The effect of competition among three salmonids on dominance and growth during the juvenile life stage

Although non‐native species can sometimes threaten the value of ecosystem services, their presence can contribute to the benefits derived from the environment. In the Great Lakes, non‐native brown trout (Salmo trutta) and rainbow trout (Oncorhynchus mykiss) support substantial recreational fisheries. With current efforts underway to restore once‐native Atlantic salmon (Salmo salar) to Lake Ontario, there is some concern that Atlantic salmon will impede non‐native contributions to the recreational fishery because Atlantic salmon exhibit niche overlap with brown trout and rainbow trout, particularly during the juvenile life stage. We therefore examined competition and growth of juvenile Atlantic salmon, brown trout and rainbow trout in semi‐natural streams. We found that brown trout were the most dominant and had the greatest growth rate regardless of what other species were present. Rainbow trout were more dominant than Atlantic salmon and consumed the most food of the three species. However, in the presence of brown trout, rainbow trout fed less frequently and exhibited negative growth as compared to when the rainbow trout were present with only Atlantic salmon. These data suggest that, outside of density‐dependent effects, Atlantic salmon will not impact stream production of brown trout and rainbow trout.     

Food habits of introduced brown trout and native masu salmon are influenced by seasonal and locational prey availability

A knowledge of food habits is important for evaluating interspecific competition and predation between sympatric species. Data on food availability should be combined with data on food habits in this type of survey. Although food availability differs between habitats or seasons, these differences had never been considered in previous studies. We conducted year-round field surveys throughout a stream to compare the food habits of an introduced salmonid, brown trout Salmo trutta, and a native salmonid, masu salmon Oncorhynchus masou. Masu salmon did not constitute a large proportion of the diet of brown trout and vice versa. Thus, predation will likely not affect the population level of either species. The dietary overlap between brown trout and masu salmon varied depending on the presence of Gammaridae and terrestrial invertebrates; i.e., the intensity of interspecific competition for food resources may differ according to food conditions.     

Tagging effects on three non-native fish species in England (Lepomis gibbosus, Pseudorasbora parva, Sander lucioperca) and of native Salmo trutta

Abstract –  To address the dearth of information on tagging effects and long-term survivorship of tagged fish in native and introduced species, laboratory and field investigations were undertaken on three non-native fish species (pumpkinseed Lepomis gibbosus ; topmouth gudgeon Pseudorasbora parva ; pikeperch Sander lucioperca ) tagged with coded-wire (CW), passive integrated transponder (PIT), radio (RT) telemetry and/or acoustic tags (AT), with survivorship of native brown trout ( Salmo trutta ) examined in the field. Laboratory results revealed high survivorship following tag attachment/insertion and resumption of feeding within 24–48 h of tagging (all mortalities could be attributed to an unrelated outbreak of fungal infection), with retention rates being high in both pumpkinseed and pikeperch but low in topmouth gudgeon (excluded from field studies). In the field, short-term post-operation survival was high in pikeperch, pumpkinseed and brown trout. In pumpkinseed and trout, 100% of RT fish survived a 24–30 day tracking study, with 60% and 80%, respectively, recaptured alive at least 3 months post-tagging. Of PIT tagged pumpkinseed, 44% were recaptured (after 6–18 months), with small-sized, CW-tagged fish (0.38 g weight) captured up to 1 year after tagging. In pikeperch, all AT fish except one (the smallest specimen) survived their full expected tracking period (i.e. tag life) – the single lost specimen survived at least half of its expected tracking period (i.e. 6 month battery life). Overall, the tagging methods used were highly effective in pumpkinseed and pikeperch, showing good retention and survival, but PIT tagging of topmouth gudgeon was plagued by low survivorship and tag rejection.     

Stocked rainbow trout (Oncorhynchus mykiss) affect space use by Warpaint Shiners (Luxilus coccogenis)

Due to widespread stocking, Rainbow Trout (Oncorhynchus mykiss Walbaum) are perhaps the most widely distributed invasive species in the world. Nonetheless, little is known about the effects of stocked Rainbow Trout on native non‐game species. We conducted experiments in an artificial stream to assess the effects of hatchery Rainbow Trout on home range and behaviour of Warpaint Shiners (Luxilus coccogenis Cope), a common minnow frequently found in stocked Southern Appalachian streams. We used the LoCoH algorithm to generate polygons describing the home ranges used by Warpaint Shiners. When a stocked trout was present Warpaint Shiners: (a) increased home range size by 57%, (b) were displaced into higher velocity mesohabitats, and (c) reduced mean overlap between the home ranges of individual warpaint shiners. Rainbow Trout did not significantly affect the edge/area ratio of Warpaint Shiner home ranges. Warpaint Shiner density (two and five fish treatments) did not significantly affect any response variable. Displacement from preferred microhabitats and increases in home range size likely result in increased energy expenditure and exposure to potential predators (i.e., decreased individual fitness) of Warpaint Shiners when stocked trout are present.     

Foraging mode variation in three stream‐dwelling salmonid fishes

Despite long‐standing interest in foraging modes as an important element of animal space use, few studies document and compare individual foraging mode differences among species and ecological conditions in the wild. We observed and compared foraging modes of 61 wild Arctic charr, Salvelinus alpinus, 42 brown trout, Salmo trutta, and 50 Atlantic salmon, Salmo salar, in their first growing season over a range of habitats in 10 Icelandic streams. We found that although stream salmonids typically sit‐and‐wait to ambush prey from short distances, Arctic charr were more mobile during prey search and prior to prey attack than Atlantic salmon, whereas brown trout were intermediate. In all three species, individuals that were mobile during search were more likely to be moving when initiating attacks on prey, although the strength and the slope of this relationship differed among species. Arctic charr also differed from salmon and trout as more mobile individuals travelled longer distances during prey pursuits. Finally, coupled with published data from the literature, salmonid foraging mobility (both during search and prior to attack) clearly decreased from still water habitats (e.g., brook charr), to slow‐running waters (e.g., Arctic charr) to fast‐running waters (e.g., Atlantic salmon). Hence, our study suggests that foraging mode of young salmonids can vary distinctly among related species and furthers our understanding of the behavioural mechanisms shaping the geographical distribution of wild salmonids.     

Effect of predation and habitat quality on growth and reproduction of a stream fish

Billman EJ, Tjarks BJ, Belk MC. Effect of predation and habitat quality on growth and reproduction of a stream fish.
Ecology of Freshwater Fish 2011: 20: 102–113. © 2010 John Wiley & Sons A/S Abstract – Anthropogenic disturbances are rarely independent, requiring native fishes to respond to multiple factors to persist in changing environments. We examined the interaction of predation environment (presence of introduced brown trout, Salmo trutta) and habitat quality on growth and reproduction of southern leatherside chub, Lepidomeda aliciae, a small‐bodied stream fish native to central Utah, USA. Southern leatherside chub were sampled from four streams representing a complete two‐factor cross of predation environment and habitat quality. Growth was estimated using increment analysis of annuli on otoliths, and reproductive traits were measured for both sexes. Southern leatherside chub growth was greater in high‐quality than in low‐quality habitats, and greater in predator than in nonpredator environments. However, fish exhibited a greater growth response to presence of brown trout in low‐quality habitats. Southern leatherside chub growth followed predictions of plastic responses to resource availability based on habitat quality and predation environment (lethal vs. nonlethal effects). Reproductive allocation (gonad wet mass) was significantly greater in low‐quality versus high‐quality habitats, but was unaffected by predation environment. Other female life‐history traits were affected either by both effects or their interaction. Reproductive responses to habitat quality and predation environment were consistent with predictions based on differential mortality. Southern leatherside chub growth and reproduction responded differently to the combination of habitat quality and predation environment, thus demonstrating the importance of assessing interacting effects of anthropogenic disturbances to more fully comprehend impacts on native species and to appropriately manage, recover and restore these species and their habitats.     

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.     

Competitive interactions among multiple non‐native salmonids and two populations of Atlantic salmon

Competitive interactions with non‐native species can have negative impacts on the conservation of native species, resulting in chronic stress and reduced survival. Here, juvenile Atlantic salmon (Salmo salar) from two allopatric populations (Sebago and LaHave) that are being used for reintroduction into Lake Ontario were placed into semi‐natural stream tanks with four non‐native salmonid competitors that are established in Ontario streams: brown trout (S. trutta), rainbow trout (Oncorhynchus mykiss), Chinook salmon (O. tshawytscha) and coho salmon (O. kisutch). Brown trout and rainbow trout reduced the survival and fitness‐related traits of Atlantic salmon, whereas Chinook salmon and coho salmon had no impact on these traits. These data support theories on ecological niche overlap and link differences in observed aggression levels with competitive outcomes. Measurements of circulating hormones indicated that the Atlantic salmon were not chronically stressed nor had a change in social status at the 10‐month time point in the semi‐natural stream tanks. Additionally, the Sebago population was better able to coexist with the non‐native salmonids than the LaHave population. Certain populations of Atlantic salmon may thus be more suitable for some environments of the juvenile stream phase for the reintroduction into Lake Ontario.     

Competitive interactions between native and exotic salmonids: a combined field and laboratory demonstration

Abstract –  We studied the impact of two exotic salmonid species (brook trout, Salvelinus fontinalis and rainbow trout, Oncorhynchus mykiss ) on native brown trout ( Salmo trutta fario ) habitat, growth and survival. Habitat selection and vertical distribution between young-of-the-year of the three species were examined in a stream aquarium under different sympatric and allopatric combinations. In addition, similar species combinations were introduced in a Pyrenean mountain stream (southwest France) in order to extend laboratory results to growth and apparent survival. Both laboratory and field results indicated that rainbow trout significantly affected native brown trout habitat selection and apparent survival. On the contrary, brown trout habitat, growth and apparent survival were hardly affected by brook trout. These results support the idea that rainbow trout negatively influence native brown trout, and that competition could influence the outcome of fish biological invasions in freshwater ecosystems.     

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.     

Projected impacts of climate change on stream salmonids with implications for resilience‐based management

The sustainability of freshwater fisheries is increasingly affected by climate warming, habitat alteration, invasive species and other drivers of global change. The State of Michigan, USA, contains ecologically, socioeconomically valuable coldwater stream salmonid fisheries that are highly susceptible to these ecological alterations. Thus, there is a need for future management approaches that promote resilient stream ecosystems that absorb change amidst disturbances. Fisheries professionals in Michigan are responding to this need by designing a comprehensive management plan for stream brook charr (Salvelinus fontinalis), brown trout (Salmo trutta) and rainbow trout (Oncorhynchus mykiss) populations. To assist in developing such a plan, we used stream‐specific regression models to forecast thermal habitat suitability in streams throughout Michigan from 2006 to 2056 under different predicted climate change scenarios. As baseflow index (i.e., relative groundwater input) increased, stream thermal sensitivity (i.e., relative susceptibility to temperature change) decreased. Thus, the magnitude of temperature warming and frequency of thermal habitat degradation were lowest in streams with the highest baseflow indices. Thermal habitats were most suitable in rainbow trout streams as this species has a wider temperature range for growth (12.0–22.5 °C) compared to brook charr (11.0–20.5 °C) and brown trout (12.0–20.0 °C). Our study promotes resilience‐based salmonid management by providing a methodology for stream temperature and thermal habitat suitability prediction. Fisheries professionals can use this approach to protect coldwater habitats and drivers of stream cooling and ultimately conserve resilient salmonid populations amidst global change.     

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.     

Resource use of native and stocked brown trout Salmo trutta L., in a subarctic lake

Habitat use, food composition and growth of stocked and native brown trout, Salmo trutta L., were studied in the subarctic Lake Muddusjärvi in northern Finland. Stocked brown trout and native brown trout preferred littoral and pelagic areas. Trout were stocked in October. In June stocked trout fed primarily on invertebrates while native fish were piscivorous. From July onwards the composition of the diet of both stocked and native trout was similar and consisted almost entirely of small‐sized whitefish. Brown trout were already piscivorous at a length of about 20 cm. The mean length of prey consumed was about 12 cm. Mean length‐at‐age was similar from the second year in the lake despite of the larger size of stocked fish during the first year in the lake.     

Swimming performance of brown trout and grayling show species‐specific responses to changes in temperature

Fishways have historically been constructed to restore and preserve the ecological connectivity for fish in fragmented rivers. However, the fishways are often selective on species due to different size and swimming capacity. As the proportion of dammed rivers is still increasing, there is a growing need for more information on wild fish and their migration potential. In this study, we compared the swimming capacity of wild caught brown trout (Salmo trutta) and European grayling (Thymallus thymallus) until the fish were exhausted in a critical swimming speed (U_crit) test, under three different naturally occurring stream temperatures in Norway: 1.7, 5.5 and 10°C. The results indicated that trout swim better at the warmer temperatures than at colder temperatures. The grayling showed consistent swimming patterns with little variation across all tested temperatures. The results therefore signify the need to have operational fishways already early in the spring when the grayling migration starts and highlight the need for more studies on fish migration abilities across a wider range of species and seasons.     

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.     

Different longitudinal distribution patterns of native white-spotted charr and non-native brown trout in Monbetsu stream, Hokkaido, northern Japan

Abstract  This study focused on longitudinal distribution patterns of native white-spotted charr and non-native brown trout in a mountainous stream in Hokkaido, Japan. Brown trout ratio, which is the proportion of brown trout in the catch of salmonids, was decreasing from downstream to upstream. Brown trout ratio correlated negatively with water temperature. Thus, our results suggested that temperature may influence the possible competition between native white-spotted charr and non-native brown trout.     

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.     

Habitat niche separation of the nonnative rainbow trout and native masu salmon in the Atsuta River,Hokkaido, Japan

The mechanisms by which nonnative species establish populations can be classified into two broad categories: they usurp the niches of native species through interspecific competition, or they avoid this intense interspecific competition by making use of minimal niche overlap with the native species. In this study, we considered how a nonnative salmonid species, the rainbow trout Oncorhynchus mykiss, established a population in the presence of the native salmonid species, the masu salmon O. masou, in Hokkaido, Japan. Circumstantial field evidence shows that the masu salmon exceeds the rainbow trout in abundance and suggests that these species use different types of cover habitat (rainbow trout abundance increases with increasing abundance of large woody debris aggregates, whereas masu salmon abundance increases with increasing abundance of undercut banks). These results imply that the rainbow trout established a population due to minimal niche overlap with the masu salmon, and not by competitive exclusion of the native species.     

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.     

Competitive effects between rainbow trout and Atlantic salmon in natural and artificial streams

Competition with non‐native species may impede the restoration of native species, but differences in competitive abilities among intraspecific native populations may make some populations more suitable for reintroduction than others. Here, juvenile Atlantic salmon (Salmo salar) from two allopatric populations (LaHave and Sebago) being used for reintroduction into Lake Ontario were placed into two natural stream sites differing in the presence of ecologically similar rainbow trout (Oncorhynchus mykiss). We assessed the effects of competition in the natural streams on fitness‐related traits and habitat use of the Atlantic salmon. We then compared these effects to those observed in artificial streams from a previous study. Atlantic salmon in natural streams had reduced fitness‐related traits that were associated with suboptimal microhabitats in the presence of rainbow trout, but utilised optimal microhabitats in their absence. In the presence of rainbow trout, the two Atlantic salmon populations exhibited comparable recapture proportions to each other, but the individuals from the Sebago population had better performance (body size and condition) than those from the LaHave population. Responses of both Atlantic salmon populations to competition with rainbow trout were generally similar in both direction and magnitude when compared to results from the artificial stream study. The combined results suggest that native species restoration efforts should be focused on candidate populations that are ecologically suitable to reintroduction environments, as well as on suitable habitats that do not contain exotic competitors. Moreover, this study highlights the value of controlled experiments in artificial environments for predicting fitness‐related performance in natural environments.