Seasonal movement,growth and survival of brook trout in sympatry with brown trout in Midwestern US streams

Seasonal patterns in growth, survival and movement of brook trout Salvelinus fontinalis were monitored in two southeastern Minnesota streams divided into study reaches based on brown trout Salmo trutta abundance. We estimated survival and movement while testing for effects of stream reach and time using a multistrata Cormack–Jolly–Seber model in Program MARK. Multistrata models were analysed for three age groups (age‐0, age‐1 and age‐2+) to estimate apparent survival, capture probability and movement. Survival varied by time period, but not brown trout abundance and was lower during flood events. Age‐0 brook trout emigrated from reaches with low brown trout abundance, whereas adult brook trout emigrated from downstream brown trout‐dominated reaches. Growth was highest in spring and summer and did not differ across streams or reaches for the youngest age classes. For age‐2+ brook trout, however, growth was lower in reaches where brown trout were abundant. Interspecific interactions can be age or size dependent; our results show evidence for adult interactions, but not for age‐0. Our results suggest that brook trout can be limited by both environmental and brown trout interactions that can vary by season and life stage.     

Seasonal habitat use by brook trout,Salvelinus fontinalis (Mitchill), in a second-order stream

Abstract Seasonal habitat use by over-yearling and under-yearling brook trout, Salvelinus fontinalis (Mitchill), was examined in a second-order stream in north-central Pennsylvania, USA. The habitat occupied by brook trout and available habitat were determined in a 0.5-km stream reach during the spring, summer and autumn of 1989 and the spring and summer of 1990. Cover, depth, substrate and velocity were quantified from over 2000 observations of individual brook trout. Habitat used by under-yearling brook trout was more uniform between seasons and years than that used by over-yearling brook trout. Over-yearling brook trout occupied areas with more cover and greater depth than did under-yearling brook trout, suggesting ontogenetic shifts in these variables. Differences for velocity and substrate were not as great as those for cover and depth. The selection of areas with low water velocities governed trout habitat use in spring, whereas cover and depth were the most important habitat variables in summer and autumn. Principal component analysis showed that available habitat and trout habitat centroids diverged most in spring, indicating that habitat selection by brook trout may be greatest at this time.     

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.     

Ontogenetic and spatial variations in brown trout habitat selection

Ayllón D, Almodóvar A, Nicola GG, Elvira B. Ontogenetic and spatial variations in brown trout habitat selection.
Ecology of Freshwater Fish 2010: 19: 420–432. © 2010 John Wiley & Sons A/S Abstract – Habitat quality and quantity determine many biological processes and traits that directly affect the population dynamics of stream fishes. Understanding how habitat selection is adjusted to different ecological conditions is essential to improve predictive modelling of population dynamics. We describe brown trout Salmo trutta summer habitat selection patterns through univariate and multivariate habitat selection functions across defined river reach typologies. We sampled 44 sites and performed a principal component analysis that defined eight reach types differing in both local site and catchment‐scale physical features. We observed ontogenetic changes in habitat selection, as trout preferred deeper and slower flowing water as they increased in size. Likewise, selectivity for different types of structural habitat elements changed through ontogeny. Both patterns were consistent across reach types. Moreover, we detected spatial variations in habitat selection patterns within age‐classes among different reach types. Our results indicate that brown trout is a habitat generalist and suggest that spatial variations in habitat selection patterns are driven by physical and environmental factors operating at multiple spatial scales.     

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.     

The loss of large wood affects rocky mountain trout populations

Western U.S. rivers are currently influenced by legacy effects of reduced large wood (LW) loading and retention that has substantially reduced in‐stream habitat complexity. Large wood is typically associated with streams in undisturbed old‐growth forest and in the correct geomorphic context can drastically alter stream and valley habitat complexity. Streams with LW are typically multichannel and depositional, while streams lacking LW, due to relatively recent wildfire or logging (<200 years ago), are usually single channelled and erosional. We compared population biomass and individual growth rates of Brook Trout Salvelinus fontinalis in streams across a gradient of wood volumes. At both the square metre and valley length scales, standing stock biomass of aquatic invertebrates was the best predictor of trout biomass. However, at the valley scale, the number of pools was important in predicting trout biomass in combination with standing stock biomass of aquatic invertebrates. Individual growth rates of age‐1 Brook Trout were negatively affected by increasing density; however, growth rates for the largest and smallest individuals at each site were unaffected by density. Our results suggest the pool habitat created by LW acts synergistically with prey availability to dramatically increase trout populations. However, in streams lacking LW, negative effects of detrimental land use practices have persisted >100 years, suggesting that recovering lost animal production in mountain stream networks will only occur at decadal to century time scales.     

Cohort‐specific variation in juvenile coho salmon habitat use

In this study, we examined summer and fall freshwater rearing habitat use by juvenile coho salmon (Oncorhynchus kisutch) in the quickly urbanising Big Lake drainage in south‐central Alaska. Habitat use was assessed by regressing fish count data against habitat survey information across thirty study sites using generalised linear mixed models. Habitat associations were examined by age‐0 and age‐1+ cohorts separately, providing an opportunity to compare habitat use across different juvenile coho salmon life stages during freshwater rearing. Regression results indicated that the age‐0 cohorts were strongly associated with shallow, wide stream reaches with in‐stream vegetation, whereas age‐1+ cohorts were associated with deeper stream reaches. Furthermore, associations between fork length and habitat characteristics suggest cohort‐specific habitat use patterns are distinct from those attributable to fish size. Habitat use information generated from this study is being used to guide optimal fish passage restoration planning in the Big Lake drainage. Evidence for habitat use partitioning by age cohort during freshwater juvenile rearing indicates that pooling age cohorts into a single “juvenile” stage for the purposes of watershed management may mask important habitat use dynamics.     

Effects of reduced summer flows on the brook trout population and temperatures of a groundwater‐influenced stream

Withdrawal of water from streams and groundwater is increasing in Midwestern North America and is a potential threat to coldwater fishes. We examined the effects of summer water withdrawals on brook trout Salvelinus fontinalis populations and water warming rates by diverting 50–90% of summer baseflow from a 602‐m treatment zone (TZ) in a groundwater‐influenced Michigan stream during 1991–1998. We compared density of brook trout in fall, and spring‐to‐fall growth and survival of brook trout, between the TZ and an adjacent reference zone (RZ) whose flows were not altered. Flow reductions had no significant effects on the density of brook trout in fall or spring‐to‐fall survival of brook trout. However, spring‐to‐fall growth of brook trout in the TZ declined significantly when 75% flow reductions occurred. Cold upstream temperatures and the relatively short study reach kept thermal habitat conditions excellent for brook trout in the TZ throughout the dewatering experiments. These findings suggest that brook trout can tolerate some seasonal loss of physical habitat if temperature conditions remain suitable. In summer 1999, we experimentally assessed the influence of flow reduction on the warming rate through the TZ by diverting from 0% to 90% of flow around the TZ in 3‐ or 4‐day trials on a randomised schedule. Average daily temperature increased exponentially as stream flows declined from normal summer levels. Our findings suggest the risk of trout habitat loss from dewatering is potentially large and proportional to the magnitude of withdrawal, especially as thermal conditions approach critical levels for trout.     

Season-specific survival rates and densities of coastal cutthroat trout across stream sizes in southwestern British Columbia

Variation in seasonal survival rates, densities and growth rates of coastal cutthroat trout (Oncorhynchus clarkii clarkii) were assessed across a size gradient of small, forested streams in the Pacific Northwest. We used a robust, mark-recapture study, stratified seasonally to estimate monthly survival rates of trout in coastal British Columbia (not including young-of-the-year). Survival estimates showed that the summer season had the lowest monthly survival rates (0.907) across all streams in our study (0.927 remainder of year). Within the size range of the seven small streams studied, low-flow habitat availability (defined by residual pool depth in summer) was the best predictor of mean monthly survival rates, supporting the hypothesis that trout survival increases with the quantity of aquatic habitat, particularly depths of residual pools. In addition, there was an asymptotic relation between water depth and survival rates, where beyond ~20 cm of residual pool depth, greater depth did not confer greater rates of trout survival. Growth rates in all but the largest stream were also lowest during summer. While densities tended to be higher in streams with greater residual depth, this was not significant. Body mass in a given season was a good predictor of survival to the next sampling period. The distribution and success of resident cutthroat trout populations in small streams appear to be constrained by summer low-flow periods and specific geomorphologies that support deeper pools.     

The complementary role of lentic and lotic habitats for Arctic grayling in a complex stream‐lake network in Arctic Alaska

Lakes can be important to stream dwelling fishes, yet how individuals exploit habitat heterogeneity across complex stream‐lake networks is poorly understood. Furthermore, despite growing awareness that intermittent streams are widely used by fish, studies documenting the use of seasonally accessible lakes remain scarce. We studied Arctic grayling (Thymallus arcticus) in a small seasonally flowing (June–October) stream‐lake network in Alaska using PIT telemetry. Overall, 70% of fish visited two lakes, 8% used a single lake, and 22% used only stream reaches. We identified five distinct behavioural patterns that differed in dominant macrohabitat used (deep lake, shallow lake or stream reaches), entry day into the network and mobility. Some juvenile fish spent the entire summer in a shallow seasonally frozen lake (average 71 days), whereas others demonstrated prospecting behaviour and only entered the stream channel briefly in September. Another group included adult and juvenile fish that were highly mobile, moving up to 27 km while in the 3‐km stream‐lake network, and used stream reaches extensively (average 59 days). Lentic and lotic habitats served differing roles for individuals, some fish occupied stream reaches as summer foraging habitat, and other individuals used them as migration corridors to access lakes. Our study emphasises the importance of considering stream‐lake connectivity in stream fish assessments, even to shallow seasonally frozen habitats not widely recognised as important. Furthermore, we demonstrate that individuals may use temporary aquatic habitats in complex and changing ways across ontogeny that are not captured by typical classifications of fish movement behaviour.     

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.     

Patch size but not short‐term isolation influences occurrence of westslope cutthroat trout above human‐made barriers

Habitat fragmentation in aquatic systems has led to widespread isolation of stream fishes. Metapopulation theory predicts that persistence is directly related to local patch size and its characteristics, but because these relationships tend to be taxon‐specific, empirical data are important. We assembled 246 observations of occurrence of westslope cutthroat trout (WCT), a taxon of concern in the western U.S. and Canada, in stream networks isolated for up to 100 years (median 40 years) above human‐made barriers, mostly culverts, at road crossings within U.S. National Forests. We used logistic regression to analyse how WCT occurrence varied with patch size, isolation time and stream‐level covariates. Occurrence was positively related to stream length and habitat quality within the isolated stream network and negatively related to elevation and channel gradient. Unexpectedly, the probability of occurrence was not related to how long a habitat patch had been isolated. At the median elevation (1354 m) and channel gradient (14%), and where habitat quality was poor, WCT were likely to occur (probability >0.5) if an isolated stream network was at least 1.7 km. If habitat quality was high, about 0.2 km of habitat produced the same probability. Although there are important limitations, this analysis provides the first empirical estimate for how patch size and patch‐level characteristics influence persistence of WCT in isolated stream networks.     

Census and effective population sizes of white‐spotted charr (Salvelinus leucomaenis) in a fragmented landscape

Several habitat models have been proposed to predict population size for stream fishes and to guide habitat assessment and monitoring techniques. However, most models do not incorporate the potential advantage of molecular genetic markers. We conducted a field survey and microsatellite DNA analyses to quantify the relationships among genetic diversity, census/effective population size and habitat variables in fragmented populations of white‐spotted charr (Salvelinus leucomaenis). The census population size significantly increased with the stream length, the number of pools and a pool‐riffle sequence index, a proxy for channel‐unit habitat type complexity within reaches. Population density was correlated with the pool‐riffle sequence index only. Genetic diversity and effective population size were not correlated with the habitat variables or census population size. There was a lack of isolation‐by‐distance population structure in the studied populations. Our results suggest that stream length and the number of pools within reaches associated with habitat complexity are the habitat variables that explain the majority of variation in population size of white‐spotted charr. Our findings provide further evidence that census population size per se is a poor indicator of the inclusive genetic diversity within populations in a fragmented landscape.     

Modelling functional fish habitat connectivity in rivers: A case study for prioritizing restoration actions targeting brown trout

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Seasonal effects of macrophyte growth on rainbow trout habitat availability and selection in a low‐gradient,groundwater‐dominated river

Rainbow trout habitat use is often described in high‐gradient, runoff‐driven, heterotrophic streams where geomorphic features and overhanging riparian vegetation provide channel complexity and cover. However, many rainbow trout populations thrive in rivers with contrasting aquatic habitat. We describe rainbow trout habitat use in a low‐gradient, groundwater‐dominated tailwater river where river flow management and macrophyte growth and senescence largely govern available trout habitat. In the summers of 2013 and 2014, available aquatic habitat (depth, velocity, macrophyte cover, substrate size) was quantified, while individual trout location was determined by radio telemetry and linked to environmental variables. Detailed habitat surveys indicate that macrophyte cover increases throughout the summer and is a strong determinant of in‐stream habitat characteristics. Paired logistic regression shows that adult rainbow trout prefer greater depths. Water depth increases with macrophyte abundance at both reach and local scales as plants restrict flow, and available trout habitat is linked to this seasonal pattern. When macrophyte abundance is high, adult trout show secondary preference for localised areas of lower macrophyte cover but otherwise show no selectivity for macrophyte cover, velocity or substrate size. Results suggest that submerged aquatic plants increased the quantity and quality of rainbow trout habitat as a source of channel complexity and cover. Macrophytes may play a similar role in other low‐gradient streams and should not be overlooked by fisheries managers considering habitat suitability.     

Are native brook charr and introduced rainbow trout differentially adapted to upstream and downstream reaches?

Abstract— Due to species introductions, brook charr (Salvelinus fontinalis) and rainbow trout (Oncorhynchus mykiss) occur together in many North American streams and typically exhibit a pattern of distribution in which brook charr numerically dominate headwaters and rainbow trout dominate downstream reaches. It has been suggested that 1) the two species compete or 2) the two species do not compete because they are differentially adapted to environmental conditions found in upstream and downstream zones. We assessed whether there were differences in growth and macrohabitat (pool, run and riffle) selection of brook charr and rainbow trout in upper, middle and lower stream zones of a small Pennsylvania stream. Brook charr and rainbow trout placed in replicate paired enclosures set in upstream and downstream reaches showed no significant differences in growth and survival rates upstream, but brook charr had significantly greater growth rates than rainbow trout downstream. Enclosed fish and free-ranging fish both had negative growth rates during the summer. Enclosed fish lost significantly less weight than free-ranging fish. Instantaneous growth rates of free-ranging adult brook charr and rainbow trout from May to August were negative for both species in all stream zones. Underwater observations of adult brook charr and rainbow trout showed both species occurred significantly more often in pool macrohabitats than expected on the basis of macrohabitat availability, except for rainbow trout in the upstream zone. The proportion of pool macrohabitat was not significantly different among stream zones. Brook charr do not appear to be better adapted to upstream environments in Powdermill Run based on growth, survival and macrohabitat selection during summer. Negative biotic interactions acting along with differential environmental adaptations may explain the pattern of distribution of brook charr and rainbow trout in streams, but long-term transplant experiments with additional life stages will be necessary to examine this hypothesis.     

Broad‐scale habitat classification variables predict maximum local abundance for native but not non‐native trout in New York streams

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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.     

Summer habitat and movements of juvenile salmonids in a coastal river of Washington State

We investigated the summer ecology of juvenile steelhead trout Onchorhynchus mykiss and Chinook salmon O. tshawytscha in the context of habitat use and movement behaviour. The study area was a 14.8 km section of the Chehalis River, Washington, and is of particular interest due to recent proposals for both a flood retention dam and restoration actions in this watershed. Ten study reaches were paired in distance upstream and downstream from a central point where a passive integrated transponder antenna array was operated between late June and September 2014. Juvenile densities for each species were associated with reach‐scale habitat and temperature characteristics. Juvenile steelhead underwent upstream and downstream movements up to 7 km, although more fish from further away moved downstream than upstream. Juvenile steelhead repeated horizontal movements throughout the study period, but daily detections were not associated with temperature or flow. The majority (81%) of steelhead movements occurred between the hours of 04:00–07:00 and 18:00–21:00. Juvenile Chinook underwent a downstream migration that was nearly complete by the end of August. Most juvenile Chinook were detected just once and movements occurred on days with warmer stream temperature and higher flows. The majority of Chinook movements occurred at night. Although juvenile salmonids are often thought to have small home ranges during summer months, our results suggest that horizontal movements may be more prevalent than previously thought. Summer habitat should be defined by a network of suitable rearing reaches with connectivity available in both upstream and downstream directions.     

Polymorphic mountain whitefish (Prosopium williamsoni) in a coastal riverscape: size class assemblages,distribution, and habitat associations

We compared the assemblage structure, spatial distributions, and habitat associations of mountain whitefish (Prosopium williamsoni) morphotypes and size classes. We hypothesised that morphotypes would have different spatial distributions and would be associated with different habitat features based on feeding behaviour and diet. Spatially continuous sampling was conducted over a broad extent (29 km) in the Calawah River, WA (USA). Whitefish were enumerated via snorkelling in three size classes: small (10–29 cm), medium (30–49 cm), and large (≥50 cm). We identified morphotypes based on head and snout morphology: a pinocchio form that had an elongated snout and a normal form with a blunted snout. Large size classes of both morphotypes were distributed downstream of small and medium size classes, and normal whitefish were distributed downstream of pinocchio whitefish. Ordination of whitefish assemblages with nonmetric multidimensional scaling revealed that normal whitefish size classes were associated with higher gradient and depth, whereas pinocchio whitefish size classes were positively associated with pool area, distance upstream, and depth. Reach‐scale generalised additive models indicated that normal whitefish relative density was associated with larger substrate size in downstream reaches (R² = 0.64), and pinocchio whitefish were associated with greater stream depth in the reaches farther upstream (R² = 0.87). These results suggest broad‐scale spatial segregation (1–10 km), particularly between larger and more phenotypically extreme individuals. These results provide the first perspective on spatial distributions and habitat relationships of polymorphic mountain whitefish.