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.     

Comparing competitive ability and associated metabolic traits between a resident and migratory population of bull trout against a non-native species

Juvenile bull trout Salvelinus confluentus from two geographically and ecologically distinct populations were compared with regard to their ability to compete with non-native brook trout Salvelinus fontinalis in an artificial stream, and with respect to their rates of oxygen consumption. Bull trout collected from a migratory population foraged more successfully against brook trout competitors than those from a resident population, capturing more of a limited amount of food items presented. The migratory population was also more aggressive (measured by the number of nips, chases and lateral threat displays) against brook trout competitors than the resident population. Bull trout from the migratory population had a higher oxygen consumption rate (203 mg O₂ kg · hr^-1) in the field than similar sized fish from the resident population (183 mg O₂ kg · hr^-1). These results suggest native bull trout have population-level variation in competitive ability against a non-native species and such competitive ability is positively associated with metabolism and migratory life history.     

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.     

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.     

Assessing the impact of stocking northern-origin hatchery brook trout on the genetics of wild populations in North Carolina

The release of hatchery-origin fish into streams with endemics can degrade the genetics of wild populations if interbreeding occurs. Starting in the 1800s, brook trout descendent from wild populations in the northeastern United States were stocked from hatcheries into streams across broad areas of North America to create and enhance fishery resources. Across the southeastern United States, many millions of hatchery-origin brook trout have been released into hundreds of streams, but the extent of introgression with native populations is not well resolved despite large phylogeographic distances between these groups. We used three assessment approaches based on 12 microsatellite loci to examine the extent of hatchery introgression in 406 wild brook trout populations in North Carolina. We found high levels of differentiation among most collections (mean F′_ST = 0.718), and among most wild collections and hatchery strains (mean F′_ST = 0.732). Our assessment of hatchery introgression was consistent across the three metrics, and indicated that most wild populations have not been strongly influenced by supplemental stocking. However, a small proportion of wild populations in North Carolina appear to have been strongly influenced by stocked conspecifics, or in some cases, may have been founded entirely by hatchery lineages. In addition, we found significant differences in the apparent extent of hatchery introgression among major watersheds, with the Savannah River being the most strongly impacted. Conversely, populations in the Pee Dee River watershed showed little to no evidence of hatchery introgression. Our study represents the first large-scale effort to quantify the extent of hatchery introgression across brook trout populations in the southern Appalachians using highly polymorphic microsatellite markers.     

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.     

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.     

Non‐native trout limit native brook trout access to space and thermal refugia in a restored large‐river system

We used direct observation via snorkeling surveys to quantify microhabitat use by native brook (Salvelinus fontinalis) and non‐native brown (Salmo trutta) and rainbow (Onchorynchus mykiss) trout occupying natural and restored pool habitats within a large, high‐elevation Appalachian river, United States. Permutational multivariate analysis of variance (PERMANOVA) and subsequent two‐way analysis of variance (ANOVA) indicated a significant difference in microhabitat use by brook and non‐native trout within restored pools. We also detected a significant difference in microhabitat use by brook trout occupying pools in allopatry versus those occupying pools in sympatry with non‐native trout—a pattern that appears to be modulated by size. Smaller brook trout often occupied pools in the absence of non‐native species, where they used shallower and faster focal habitats. Larger brook trout occupied pools with, and utilized similar focal habitats (i.e. deeper, slower velocity) as, non‐native trout. Non‐native trout consistently occupied more thermally suitable microhabitats closer to cover as compared to brook trout, including the use of thermal refugia (i.e. ambient–focal temperature >2°C). These results suggest that non‐native trout influence brook trout use of restored habitats by: (1) displacing smaller brook trout from restored pools, and (2) displacing small and large brook trout from optimal microhabitats (cooler, deeper, and lower velocity). Consequently, benefits of habitat restoration in large rivers may only be fully realized by brook trout in the absence of non‐native species. Future research within this and other large river systems should characterize brook trout response to stream restoration following removal of non‐native species.     

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.     

Selenium Species and Their Distribution in Freshwater Fish from Argentina

The distribution and speciation of selenium (Se) in freshwater fish (muscle and liver tissue) from lakes in Argentina was investigated. Three introduced species, brown trout (Salmo trutta), rainbow trout (Oncorhynchus mykiss) and brook trout (Salvelinus fontinalis), and one native species, creole perch (Percichthys trucha), were investigated. Values for total selenium in muscle ranged from 0.66 to 1.61 μg/g, while in the liver, concentrations were much higher, from 4.46 to 73.71 μg/g on a dry matter basis. Separation of soluble Se species (SeCys₂, selenomethionine (SeMet), SeMeSeCys, selenite and selenate) was achieved by ion exchange chromatography and detection was performed by inductively coupled plasma–mass spectrometry. The results showed that in fish muscle, from 47 to 55 % of selenium was soluble and the only Se species identified was SeMet, which represented around 80 % of soluble Se, while in the liver, the amount of soluble Se ranged from 61 to 76 % and the percentage of species identified (SeMet and SeCys₂) was much lower and ranged from 8 to 17 % of soluble Se.     

Shifting thermal regimes influence competitive feeding and aggression dynamics of brook trout (Salvelinus fontinalis) and creek chub (Semotilus atromaculatus)

The natural distributions of freshwater fish species are limited by their thermal tolerances via physiological constraints and increased interspecific competition as temperatures shift toward the thermal optima of other syntopic species. Species may mediate stress from temperature change physiologically, behaviorally, or both; but these changes may compromise competitive advantages through effects on feeding and social behavior. In the Appalachian Mountains of North America, creek chub (Semotilus atromaculatus) are found in warm‐water and cold‐water streams and overlap in range with brook trout (Salvelinus fontinalis) across lower thermal maxima, where they compete for food and space. As stream temperatures continue to increase due to climate change, brook trout are under increasing thermal stress which may negatively affect their ability to compete with creek chub. To examine the influences of temperature on competitive interactions between these species, we observed feeding behavior, aggression, and habitat use differences at three temperatures approaching brook trout thermal maxima (18°C, 20°C, and 22°C) among dyad pairs for all combinations of species in experimental flow‐through tanks. We also examined feeding and habitat use of both species under solitary conditions. We found as temperature increased, feeding and aggression of brook trout were significantly reduced in the presence of creek chub. Creek chub pairs were more likely to occupy benthic areas and refugia while brook trout pairs used surface water more. Space use patterns significantly changed by pairing treatment. Aggression and space use shifts allowed increased exploitative and interference competition from creek chub when paired with brook trout that was not present in conspecific pairs. The decreased dominance of a top predator may lead to diverse impacts on stream community dynamics with implications for the future range restriction of brook trout and demonstrate possible mechanisms to facilitate competitive advantages of warm water generalist species under thermal stress.     

Joint Segregation of Biochemical Loci in Salmonidae. II. Linkage Associations from a Hybridized Salvelinus Genome (S. NAMAYCUSHxS. FONTINALIS)

The results of more than 300 pairwise examinations of biochemical loci for joint segregation in brook trout (Salvelinus fontinalis) and in the hybridized genome of lake trout (S. namaycush) x brook trout are summarized. Nineteen loci have been assigned to the following eight linkage groupings on the basis of nonrandom assortment, including cases of both classical linkage and pseudolinkage: ODH with PMI with PGI-3, PGI-2 with SDH, ADA-1 with AGP-2, AAT-(1,2) with AGP-1 with MDH-1, MDH-3 with MDH-4, LDH-3 with LDH-4, IDH-3 with ME-2 and GUS with CPK-1. Pseudolinkage (an excess of nonparental progeny types) was observed only for male testcross parents. The results suggest that this phenomenon involves homeologous chromosome arms as evidenced by the de novo association of presumed duplicate loci in each case. Classical linkage has not been found for the five pairs of duplicate loci examined in Salvelinus, suggesting that not all of the eight metacentrics in the haploid complement involve fusions of homeologous chromosomes. Females consistently showed a greater degree of recombination.     

Can replacement of native by non‐native trout alter stream‐riparian food webs?

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Heterochromatins and band karyotypes in three species of salmonids

Summary The heterochromatins of rainbow trout (Salmo gairdneri R.), brown trout (Salmo trutta fario L.) and brook trout (Salvelinus fontinalis M.) were characterized by sequential chromomycin A₃/distamycin A/DAPI (CDD) and DAPI/actinomycin D (DAPI/AmD) fluorescence. On most biarmed chromosomes, an equilocal localization of prominent DAPI/AmD positive, chromomycin A₃ negative, AT-rich blocks at the centromeres were observed in all three species. Band karyotypes of the three species were established. In rainbow trout, several DAPI/AmD positive heterochromatin blocks behaved positive in a silver-staining method. Mitotic and interphase studies proved the presence of inter-individual NOR variation in brown trout. The NORs of brook trout were localized on chromosomes 5, 10, 14, 15 and 29.     

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.     

Invasion of north European streams by brook trout: hostile takeover or pre-adapted habitat niche segregation?

We combine evidence from small-scale experiments with a large-scale field survey to clarify the roles of biotic resistance and pre-adapted habitat niche segregation to the invasion success of the North American brook trout (Salvelinus fontinalis) in North European streams previously dominated by brown trout (Salmo trutta). Interspecific aggressions among the two species were negligible, yet there was distinct habitat niche segregation between them: brook trout occupied mainly pool habitats while brown trout tended to reside in fast-flowing riffles. Habitat niche segregation among brook trout and brown trout prevailed across a wide array of scales from experimental flumes to entire drainage systems, although the segregation pattern was weaker in the field. Habitat differentiation among the two species reflected their differential habitat requirements, suggesting that a match between a species’ niche requirements in its native range and habitat availability in the new environment is a prerequisite for understanding invasion success.     

Limestone Treatment of Whetstone Brook, Massachusetts. II. Changes in the Brown Trout (Salmo trutta)and Brook Trout (Salvelinus fontinalis)Fishery

Density, age structure, and growth rates of wild brook trout (Salvelinus fontinalis)and brown trout (Salmo trutta)in Whetstone Brook in northcentral Massachusetts were monitored for 4 years before and 3 years during limestone treatment to mitigate acidic conditions. The population density of brook trout increased significantly during treatment. Liming did not have any significant effects on the growth rates of brook trout or brown trout. Actual survival rates of brook trout and brown trout were not calculated due to the low density of both species, but more older individuals of both species were captured during the treatment period. Fulton condition factors (an index of fish condition) increased significantly for both brook trout and brown trout during treatment. Seven-day in situ bioassays of brown trout and rainbow trout demonstrated that liming improved the chemical environment for fish in Whetstone Brook. During a pretreatment bioassay in 1987, 100% rainbow trout mortality was observed at both the control and treatment stations in Whetstone Brook. Brown trout mortality was 67% in the control station and 70% in the treatment station. The pH during the 1987 bioassay averaged 4.90 in the control station and 4.99 in the treated station. During a bioassay conducted in 1990 after treatment began, rainbow trout mortality was 100% in the control station and 0% in the treatment station. Brown trout mortality was 17% in the control station and 0% in the treatment station. The pH during the 1990 bioassay averaged 5.23 in the control station and 6.60 in the treatment station. Analysis of total aluminum in the gills of fish from the 1990 bioassay revealed higher levels in fish from the control station than in those from the treatment station.     

Cytogenetic identification of genomic elements of brook trout (Salvelinus fontinalis) in the karyotype of Arctic char (Salvelinus alpinus) from the aquaculture broodstock

Growing interest of Arctic char (Salvelinus alpinus) aquaculture in Europe, and the fact that it can easily hybridize with brook trout (Salvelinus fontinalis) resulting in fertile progeny, led us to investigate fish from the farmed stocks. Chromosomes of sampled Arctic char were examined using conventional and molecular cytogenetic (FISH) techniques in order to determine possible contamination of genomic elements of brook trout. Investigated fish possessed karyotypes composed of 80–82 chromosomes and up to three chromosome fragments. Using staining methods and FISH approach enabled identification of the brook trout chromosomes in the eight out of twenty‐two examined Arctic char. Specific location of AT‐, GC‐ positive and NOR sites observed on chromosomes as well as chromosome fragments in the karyotypes of several individuals points on past chromosomal rearrangements in fish from examined broodstock. Based on our results, it may be assumed that individuals with the brook trout genomic elements, although phenotypically identified as Arctic chars, were hybrids. Our results highlights that special care should be taken to protect gene pools of brook trout and Arctic char in farms where both species are cultured.     

Watershed‐level brook trout genetic structuring: Evaluation and application of riverscape genetics models

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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.