Demographic changes following mechanical removal of exotic brown trout in an Intermountain West (USA), high‐elevation stream

Exotic species present a great threat to native fish conservation; however, eradicating exotics is expensive and often impractical. Mechanical removal can be ineffective for eradication, but nonetheless may increase management effectiveness by identifying portions of a watershed that are strong sources of exotics. We used mechanical removal to understand processes driving exotic brown trout (Salmo trutta) populations in the Logan River, Utah. Our goals were to: (i) evaluate the demographic response of brown trout to mechanical removal, (ii) identify sources of brown trout recruitment at a watershed scale and (iii) evaluate whether mechanical removal can reduce brown trout densities. We removed brown trout from 2 km of the Logan River (4174 fish), and 5.6 km of Right Hand Fork (RHF, 15,245 fish), a low‐elevation tributary, using single‐pass electrofishing. We compared fish abundance and size distributions prior to, and after 2 years of mechanical removal. In the Logan River, immigration to the removal reach and high natural variability in fish abundances limited the response to mechanical removal. In contrast, mechanical removal in RHF resulted in a strong recruitment pulse, shifting the size distribution towards smaller fish. These results suggest that, before removal, density‐dependent mortality or emigration of juvenile fish stabilised adult populations and may have provided a source of juveniles to the main stem. Overall, in sites demonstrating strong density‐dependent population regulation, or near sources of exotics, short‐term mechanical removal has limited effects on brown trout populations but may help identify factors governing populations and inform large‐scale management of exotic species.     

Landscape features determine brown trout population structure and recruitment dynamics

Variation in brown trout (Salmo trutta L.) population recruitment and structure is related to migratory patterns, which should depend on ease of access to habitats providing increased opportunity for growth. We quantified the number of young of year (YOY) as a proportion of the total number of brown trout at 24 locations on 11 streams within the Taieri catchment, New Zealand, including back calculated growth rates and emergence dates from otoliths. Locations with high absolute and relative abundance of YOY fish were related to elevation and distance from the river mainstem (habitat used by migratory fish), fish density, and the interaction between invertebrate food biomass, distance and elevation. Hatch date and growth were not related to the proportion of YOY fish, though growth was negatively correlated to total fish density. We suggest landscape features play a large role in determining recruitment and population structure. Locations at lower elevations have a high YOY density, high competition and lower growth, likely prompting out‐migration. These conditions could be created by successful return migration and spawning of large fecund fish resulting in YOY densities exceeding the habitat carrying capacity. Environmental factors, such as food availability, also played a role in determining population structure. These results provide an example of how population structure and recruitment might be controlled by local conditions and access to high growth environments in wild populations of introduced brown trout across a catchment.     

Potential population and assemblage influences of non‐native trout on native nongame fish in Nebraska headwater streams

Non‐native trout are currently stocked to support recreational fisheries in headwater streams throughout Nebraska. The influence of non‐native trout introductions on native fish populations and their role in structuring fish assemblages in these systems is unknown. The objectives of this study were to determine (i) if the size structure or relative abundance of native fish differs in the presence and absence of non‐native trout, (ii) if native fish‐assemblage structure differs in the presence and absence of non‐native trout and (iii) if native fish‐assemblage structure differs across a gradient in abundances of non‐native trout. Longnose dace Rhinichthys cataractae were larger in the presence of brown trout Salmo trutta and smaller in the presence of rainbow trout Oncorhynchus mykiss compared to sites without trout. There was also a greater proportion of larger white suckers Catostomus commersonii in the presence of brown trout. Creek chub Semotilus atromaculatus and fathead minnow Pimephales promelas size structures were similar in the presence and absence of trout. Relative abundances of longnose dace, white sucker, creek chub and fathead minnow were similar in the presence and absence of trout, but there was greater distinction in native fish‐assemblage structure between sites with trout compared to sites without trout as trout abundances increased. These results suggest increased risk to native fish assemblages in sites with high abundances of trout. However, more research is needed to determine the role of non‐native trout in structuring native fish assemblages in streams, and the mechanisms through which introduced trout may influence native fish populations.     

Parallelism in thermal growth response in otoliths and scales of brown trout (Salmo trutta L.) from alpine lakes independent of genetic background

Low density in natural populations of salmonids has predominantly been managed by stocking of non‐native conspecifics. Due partly to domestication, introduced non‐native fish may be maladapted under natural conditions. Interbreeding between introduced and wild individuals may therefore impair local adaptation and potentially population viability. Brown trout (Salmo trutta L.) from three headwaters (with stocked fish) and three interconnected lakes (with native fish) on the Hardangervidda mountain plateau, southern Norway, were tested for differences in thermal effects on scale and otolith growth. Otolith and scale annuli widths from immature brown trout showed positive correlation with mean annual summer temperature for all six sampled populations. In mature individuals, a similar positive thermal correlation was evident for the otoliths only. Interannuli width measurements from scales indicate a halt in somatic growth for brown trout in this alpine environment when reaching ages between 7 and 9 winters, coinciding with age at maturity. Our study indicates that otolith growth follows summer temperature even when individuals do not respond with somatic growth in these populations and that introduced brown trout and introgressed populations have similar thermal growth responses. Due to the continued otolith growth after stagnation in somatic growth and the impact of fluctuations in summer temperature, the utilisation of otolith annuli widths for back calculation of length at age should be treated with caution.     

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.     

Biotic and abiotic regulation of a low‐density stream‐dwelling brown trout (Salmo trutta L.) population: effects on juvenile survival and growth

The effects of biotic (density‐dependent) and environmental (flow and temperature) factors on the apparent survival, mean length and size variation of a low‐density brown trout population in the juvenile stage, that is, from their first summer (0+) to the end of the second year (1+), were determined. Apparent survival was negatively related to the age class density during the three periods (first summer, first winter and second summer). A significant interaction between the mean flow and 0+ density highlighted a gradient towards strong density dependence acting on fish loss (i.e., mortality or migration) with decreasing summer flow. Conversely, no density dependence was reported at higher mean flows. The mean length was determined by density‐dependent and density‐independent (temperature and flow) factors throughout the study period. The negative relationship between fish length and intracohort density was highly significant during the three periods. The yearling (1+) density was negatively related to 0+ fish length measured after the first summer, suggesting intercohort effects. A positive effect of temperature on fish length was observed. Mean length after the summer seasons (0+ and 1+ fish) was also positively related to mean flow. Fish size variation around the mean measured with the coefficient of variation (CV) increased with increasing 0+ densities, both at the end of the first summer and the first winter. Results suggested that density‐dependent and density‐independent factors acted jointly on apparent survival and growth with a predominance of biotic processes. We discussed the potential implications of density‐dependent regulations on growth and survival for population resilience after catastrophic events.     

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.     

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.     

Addition of structural complexity – contrasting effect on juvenile brown trout in a natural stream

In a field experiment, we examined the effects of structural complexity in the form of added artificial plastic plants and shredded plastic bags on growth and abundance of juvenile brown trout (Salmo trutta). Just after emergence, the added complexity had a positive effect on the density, biomass and condition factor of young‐of‐the‐year (0+) brown trout. This difference in density was not present six weeks later. In contrast, both young‐of‐the‐year and older brown trout generally tended to be larger in the simple habitat. Hence, our data suggest that increased complexity initially is beneficial for young‐of‐the‐year individuals probably due to lower risk of predation and increased densities of prey. However, as density increases in the complex environment, it may induce negative density‐dependent effects, here reflected in smaller sized fish in the complex environment. This might force fish to redistribute to habitats with lower densities of conspecifics as they grow larger. We propose that habitat complexity can increase survival of yearlings in early phases and thereby also affect the overall population structure of brown trout in natural streams.     

A conceptual model for examining density dependence in the growth of stream trout

Determination of the occurrence and importance of densitydependent responses is central to understanding stream trout population dynamics. I propose a conceptual model of growth, based on a distribution of feeding site quality, that considers the effects of density on growth. The site quality model assumes that trout select the best feeding sites available and, as the number of trout increases, they will be forced to use less energetically profitable sites, resulting in decreased growth, but also an increase in variance of size or growth. Results from a 3-year study of a brown trout (Salmo trutta) population show that growth to age 1 was significantly reduced for the 1981 year class, which was about 4 times more dense than other year classes. The reduction in growth was not due to a decrease in the growth of all trout but rather an increase in the number of slower growing fish. These results are consistent with my site quality distribution model and suggest that individual fish growth, the distribution of growth, and the variance of growth rates should be considered in addition to mean size and growth.     

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.     

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.     

Increasing connectivity of Great Lakes tributaries: Interspecific and intraspecific effects on resident brook trout and brown trout populations

Understanding resident fish population responses to restored connectivity would enhance decision-making on dam removal and fish passage. Since such evaluations are limited in the Great Lakes region of North America, we compared abundance, survival, and growth of resident brook trout and brown trout between sets of Michigan streams where populations were or were not interacting with salmonid species that might be present if connectivity existed. We analysed data from 34 electrofishing index sites to compare resident trout populations between streams without versus with Great Lakes access (and migratory Pacific salmonids), and brook trout populations in Great Lakes inaccessible (land-locked) streams where brown trout were present versus absent. Great Lakes accessibility effects on fish density became increasingly positive for older age groups of brown trout while generally negative for all age classes of brook trout. Brown trout had consistently negative effects on brook trout density in land-locked streams. Increased connectivity had significant effects on annual survival for only one of seven trout age classes modelled, while intraspecific density-dependent effects on survival were significant in six models. Significant intraspecific effects on resident trout growth occurred for seven of eleven age classes examined. Negative interspecific effects of Great Lakes access on resident trout growth were most noticeable for age-0 and age-1 resident trout, age classes that likely compete with juvenile Pacific salmonids. Our findings provide a more robust understanding of how Great Lakes connectivity affects resident trout populations, highlighting negative influences of brown trout on brook trout and intraspecific density-dependent effects.     

Recruitment of the threatened mussel Margaritifera margaritifera in relation to mussel population size,mussel density and host density

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Driving factors of synchronous dynamics in brown trout populations at the rear edge of their native distribution

The classical debate about the roles of endogenous factors and external drivers in regulating populations continues to be a very active area of scientific research, and these roles are even more complex in populations located in geographically marginal areas. We tested three hypotheses on two separate brown trout populations inhabiting the southernmost basins of the Iberian Peninsula, with the aims of determining the relationships among (a) rainfall and air temperature for each season preceding fish sampling and the observed age group densities; (b) environmental synchrony and density synchrony for all possible pairs of sites; and (c) physical habitat similarity and the similarity of the density response to climatic drivers among pairs of sites. For this purpose, demographic series (98 surveys at 14 sampling sites), climatic conditions (rainfall and air temperature) and site habitat characteristics (physiogeographic and quality index values) were analysed between 2006 and 2014. The results show how the synchrony in the study populations is the result of a Moran effect directed mainly by winter precipitation and conditioned by the habitat similarity (highlighting the altitude, the distance to the upper limit and IHF index, along with the Euclidian distance among pairs of sites of the whole set of habitat variables). Preserving the heterogeneity of these habitat characteristics will increase the resilience of trout populations that inhabit this rear edge of distribution. The current context of global climate warming predicts extremely drastic changes in the dynamics of external drivers that regulate the trout populations in the study region.     

Dispersion of stocked brown trout, Salmo trutta L., and landlocked salmon, Salmo salar L., from stocking sites in Lake Oulujärvi, Finland

Movement and recaptures of two hatchery-reared brown trout, Salmo trutta L., stocks and landlocked salmon, Salmo salar L., released at different sites in regulated Lake Oulujärvi, were studied in relation to release site. Five groups of fish from each stock were released in approximately equal numbers. Most of the fish released in June and July were recaptured within 3 months, whereas the majority of the fish released in early winter (October and November) were caught the following spring, about 7–9 months after stocking. The release site had a significant effect on recapture rate. The results showed that fishing restrictions targeted mainly at gill net fishing are needed to preserve the stocked fish from overfishing. Significantly fewer recaptures were observed from the landlocked salmon stocking compared with brown trout. The recaptures from the landlocked salmon stocking indicated more active movement and less clumping compared with the two brown trout stocks.     

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.     

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

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

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.     

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

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