FW density and differs from the response to LW as reported by others.     

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.     

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.     

River bed construction: impact and habitat restoration for juvenile Atlantic salmon, Salmo salar L., and brown trout, Salmo trutta L.

The River Soya, Norway, was canalized for agricultural purposes. In order to compensate for damage to the Atlantic salmon, salmo salar L. and brown trout, Salmo trutta L. populations, different weirs were built. The aims of this study were firstly to analyse the effects of weirs covering the river bank and entire river bottom with blasted stones and secondly, to analyse the effects of sediments transported by freshets on the downstream salmon and trout populations after canalization. Restoration of the river bottom with blasted stones provided salmon with more substrate spaces. Densities of trout increased after the river bank was covered with stones. Sediments transported downstream from the canalized river stretch decreased the densities of juvenile salmon and trout.     

Assessing near‐bed velocity in a recruiting population of the endangered freshwater pearl mussel (Margaritifera margaritifera) in Ireland

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Reproductive biology of Oreochromis mossambicus populations of man-made lakes in Sri Lanka: a comparative study

Abstract. The sex ratio, mean size at maturity (L_m), gonosomatic index (GSI), frequency of distribution of egg diameter of the mature ovaries and fecundity of Oreochromis mossambicus (Peters) populations of 12 man-made lakes were studied using commercial landings from the gill-net fishery. The overall sex ratio was 1·0:0·88 of females to males and that of the individual populations varied from 1·0:0·54 to 1·0:1·34. The L_m ranged from 16·0 to 21·0 cm (TL) and was found to be correlated to the size of the water-body. The GSI of the females in stage V, the stage prior to spawning, did not differ from each other, except in one reservoir. The variations in the mean egg size (μ) of different populations were negatively correlated to the fecundity of a Ash of standard weight (FW₂₀₀) of the different populations and the relationship was, y =3487·4– 1·14FW₂₀₀(df = 10; r= -0·64; P < 0·05) The overall fecundity ranged from 318 to 3169 in fish ranging in body length (TL) and body weight from 16·5 to 32·0 cm and 80 to 651 g respectively. Fecundity was linearly related to body weight and curvilinearly to body length. The exponent b of the fecundity (F) length (1) regressions (F = aL^b) of the different populations varied between 1·20 and 3·83. The overall relationship of fecundity to body weight (W) and body length (1) for O. mossambicus reservoir populations in Sri Lanka were: F = 3·23W + 357·8 (df = 203; r= 0·73; P < 0·001) F = l·52L^2·11(df = 203; r= 0·68; P < 0·001) The variations in the theoretical fecundity calculated for a standard size individual of 200 g in weight (FW₂₀₀) of the reservoir populations were correlated to the mean fishing pressure (X) exerted on each population according to the equations. (a) FW₂₀₀= 10–24X+ 896·8 (df 9; r= 0·58; P < 0·05) (b) FW₂₀₀= 1200 – 422·76e^?0·102X (df 9; P < 0·05)     

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.