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.     

Invasions of rainbow trout and brown trout in Japan: A comparison of invasiveness and impact on native species

Many species of salmonids have been stocked into waters outside of their native range. The invasiveness and impact of these species on native species varies depending on their biological traits, and on environmental conditions, such as climate. In Japan, rainbow trout and brown trout, both listed in 100 of the world's worst invasive alien species by the International Union for Conservation of Nature, occur as non-native species. The invasiveness of these two species is thought to be related to seasonal flooding, given flood waters can physically damage fry and prevent population establishment. Rainbow trout have successfully invaded waters in Hokkaido, northern Japan, where the likelihood of flooding is low between June and July, when their fry emerge, but successful invasions are rare in regions south of Hokkaido. Brown trout, however, have successfully invaded waters not only in Hokkaido, but also other regions. Since brown trout have a similar life history to the native white-spotted charr and masu salmon, with fry emerging before the flood season, they are more suited to the Japanese climate than Rainbow trout. Rainbow and brown trout interact with native species in various ways, but a common outcome of these interactions is the displacement of native charr species. Legal regulations of non-native salmonids should be based on understandings of the ecological traits of each invasive species and regional impacts on native species. Given the ongoing nature of climate change, the nature and extent of the effects of rainbow and brown trout on native species might also change.     

Resource partitioning and spatial segregation in native and stocked brown trout, Salmo trutta L., and Arctic charr, Salvelinus alpinus (L.), in a hydroelectric reservoir

Abstract. Habitat use, food and spatial segregation in native and stocked brown trout, Salmo trutta L., and Arctic charr, Salvelinus alpinus (L.), were studied during summer 1989 and 1990 in the hydroelectric reservoir Lake Tunhovdfjorden. There was no difference in habitat use and feeding habits between wild and stocked brown trout. In epibenthic areas brown trout lived chiefly down to 2 Secchi disc units, whereas Arctic charr were most abundant between 1 and 4 Secchi disc units. In pelagic areas the catches were low for both species, and they were chiefly confined to surface waters down to 1 Secchi disc unit. The food segregation between brown trout and Arctic charr was almost complete. Both pelagic and epibenthic Arctic charr fed mainly on cladocerans ( Bosmina longispina and Daphnia galeata ), whereas surface insects of terrestrial origin and Arctic charr were the dominant food items for brown trout. Pelagic Arctic charr were significantly older, larger and more homogeneous in size than epibenthic charr. During calm weather schools of Arctic charr were observed cruising with the dorsal fin above the surface.     

Long-term variation in piscivory in a brown trout population: effect of changes in available prey organisms

Abstract – The piscivorous behaviour in a brown trout ( Salmo trutta L.) population was studied in four discrete periods over seven decades (1917–94) in the hydroelectric reservoir Tunhovdfjord in Norway established in 1919. Piscivorous brown trout were extremely scarce prior to the introduction of two fish species Arctic charr ( Salvelinus alpinus L.) and European minnow ( Phoxinus phoxinus L.) in the 1920s. Brown trout started eating minnow at 17 cm and Arctic charr at 22 cm of length. In the 1950s, the brown trout predated extensively (60% of analysed trout) on Arctic charr and minnow. During the next four decades, the incidence of piscivorous brown trout declined to 15%, whereas the frequency of brown trout eating Arctic charr remained constant at 10%. The growth pattern, expressed as back-calculated length, demonstrated similarity in three periods (1920s, 1960s and 1990s) and improved growth in the 1950s. The improvement was addressed the impoundment of a reservoir upstream. We did not find any marked change in growth rate due to piscivority, but coefficient of variance of back-calculated lengths indicated significant variation in individual growth in age group ≥6 years from 1950 onwards. We accredit this variation to the rise of piscivorous brown trout.     

The relationship between the snowmelt flood and the establishment of non‐native brown trout (Salmo trutta) in streams of the Chitose River,Hokkaido, northern Japan

Flow regime is one of the major determinants of establishment success for non‐native aquatic organisms. Here, we examine the influence of flow variability associated with snowmelt flood on the establishment success of non‐native brown trout in 10 streams in northern Japan. We regarded the presence of Age‐0 brown trout as the index of the successful establishment. The emergence of Age‐0 brown trout in our study region begins in May, a time that overlaps with the occurrence of snowmelt flood. The presence of Age‐0 brown trout was negatively associated with flow variability, and it was also negatively associated with summer water temperature. Our results indicate that the non‐native brown trout tends to establish in the streams with smaller snowmelt floods and lower summer water temperatures. Brown trout is an invasive, non‐native species that is problematic all over the world, and effective management strategies for preventing their further expansion are urgently needed. This study suggests that river managers should recognise that stable streams such as spring‐fed streams (i.e., low flow and summer water temperature) and flow‐regulated streams, have a higher potential risk of brown trout invasion.     

Hybrids as potential mediators spreading non-native genes: Comparison of survival,growth, and movement among native,introduced and their hybrid salmonids

Comparing multiple fitness components and potential movement of wild hybrids with their parental species is necessary to fully understand the consequences of human-mediated introgression, but studies tracking both parental species and their hybrids at the individual-level are limited. Here, we compared growth, survival and movement of sympatric introduced brook trout (BT: Salvelinus fontinalis) and native white-spotted charr (WSC: S. leucomaenis) with their hybrids (HYB) in a northern Japanese stream, using mark-recapture data (1,087 marked individuals) collected over 4 years (2013–2016). The mark-recapture data with a single cohort showed that HYB had a comparable or even higher growth rate to BT and WSC. In addition, there is no evidence that hybrid survival was lower than both parental species throughout the entire study period. Furthermore, HYB showed high mobility equivalent to WSC, while BT showed the lowest mobility. Although our previous studies have documented the reduction of BT distribution and lowered reproductive success of HYB, non-native genes can pose a threat to native WSC via relatively high survival, growth and/or mobility of HYB.     

Brown trout (Salmo trutta L.) and Arctic charr (Salvelinus alpinus (L.)) as predators on three sympatric whitefish (Coregonus lavaretus (L.)) forms in the subarctic Lake Muddusjärvi

Abstract  – Brown trout ( Salmo trutta L.) and Arctic charr ( Salvelinus alpinus (L.)) use whitefish ( Coregonus lavaretus (L.)) as their main prey in the subarctic Lake Muddusjärvi. Brown trout dwelled in littoral and pelagic habitat, whereas Arctic charr lived only in epibenthic habitat. Both species shifted to whitefish predation at a length of 20–30 cm. At this size, brown trout fed on larger whitefish than Arctic charr. Whitefish occur in three sympatric forms, differing in their habitat, ecology and morphology. Both the predators preyed primarily upon the small-sized, densely rakered whitefish form (DR), which was the most numerous whitefish form in the lake. DR used both epibenthic and pelagic habitat, whereas two sparsely rakered whitefish forms dwelled (LSR and SSR) only in epibenthic habitat: LSR in littoral and SSR in profundal areas. Sparsely rakered whitefish forms had minor importance in predator diet.     

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.     

Threats to at-risk salmonids of the Canadian Rocky Mountain region

Trout and charr, members of the salmonid family, have high conservation value but are also susceptible to anthropogenic threats in part due to the specificity of their habitat requirements. Understanding historical and future threats facing these species is necessary to promote their recovery. Of freshwater trout and charr in the Canadian Rocky Mountain region, westslope cutthroat trout (Oncorhynchus clarkii lewisi), bull trout (Salvelinus confluentus; a charr species) and Athabasca rainbow trout (Oncorhynchus mykiss) are of conservation concern. And indeed, range contractions and declining populations are evident throughout much of their ranges. Range contraction was most evident in the southern Alberta designatable unit (DU) of westslope cutthroat trout. Diminished populations were also evident in the downstream watersheds of the Alberta bull trout range, and throughout the Athabasca rainbow trout range. We assessed historical and future threats to evaluate the relative importance of individual threats to each DU and compare their impact among species. Individual threats fall into the broad categories of angling, non-native species and genes, habitat loss and alteration, and climate change. Severity of each threat varies by DU and reflects the interaction between species’ biology and the location of the DU. Severity of threats facing each DU has changed over time, reflecting extirpation of native populations, changes in management and industry best practices, expansion of non-native species and progressing climate change. The overall threat impact for each DU indicates a high probability of substantial and continuing declines and calls for immediate action.     

Comparative life-history characteristics of native and hatchery-reared brown trout, Salmo trutta L., in a sub-Alpine reservoir

Wild and non-native hatchery-reared brown trout, Salmo trutta L., released when 2 summers old, were caught in the littoral habitat of Vinstervatna Reservoir, southern Norway. Hatchery-reared brown trout grew more slowly and had a smaller asymptotic length (293 ± 71 mm CL) than native fish (391 ± 56 mm CL). Hatchery-reared brown trout also exhibited significantly shorter life spans than native fish. This category consisted mainly of individuals aged 2+ and 3+ years, and only 1.5% of the specimens were aged ≥5 years. The ages of the native fish in the sample were between 2 and 8 years, and the most abundant age groups of trout were 4+ and 5+ years. It is suggested that the differences in life-history characteristics are related to adaptations by the native trout to the local environmental conditions. In this reservoir, which has a limited food supply as a result of water level fluctuations and a high level of inter- and intraspecific competition, environmental effects might be significant.     

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.     

Habitat use, size and age structure in sympatric brown trout (Salmo trutta) and Arctic charr (Salvelinus alpinus) stocks: resistance of populations to change following harvest

Abstract– Habitat use and population dynamics in brown trout Salmo trutta and Arctic charr Salvelinus alpinus were studied in an oligotrophic lake over a period of 10 years. Previous studies showed that the species segregated by habitat during summer. While brown trout occupied the surface water down to a depth of 10 m, Arctic charr were found deeper with a maximum occurrence at depth 10–15 m. Following the removal of a large number of intermediate sized fish in 1988–89, habitat segregation between the species broke down and Arctic charr were found in upper waters, while brown trout descended to deeper waters. The following year, both species were most frequently found in surface waters at depths of 0–5 m. During the last four years, the species reestablished their original habitat segregation despite another removal experiment of intermediate-sized fish in 1992–1994. The removal of fish resulted in an increased proportion of large (≥ 25 cm) fish in both species. Furthermore, the charr stock responded by reduced abundance and increased size-at-age. The results revealed plasticity and strong resistance to harvest populations of brown trout and Arctic charr. This is probably due to internal mechanisms of intraspecific competition within each population, which result in differential mortality among size classes.     

Effects of stocking of piscivorous brown trout, Salmo trutta L., on stunted Arctic charr, Salvelinus alpinus (L.)

To study the effects on a stunted freshwater population of Arctic charr, Salvelinus alpinus (L.), two groups of large (26–45 cm) individually tagged brown trout, Salmo trutta L., were released and recaptured with gillnets after 1, 7, 11 and 63 weeks. One group of trout was trained on a fish diet before release, and the other, reared on commercial dry pellets, served as a control. Specific growth rates in both groups were negative 1 week after release and approached zero after 63 weeks. Condition factor and internal fat content decreased during the experiment. Although only 11% of the trout stomachs examined contained fish prey, charr represented 79% of the total stomach weight content. Gillnet samples of charr before and 63 weeks after the release of trout indicated a decreasing population size of charr. Individual growth and mean length of charr increased after release of trout, especially for charr at age 4 years. After the release of trout, 35% of the charr were longer than 20 cm as compared with 6% before the release.     

Long-term variation in the population structure of Arctic charr, Salvelinus alpinus, and brown trout, Salmo trutta

The effects of induced water level fluctuations and introduction of the mysid Mysis relicta Lovén on population structure of brown trout, Salmo trutta L., and Arctic charr, Salvelinus alpinus (L.), were studied during 1953–1995 in Limingen hydroelectric reservoir, Norway. The main response was a marked reduction in catch‐per‐unit‐effort (CPUE) for trout and charr, probably caused by reduced recruitment following increased variation in water level. For both species, mean length decreased until 1967 and increased thereafter, whereas mean mass‐at‐length increased for the whole period. Both length and mass‐at‐length were negatively correlated with CPUE. The increases in mean length and mass‐at‐length were probably because of reduced competition following the reduced recruitment. Mysis relicta has become an important food item for charr but not for brown trout, but the increases in mean length and mass‐at‐length of charr started prior to the appearance of M. relicta in the charr diet.     

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.     

Hatch timing of two subarctic salmonids in a stream network estimated by otolith increments

Hatch timing in autumn-spawning stream salmonids is poorly understood in the subarctic region because snow cover prevents direct sampling of cryptic early life stages. Otolith micro-increment analysis was used to infer hatch dates of white-spotted charr Salvelinus leucomaenis (Pallas) and masu salmon Oncorhynchus masou (Brevoort) in a mainstem-tributary network in northern Japan. Accuracy and precision were validated by ageing hatchery individuals with known hatch date ranges. In July 2018, 93 wild young-of-the-year white-spotted charr and 81 masu salmon were collected and aged. Masu salmon hatched, on average, 24 days earlier (mean = February 8) than white-spotted charr (March 4), and hatch dates spanned a minimum of 2 months for each species. In masu salmon, hatch dates of individuals collected in the mainstem were nearly 3 weeks earlier than those in a tributary. This study provided knowledge on intra- and inter-specific variation in hatch timing of native salmonids in a subarctic stream network.     

Recovery of white-spotted charr Salvelinus leucomaenis following the removal of stocked red-spotted masu salmon Oncorhynchus masou ishikawae in a small headwater tributary of Lake Biwa,central Japan

The possible recovery of a white-spotted charr population in a small tributary of a river to Lake Biwa, following removal of previously stocked red-spotted masu salmon, was investigated by electrofishing. We captured 30 red-spotted masu salmon and two hybrids of these two species in 2014. Prior to the stocking of red-spotted masu salmon, the estimated number of white-spotted charr?≥?100 mm standard length (SL) in the tributary was more than 200 individuals in 2005, which had reduced to fewer than 30 individuals by 2014 when 30 red-spotted masu salmon plus two charr/masu salmon hybrids were captured. However, no red-spotted masu salmon were captured from 2015 to 2017, indicating the success of a red-spotted masu salmon removal program. The estimated number of white-spotted charr?≥?100 mm SL ranged from 25 to 91 individuals between 2015 and 2017, implying a great reduction of red-spotted masu. The estimated population size of the former increased further to 171–221 individuals in 2021, comparable to 2005. These results suggest that the white-spotted charr population declined due to stocking of red-spotted masu salmon, and recovered following removal of the latter.

     

Trophic interactions between introduced lake trout (Salvelinus namaycush) and native Arctic charr (S. alpinus) in a large Fennoscandian subarctic lake

Introduced fishes may have major impacts on community structure and ecosystem function due to competitive and predatory interactions with native species. For example, introduced lake trout (Salvelinus namaycush) has been shown to replace native salmonids and induce major trophic cascades in some North American lakes, but few studies have investigated trophic interactions between lake trout and closely related native Arctic charr (S. alpinus) outside the natural distribution of the former species. We used stomach content and stable isotope analyses to investigate trophic interactions between introduced lake trout and native Arctic charr in large subarctic Lake Inarijärvi in northern Finland. Both salmonids had predominantly piscivorous diets at >280 mm total length and were mainly caught from the deep profundal zone. However, lake trout had a more generalist diet and showed higher reliance on littoral prey fish than Arctic charr, whose diet consisted mainly of pelagic planktivorous coregonids. According to length at age and condition data, lake trout showed slightly faster growth but lower condition than Arctic charr. The results indicate that introduced lake trout may to some extent compete with and prey upon native Arctic charr, but currently have only a minor if any impact on native fishes and food web structure in Inarijärvi. Future monitoring is essential to observe potential changes in trophic interactions between lake trout and Arctic charr in Inarijärvi, as well as in other European lakes where the two salmonids currently coexist.     

Resource partitioning between lake-dwelling Atlantic salmon (Salmo salar L.) parr, brown trout (Salmo trutta L.) and Arctic charr (Salvelinus alpinus (L.))

Abstract – Resource partitioning between Atlantic salmon parr, brown trout and Arctic charr was studied throughout the ice-free season in a north Norwegian lake. Juvenile salmon and trout (≤160 mm) utilized the littoral zone and juvenile charr the profundal, while adult trout and charr (>160 mm) were found in both. Juvenile salmon and trout had a similar diet, although trichopteran larvae were more important for the trout and chironomid pupae and three-spined sticklebacks for the salmon parr. Small salmon and trout parr (≤120 mm) had a higher diet overlap than larger parr (121–160 mm). The feeding habits of adult trout were similar to that of juvenile trout, but the former took larger prey items. At the population level, both salmon and trout were generalistic feeders with a broad diet, but at the individual level, both species had specialized on a single or a few prey categories. Juvenile charr were segregated from salmon and trout in both habitat and food utilization; they had a narrow diet consisting of chironomids and zooplankton, possibly reflecting their confinement to the profundal habitat which have a low diversity of potential prey. Larger charr also took zoobenthos and sticklebacks in the littoral zone. ^Note     

Trophic ecology of brown trout (Salmo trutta L.) in subarctic lakes

In subarctic lake systems, fish species like brown trout are often important predators, and their niche performance is a key characteristic for understanding trophic interactions and food web functioning at upper trophic levels. Here, we studied summer habitat use and stomach contents of brown trout under both allopatric and sympatric conditions in six subarctic lakes to reveal its trophic role, and population‐ and individual‐level niche plasticity. In allopatry, brown trout mainly used the littoral habitat, but also less commonly used the pelagic zone. In sympatry with stickleback, there was always a considerable habitat overlap between the two species. In contrast, sympatric populations of brown trout and Arctic charr generally revealed a distinct habitat segregation. In the sympatric systems, in general, there was a distinct resource partitioning between the trout and charr, whereas the observed diet overlap between trout and stickleback was much larger. Trout modified their individual dietary specialisation between the littoral and pelagic zone, always being lower in the pelagic. Piscivorous behaviour of trout was only found in sympatric systems, possibly contributing to a competitive advantage of trout over charr and stickleback. Hence, the trophic level of trout was strongly related to the fish community composition, with a higher trophic level in sympatric systems where piscivorous behaviour was frequent. These changes in the trophic level of trout linked with the observed food resource partitioning might be an important mechanism in the ecosystem functioning of subarctic lakes to allow coexistence among sympatric‐living fish species.