Non-native species limit stream restoration benefits for brook trout

Success of stream restoration can be difficult to define because many interacting abiotic and biotic factors across spatio-temporal scales can have measurable effects. Consequently, failure in habitat restoration to achieve targeted biological goals may reflect interactions of habitat restoration with unaccounted risks that have yet to be addressed on the landscape. This is particularly true within invaded landscapes, where habitat restoration can benefit non-native competitors as much as the native fishes for which restoration is designed. We tested for interacting effects of a reach scale habitat restoration effort and non-native trout competition on habitat use by a brook trout (Salvelinus fontinalis) metapopulation within a productive main stem corridor of the Shavers Fork watershed, West Virginia. We used a joint species occupancy model within a BACI sampling design to show that brook trout occupancy of main stem habitat was highest post-restoration within restored sampling reaches, but this benefit to native brook trout was conditional on brown trout (Salmo trutta) not being present within the main stem habitat. Collectively these results indicate that habitat restoration was only beneficial for native brook trout when non-native trout were absent from the restored sampling area. Proactive approaches to restoration will be integral for supporting resilient ecosystems in response to future anthropogenic threats (e.g. climate change), and we have shown that such actions will only be successful if non-native competitors do not also benefit from the restoration actions.     

Spatial patterns of hybridization between bull trout, <Emphasis Type="Italic">Salvelinus confluentus</Emphasis>, and brook trout, <Emphasis Type="Italic">Salvelinus fontinalis</Emphasis> in an Oregon stream network

Hybridization with introduced species represents a serious threat to the persistence of many native fish populations. Brook trout (Salvelinus fontinalis) have been introduced extensively throughout the native range of bull trout (S. confluentus) and hybridization has been documented in several systems where they co-exist and is seen as a significant threat to the persistence of bull trout populations. We identified a group of diagnostic microsatellite loci to differentiate bull trout and brook trout and then used these loci to examine the spatial distribution of hybrids in the Malheur River basin, Oregon USA. In random samples of approximately 100 fish from each of three creeks we identified 181 brook trout, 112 bull trout and 14 hybrids. Although bull trout, brook trout and hybrids were found in all three creeks, they were not evenly distributed; brook trout were primarily found in the lower sections of the creeks, bull trout further upstream, and hybrids in the areas of the greatest overlap. One creek with a population of brook trout in a headwater lake provided an exception to this pattern; brook trout were found distributed throughout the creek downstream of the lake. Several post-F1 hybrids were identified suggesting that hybrids are reproducing in the Malher River Basin. Mitochondrial DNA analysis indicated that both female bull trout and brook trout are involved in hybridization events. Analysis of population structure suggested that brook trout have established multiple spawning populations within the Malheur system. Data presented in this study suggest that relative abundance of brook trout and habitat quality are important factors to consider when evaluating the threat of hybridization to bull trout populations.     

Hybridization between native white-spotted charr and nonnative brook trout in the upper Sorachi River,Hokkaido, Japan

Invasion status and impacts of nonnative brook trout (Salvelinus fontinalis) in a Hokkaido stream were investigated with field surveys and genetic analyses. Nonnative brook trout was detected in nine (41 %) of the 22 sampled reaches in three tributaries of the Sorachi River, Hokkaido, Japan. Based on the external pigmentation, twelve putative hybrids between brook trout and native white-spotted charr (Salvelinus leucomaenis) were collected in two reaches. Microsatellite and mitochondrial DNA data established that 58% of these hybrids were first-generation (F₁) progenies between male brook trout and female white-spotted charr. Our results suggest potential negative impacts of nonnative brook trout on native charr populations in Hokkaido through interspecific interactions.     

Daily temperature experience and selection by adfluvial bull trout (<Emphasis Type="Italic">Salvelinus confluentus</Emphasis>)

Bull trout (Salvelinus confluentus) are a thermally sensitive cold-water species with a threatened conservation status across much of North America. To improve our understanding of bull trout thermal ecology, we tagged more than 150 adults with temperature-sensing acoustic biotelemetry transmitters and monitored the animals in a British Columbia reservoir for two years. Thermal resource selection was estimated in open water from the summer to autumn as the system transitioned to isothermal conditions. On average, bull trout thermal history tracked seasonal changes and remained close to the optimum temperatures for growth and metabolism. As summer progressed, bull trout increasingly selected temperatures of 11–15 °C as they became less available within the water column. Selection indicated the movement to shallower waters where 11–15 °C temperatures existed. The results show that bull trout mainly occupy a narrow range of temperatures which suggests the importance of cold-water refuge for the species.     

Genetic Diversity and Hybridisation between Native and Introduced Salmonidae Fishes in a Swedish Alpine Lake

Understanding the processes underlying diversification can aid in formulating appropriate conservation management plans that help maintain the evolutionary potential of taxa, particularly under human-induced activities and climate change. Here we assessed the microsatellite genetic diversity and structure of three salmonid species, two native (Arctic charr, Salvelinus alpinus and brown trout, Salmo trutta) and one introduced (brook charr, Salvelinus fontinalis), from an alpine lake in sub-arctic Sweden, Lake Ånn. The genetic diversity of the three species was similar and sufficiently high from a conservation genetics perspective: corrected total heterozygosity, H’_T = 0.54, 0.66, 0.60 and allelic richness, A_R = 4.93, 5.53 and 5.26 for Arctic charr, brown trout and brook charr, respectively. There were indications of elevated inbreeding coefficients in brown trout (G_IS = 0.144) and brook charr (G_IS = 0.129) although sibling relationships were likely a confounding factor, as a high proportion of siblings were observed in all species within and among sampling locations. Overall genetic structure differed between species, Fst = 0.01, 0.02 and 0.04 in Arctic charr, brown trout and brook charr respectively, and there was differentiation at only a few specific locations. There was clear evidence of hybridisation between the native Arctic charr and the introduced brook charr, with 6% of individuals being hybrids, all of which were sampled in tributary streams. The ecological and evolutionary consequences of the observed hybridisation are priorities for further research and the conservation of the evolutionary potential of native salmonid species.     

Effects of different protein and carbohydrate levels on growth performance and feed utilisation of brook trout,Salvelinus fontinalis (Mitchill, 1814), at two temperatures

A 12‐week feeding trial was conducted to determine the optimum dietary protein requirement of brook trout, Salvelinus fontinalis, at 15 and 19°C. Twelve iso‐energetic (22 MJ · kg^?1) and iso‐lipidic (23%) diets (36–58% protein at 2% increments) were prepared. Fish (29.45 ± 3.25 g · fish^?1) were fed 2% of body weight per day, divided into two equal rations. The specific growth rate (SGR, % · day^?1), feed efficiency ratio (FER), productive protein value (PPV), productive lipid value (PLV) and productive energy value (PEV), apparent digestibility of diet (AD_DM) and protein (AD_CP) were significantly higher at optimum temperature (15°C). Increasing PPV with increasing dietary carbohydrate and with decreasing dietary protein content was due to the protein‐sparing effect of carbohydrates. A piecewise regression (broken line) model between the SGR and digestible dietary protein level revealed that the digestible dietary protein requirement of brook trout was 44 and 40% at 15 and 19°C, respectively. When PPV (digestible protein retention basis) was modelled with a broken line, the digestible protein requirement of brook trout was 39 and 35% at 15 and 19°C, respectively. A reduction in dietary protein content balanced by increased gelatinised carbohydrate might be useful for improving the protein utilization efficiency for growth at 15 and 19°C; however, the growth and feed efficiency was lower at the elevated temperature.     

More than a corridor: use of a main stem stream as supplemental foraging habitat by a brook trout metapopulation

Coldwater fishes in streams, such as brook trout (Salvelinus fontinalis), typically are headwater specialists that occasionally expand distributions downstream to larger water bodies. It is unclear, however, whether larger streams function simply as dispersal corridors connecting headwater subpopulations, or as critical foraging habitat needed to sustain large mobile brook trout. Stable isotopes (δ¹³C and δ¹⁵N) and a hierarchical Bayesian mixing model analysis was used to identify brook trout that foraged in main stem versus headwater streams of the Shavers Fork watershed, West Virginia. Headwater subpopulations were composed of headwater and to a lesser extent main stem foraging individuals. However, there was a strong relationship between brook trout size and main stem prey contributions. The average brook trout foraging on headwater prey were limited to 126 mm standard length. This size was identified by mixing models as a point where productivity support switched from headwater to main stem dependency. These results, similar to other studies conducted in this watershed, support the hypothesis that productive main stem habitat maintain large brook trout and potentially facilitates dispersal among headwater subpopulations. Consequently, loss of supplementary main stem foraging habitats may explain loss of large, mobile fish and subsequent isolation of headwater subpopulations in other central Appalachian watersheds.     

Hydrological Variation and Fish Assemblage Structure in the Middle Wabash River

We assessed the effect of a severe drought in 1999 upon stream morphology and brook trout (Salvelinus fontinalis) populations in seven headwater streams in the Greenbrier and Potomac River watersheds, West Virginia. During the drought, stream discharge was 96% lower than in years of normal precipitation. As a result, habitat availability and quality over all study streams was significantly lower. Riffle area was greatly reduced (?54%) relative to available pool area (?2%). Fine sediment levels (<0.063 mm) significantly increased within spawning substrate (p=0.01). Water temperature and dissolved oxygen were adequate (mean 15.8?°C, >6.0 mg l^?1, respectively) for brook trout survival in all streams during the drought. Brook trout populations were significantly reduced (adult 60%, Young-of-the-year 67%), and individual fish had significantly lower body condition during the drought relative to the post-drought period. Reductions in brook trout density and population condition during, and in the-post drought period, were related to spatially-limited food resources and/or increased fine sediment levels, but not to degraded water quality. Fisheries managers should consider the effect of periodic drought on brook trout populations and consider short-term harvest restrictions to abet recovery after such stochastic events.     

Sympatric relationship between redband trout and non-native brook trout in the Southeastern Oregon Great Basin

Brook trout (Salvelinus fontinalis) and rainbow trout (Oncorhynchus mykiss) have been widely introduced outside their respective ranges within North America causing declines and displacement of native trout. Yet, successful coexistence of native and non-native trout has received little attention. Here we evaluated the effect of introduced brook trout on the size and density of native redband trout in two invaded sub-basins in southeastern Oregon. In a multi-year study, we investigated whether habitat and fish communities differed between streams and stream reaches where redband trout were allopatric versus where redband trout were sympatric with brook trout. We hypothesized that redband trout would be less dense and have smaller total length in sympatry with brook trout than in allopatry, but that total trout density would not differ. We investigated whether differences in habitat existed between sympatric and allopatric locations that would indicate differentiation in site level habitat preferences for each trout species. We found that sympatric locations had more wood but similar fish community structure. Mean length and densities of redband trout were higher at allopatric locations. However, in most years at sympatric locations total trout density was twice that of allopatric redband trout sites. Using comparable data from an eastern United States system where brook trout are native, sympatric sites had lower densities of brook trout; however, total trout density did not differ. We conclude that invading trout negatively impact native trout densities; but in southeastern Oregon system the negative impact is minimized.     

Distinguishing between juvenile anadromous and resident brook trout (Salvelinus fontinalis) using morphology

Phenotypic variation linked to habitat use has been observed in fish, both between and within species. In many river systems, migratory and resident forms of salmonids coexist, including anadromous (migrant) and resident brook trout, Salvelinus fontinalis. In such populations, juvenile anadromous (migrant) brook trout, prior to migration, inhabit regions of higher current velocity than residents. Because it is more costly to occupy fast currents than slow currents, differences in morphology minimizing the effects of drag were expected between the two forms. As predicted, migrant brook trout were found to be more streamlined (narrower and shallower bodies) than resident brook trout, and these differences persisted into the marine life of the fish. Migrants also exhibited shorter pectoral fins, which facilitate pelagic swimming, indicating that migrants, prior to their migration to the sea, possess the appropriate morphology for swimming in open water habitats. The reported differences between migrants and residents were powerful enough to derive discriminant functions, using only five of the seven measured traits, allowing for accurate classification of brook trout as either migrants or residents with an overall correct classification rate of 87%. Importantly, this study contributes to the notion that a link exists between morphology, habitat use, metabolic costs and life-history strategies. Contribution to the program of CIRSA (Centre Interuniversitaire de Recherche sur le Saumon Atlantique).     

Do Low-Mercury Terrestrial Resources Subsidize Low-Mercury Growth of Stream Fish? Differences between Species along a Productivity Gradient

Low productivity in aquatic ecosystems is associated with reduced individual growth of fish and increased concentrations of methylmercury (MeHg) in fish and their prey. However, many stream-dwelling fish species can use terrestrially-derived food resources, potentially subsidizing growth at low-productivity sites, and, because terrestrial resources have lower MeHg concentrations than aquatic resources, preventing an increase in diet-borne MeHg accumulation. We used a large-scale field study to evaluate relationships among terrestrial subsidy use, growth, and MeHg concentrations in two stream-dwelling fish species across an in-stream productivity gradient. We sampled young-of-the-year brook trout (Salvelinus fontinalis) and Atlantic salmon (Salmo salar), potential competitors with similar foraging habits, from 20 study sites in streams in New Hampshire and Massachusetts that encompassed a wide range of aquatic prey biomass. Stable isotope analysis showed that brook trout used more terrestrial resources than Atlantic salmon. Over their first growing season, Atlantic salmon tended to grow larger than brook trout at sites with high aquatic prey biomass, but brook grew two-fold larger than Atlantic salmon at sites with low aquatic prey biomass. The MeHg concentrations of brook trout and Atlantic salmon were similar at sites with high aquatic prey biomass and the MeHg concentrations of both species increased at sites with low prey biomass and high MeHg in aquatic prey. However, brook trout had three-fold lower MeHg concentrations than Atlantic salmon at low-productivity, high-MeHg sites. These results suggest that differential use of terrestrial resource subsidies reversed the growth asymmetry between potential competitors across a productivity gradient and, for one species, moderated the effect of low in-stream productivity on MeHg accumulation.     

Competitive interactions for foraging microhabitat among introduced brook charr, Salvelinus fontinalis, and native bull charr, S. confluentus, and westslope cutthroat trout, Oncorhynchus clarki lewisi, in a Montana stream

Competitive interactions for foraging microhabitat among introduced brook charr, Salvelinus fontinalis, and native bull charr, S. confluentus, and westslope cutthroat trout, Oncorhynchus clarki lewisi, were studied by species removal experiments in a tributary of the Flathead Lake and River system, northwestern Montana, focusing on brook charr influences on bull charr. When the three species were in sympatry, they interacted with each other, forming a size-structured, mixed-species dominance hierarchy in two stream pools. The influences of interference interactions were examined by measuring changes in five characteristics of foraging microhabitat and behavior, focal point height and velocity, cover use, and foraging rate and distance, after the successive removal of two species. Cutthroat trout removal resulted in increased foraging rates and distances, and decreased cover use for brook charr, but no changes for bull charr. After removal of brook charr from the two-species system, bull charr also increased foraging rates and distances and occupied more exposed positions. Moreover, total fish densities, which had initially decreased owing to the removal experiments, were partly compensated for by subsequent bull charr immigration, implying that competitive interactions with brook charr are an important factor in the mechanisms responsible for the regulation of bull charr densities, at least on a local scale.     

Re-awakening dormant life history variation: stable isotopes indicate anadromy in bull trout following dam removal on the Elwha River,Washington

Migratory species take advantage of multiple habitats during their life cycle to optimize growth, survival, and reproduction. However, migration also makes them vulnerable to habitat degradation and exploitation in each habitat, and loss of connection between habitats. Partially migratory species (i.e., migration is facultative rather than obligate) can persist after loss of connectivity and may then resume migration after the habitats are reconnected. We analyzed stable isotopes of carbon and nitrogen to investigate the possible use of marine habitats for foraging by bull trout, Salvelinus confluentus, in years immediately after removal of impassable hydroelectric dams on the Elwha River, Washington State, USA. Juveniles in the Elwha River estuary were similar in δ¹⁵N and δ¹³C values to those in the estuary of the free-flowing Dungeness River nearby, and the values of fish from the estuaries were higher than those of juveniles collected in the river, consistent with use of marine food sources. Adult bull trout collected in each of the rivers had values indicating extensive reliance on marine prey - similar to those of adult Pacific salmon that had spent several years at sea. Taken together, these data demonstrate that the Elwha River bull trout, almost entirely landlocked for a century, are rapidly resuming anadromy and that the marine prey contribute substantially to their trophic ecology and likely their growth. More broadly, the results reveal the importance of connectivity for migratory fishes, their ability to resume anadromy once the connection between habitats is restored, and the population resilience that partial migration provides for them.     

Decline of the Migratory Form in Bull Charr, Salvelinus Confluentus, and Implications for Conservation

Large-bodied, migratory life history forms of bull charr, Salvelinus confluentus, were historically abundant in northwestern North America, but many remaining populations of this now-threatened species presently persist as small-bodied residents isolated in headwater streams. We examined whether the migratory form has been lost from headwater populations of bull charr and their potential for re-establishment. Upstream and downstream movement of bull charr and other salmonids from three tributary populations in the Bitterroot River drainage, Montana, was measured with weirs over a 17-month period. The migratory life history was rare or absent in two tributaries but still present at a low level in a third. In contrast, substantial numbers (n = 1745) of juvenile and adults of other salmonids (brown trout, Salmo trutta, cutthroat trout, Oncorhynchus clarki, and mountain whitefish, Prosopium williamsoni) were captured near tributary mouths, indicating a migratory life history was common in other species. Apparent decline of the migratory life history in bull charr was not directly related to damming suggesting other downstream mortality factors (predation, temperature) also are involved. Isolated, nonmigratory forms have increased risk of extinction, and restoration of the population connectivity via the re-establishment of migratory stocks is an important conservation goal for bull charr recovery. However, the factors governing migratory tendency remain unclear.     

Cytogenetic analysis of pseudolinkage of ldh Loci in the teleost genus salvelinus

Cytological and genetic analyses provide evidence that spontaneous centric fusion and fission can account for curious patterns of pseudolinkage of two LDH loci in males of brook trout (Salvelinus fontinalis) and in the F₁, F₂ and backcross generations of lake trout (S. namaycush) x brook trout hybrids. Intraindividual polymorphisms for acrocentric and metacentric chromosomes in somatic and gonadal tissue of these fish have been related to the proposed polyploid evolution in Salmonidae.     

Opportunistic use of estuarine habitat by juvenile bull trout, <Emphasis Type="Italic">Salvelinus confluentus</Emphasis>, from the Elwha River before,during, and after dam removal

Estuaries are used by anadromous fishes, either as the definitive marine habitat or as transition habitat as they move to fully marine waters, and extent of estuary use may vary with habitat conditions and fish attributes. Bull trout (Salvelinus confluentus) are commonly fluvial or adfluvial, though anadromous populations also exist. However, little is known about estuary use, especially by juveniles of this threatened species. We sampled the estuaries of the Elwha River, where a spawning population exists, and the nearby Salt Creek, where none exists, to reveal seasonal timing of estuarine use by juvenile bull trout, size of those using the estuary, and possible use of the non-natal estuary. We captured juvenile bull trout (all ≥100 mm FL, most <300 mm) in the Elwha River estuary in all months except August, but primarily December through May. None was captured in Salt Creek’s estuary despite comparable sampling effort. We also evaluated how dam removal on the Elwha River influenced bull trout estuarine occupancy by sampling before, during, and after dam removal, because this process enlarged the estuary but also increased turbidity and sediment transport in the lower river. Catches were low before dam removal, increased during and immediately after removal, and returned to low levels in recent years, suggesting that juveniles temporarily sought refuge from conditions associated with dam removal. Our findings indicate juvenile bull trout occupy estuarine habitat opportunistically; this information may aid conservation efforts as anadromous populations occur elsewhere in rivers with estuaries altered by human development.     

Characterizing genetic integrity of rear-edge trout populations in the southern Appalachians

Vertebrate populations at the periphery of their range can show pronounced genetic drift and isolation, and therefore offer unique challenges for conservation and management. These populations are often candidates for management actions such as translocations that are designed to improve demographic and genetic integrity. This is particularly true of coldwater species like brook trout (Salvelinus fontinalis), whose numbers have declined greatly across its historic range. At the southern margin, remnant wild populations persist in isolated headwater streams, and many have a history of receiving translocated individuals through either stocking of hatchery reared fish, relocation of wild fish, or both during restoration attempts. To determine current genetic integrity and resolve the genetic effects of past management actions for brook trout populations in SC, USA, we genetically assessed all 18 documented remaining brook trout populations along with individuals acquired from six hatcheries with recorded stocking events in SC. Our results indicated that six of the 18 streams showed signs of hatchery admixture (range 57–97%) and restored patches retained genetic signatures from multiple source populations. Populations had among the lowest genetic diversity (min average H_E?=?0.147) and effective number of breeders (mean N_b?=?31.2) estimates observed throughout the native brook trout range. Populations were highly differentiated (mean pair-wise F_ST?=?0.396), and substantial genetic divergence was evident across major river drainages (max pair-wise F_ST?=?0.773). The lowest local genetic diversity and highest genetic differentiation ever reported for this species make its conservation a challenging task, particularly when combined with other threats such as climate change and non-native species. We offer recommendations on managing peripheral populations with depleted genetic characteristics and provide a reference for determining which existing populations will best serve as sources for future translocation efforts aimed at enhancing or restoring wild brook trout genetic integrity.     

Use of cover habitat by bull trout, <Emphasis Type="Italic">Salvelinus confluentus</Emphasis>, and lake trout, <Emphasis Type="Italic">Salvelinus namaycush</Emphasis>, in a laboratory environment

Lacustrine-adfluvial bull trout, Salvelinus confluentus, migrate from spawning and rearing streams to lacustrine environments as early as age 0. Within lacustrine environments, cover habitat provides refuge from potential predators and is a resource that is competed for if limiting. Competitive interactions between bull trout and other species could result in bull trout being displaced from cover habitat, and bull trout may lack evolutionary adaptations to compete with introduced species, such as lake trout, Salvelinus namaycush. A laboratory experiment was performed to examine habitat use and interactions for cover by juvenile (i.e., <80 mm total length) bull trout and lake trout. Differences were observed between bull trout and lake trout in the proportion of time using cover (F _1,22.6 = 20.08, P < 0.001) and bottom (F _1,23.7 = 37.01, P < 0.001) habitat, with bull trout using cover and bottom habitats more than lake trout. Habitat selection ratios indicated that bull trout avoided water column habitat in the presence of lake trout and that lake trout avoided bottom habitat. Intraspecific and interspecific agonistic interactions were infrequent, but approximately 10 times greater for intraspecific interactions between lake trout. Results from this study provide little evidence that juvenile bull trout and lake trout compete for cover, and that species-specific differences in habitat use and selection likely result in habitat partitioning between these species.     

On brook chars of the southern Kuril Islands and a possible mechanism of formation of dwarf forms of Dolly Varden trout Salvelinus malma curilus

The specific morphological and biological features of dwarf chars Salvelinus malma curilus from three isolated populations of the southern Kuril Islands Kunashir and Iturup were studied. It was shown that brook dwarfs, in comparison with the migratory form of Salvelinus malma curilus, are characterized by a smaller diameter of spawned eggs (which corresponds to smaller yolk reserves and shorter embryos at hatching), retarded growth and morphogenesis of early juveniles, a mosaic pattern of paedomorphic and peramorphic morphological transformations, and a short lifespan. The population and age variability of the relative sizes and shape of otoliths was revealed. Dwarf populations are typically characterized by considerable morphological differences between males and females corresponding to those in migratory fish. The strategy of dwarfism is assumed to be built into the genome of the migratory ecotype of S. malma curilus.     

Trophic relationships of two species of grebe on a prairie lake based on stable isotope analysis

Loch Leven, U.K., contains brown trout (Salmo trutta), eel (Anguilla anguilla), minnow (Phoxinus phoxinus), perch (Perca fluviatilis), pike (Esox lucius) and three-spined stickleback (Gasterosteus aculeatus), with brook lamprey (Lampetra planeri) and stone loach (Barbatula barbatula) also present in its tributaries. Arctic charr (Salvelinus alpinus), Atlantic salmon (Salmo salar) and flounder (Platichthys flesus) are now extinct. The brown trout population has supported a world-renowned recreational fishery for over a century, although a decline in fishery performance led to extensive stocking between 1983 and 2006, including with non-native rainbow trout (Oncorhynchus mykiss). This review combines historical information with contemporary gill-net and hydroacoustic surveys. In 2008, brown trout, perch and three-spined sticklebacks were abundant, but pike and stone loach were rare. The obstruction of migratory routes was probably responsible for the loss of Atlantic salmon and flounder, while a lowering of water level likely caused the extinction of Arctic charr and contributed to a reduction in pike abundance. Perch abundance has fluctuated markedly, being influenced by disease and eutrophication, although a reduction in nutrients and associated recovery of macrophytes are likely to have benefitted this species. Although the brown trout population has undoubtedly shown a long-term decline, individuals are currently in excellent condition.