The effects of introduced salmonids on two native stream‐dwelling salmonids through interspecific competition

Using an artificial stream, habitat use by two sympatric native salmonids in the presence and absence of introduced salmonid species was investigated experimentally. When only native white‐spotted charr Salvelinus leucomaenis and masu salmon Oncorhynchus masou were sympatric, they occupied different microhabitats. In the presence of introduced brown trout Salmo trutta or rainbow trout Oncorhynchus mykiss , however, white‐spotted charr and masu salmon were observed to use a similar habitat and interspecific competition between white‐spotted charr and masu salmon was initiated. The study suggested that the coexistence of native salmonids was negatively affected through interspecific competition between native and introduced salmonids.     

Redd superimposition by introduced rainbow trout,oncorhynchus mykiss, on native charrs in a japanese stream

Spawning redd superimposition of introduced, spring-spawning rainbow trout,Oncorhynchus mykiss, on native, fall-spawning Dolly varden,Salvelinus malma, and white-spotted charr,S. leucomaenis, were examined in a small stream in Hokkaido, Japan. The stream reaches in which Dolly Varden and white-sported charr redds were observed in fall 1997 greatly overlapped with the reaches in which rainbow trout redds were recorded in spring 1998. Spawning microhabitats were also similar between trout and the two charr species. Thirteen and 3% of Dolly Varden and white-spotted charr redds, respectively, were superimposed by rainbow trout redds. The eggs or alevins in the disturbed charr redds were potentially damaged because charrs were not likely to have emerged from the redds before the superimposition occurred. In sufficiently great abundance, introduced rainbow trout may negatively impact native charr populations by dislodging the latter’s spawning redds.     

Seasonal changes in stream salmonid population densities in two tributaries of a boreal river in northern Japan

We examined seasonal changes in population densities of stream salmonids (masu salmon Oncorhynchus masou, white-spotted charr Salvelinus leucomaenis, and rainbow trout O. mykiss) in two tributaries of the Shoro River, eastern Hokkaido, Japan. In one small tributary, water temperature was relatively high during the winter, and populations of salmon and trout increased through immigration at this time of the year, becoming dominant components of the salmonid assemblage; the density of charr in this stream decreased during the winter, but charr was dominant during the summer. In another medium-sized tributary, the water temperature fell to close to 0°C during the winter, and densities of salmon and charr decreased in this season, through emigration; trout were very rare in this stream. Seasonal patterns of stream salmonid densities vary among species and between localities, resulting in seasonal changes in species composition. For a comprehensive understanding of population processes, a whole-river survey across seasons will be necessary.     

Size-structured Interactions between Native and Introduced Species: Can Intraguild Predation Facilitate Invasion by Stream Salmonids?

Dynamics of biological invasions may be complicated in size-structured animal populations. Differences in timing of life history events such as juvenile emergence create complex interaction webs where different life stages of native and non-native species act as predators, competitors, and prey. Stream salmonids are an ideal group for studying these phenomena because they display competition and predation in size-structured populations and have been introduced worldwide. For example, introduced rainbow trout (Oncorhynchus mykiss) are invading streams of Hokkaido Island, Japan and have caused declines in native masu salmon (O. masou) populations. However, age-0 rainbow trout emerge later than age-0 masu salmon and are smaller, which raises the question of why they are able to recruit and therefore invade in the face of a larger competitor. We conducted experiments in laboratory stream channels to test effects of increasing density of age-0 and age-1 rainbow trout on age-0 masu salmon. Age-1 rainbow trout dominated age-0 masu salmon by aggressive interference, relegating them to less favorable foraging positions downstream and reducing their foraging frequency and growth. The age-1 trout also reduced masu salmon survival by predation of about 40% of the individuals overall. In contrast, age-0 rainbow trout had little effect on age-0 masu salmon. Instead, the salmon dominated the age-0 trout by interference competition and reduced their survival by predation of 60% of the individuals. In each case, biotic interactions by the larger species on the smaller were strongly negative due to a combination of interspecific competition and intraguild predation. We predict that together these produce a positive indirect effect in the interaction chain that will allow the recruitment of rainbow trout in the face of competition and predation from age-0 masu salmon, and thereby facilitate their invasion in northern Japan.     

Strategies for the conservation and management of isolated salmonid populations: lessons from Japanese streams

1. Endangered native populations of stream salmonids in Japan face three major threats: (i) negative interactions with introduced hatchery‐reared fish, (ii) fragmentation of habitat by impassable dams and (iii) recreational angling. 2. To prevent imminent extinction of many local populations, we evaluated these threats and possible conservation actions for red‐spotted masu salmon (Oncorhynchus masou ishikawae) and white‐spotted charr (Salvelinus leucomaenis japonicus) in the Fuji River system in central Japan. 3. Red‐spotted masu salmon and white‐spotted charr occupied only 0.73 and 2.4% of suitable thermal habitats, respectively, with masu salmon typically occupying habitats closer to human population centres. 4. Population viability analysis resulted in a 100‐year probability of extinction of 78.1% for masu salmon and 48.1% for charr. However, extinction risk of both species was predicted to be <5% if the carrying capacity increased from 141 to 303 for masu salmon and from 94 to 125 for charr, by allowing fish passage at the lower end of the habitat, and if annual adult survival rate increased by 0.04. Adult survival rate was the principal factor associated with population persistence. 5. To conserve isolated populations of stream‐dwelling salmonids, we recommend (i) assessing the distribution of remnant native and non‐native fish populations, (ii) that fishing regulations are modified to improve adult survival and population persistence and (iii) that fragmented reaches be reconnected to adjacent habitat, for example by removing or modifying artificial barriers to increase the carrying capacity of the isolated populations. Reconnection of fragmented reaches should, however, be avoided if it results in non‐native fish invading isolated populations.     

Wild masu salmon is outcompeted by hatchery masu salmon,a native invader,rather than brown trout,a nonnative invader

Artificially grown native species are released into natural environments to increase biological resources or to recover threatened populations. Such stocks typically have enhanced survivability and may outcompete wild conspecifics as so-called native invaders. In addition, it is likely that the competitive effects of native invaders on native species are more intense than those of nonnative invaders. To test these hypotheses, an enclosure experiment was conducted using young-of-the-year wild and hatchery (normally grown to a relatively large size to increase survival after stocking) native masu salmon, Oncorhynchus masou, and nonnative brown trout, Salmo trutta (which attain a smaller size than masu salmon). Competitive effects between these fishes were evaluated in terms of stomach fullness and specific growth rate of the wild masu salmon. The magnitude of the relationship between stomach fullness and growth between the experimental treatments revealed a similar pattern, suggesting that competition for foraging habitat most affected their growth. Wild masu salmon were negatively affected by hatchery conspecifics, and the effects were greater than those caused by brown trout. We propose that these outcomes were caused by competitive dominance as a consequence of body size differences. In conclusion, the results support the hypothesis that size-enhanced hatchery masu salmon have the potential to function as native invaders, and the negative effects of artificial stocks on wild masu salmon could be greater than those caused by a nonnative invader.     

Habitat fragmentation by damming threatens coexistence of stream-dwelling charr and salmon in the Fuji River,Japan

Habitat fragmentation by damming can affect the persistence of single species population and also coexistence of two or more species through intensified competition. This study examined the effects of habitat fragmentation by damming on the coexistence of two stream-dwelling salmonids: the southern form of white-spotted charr (Salvelinus leucomaenis japonicus) and the red-spotted masu salmon (Oncorhynchus masou ishikawae). We examined charr, salmon, and dam distributions in 27 streams of the Fuji River basin, central Japan. In the 1970s, there were streams with five sympatric and 22 allopatric populations (n = 13 for charr, n = 9 for salmon). However, from the 1970s to 2004, 356 impassable dams were constructed in the surveyed streams, and four of the five sympatric streams became allopatric. In the extant sympatric stream, more than 20 dams fragmented habitat. Species distributions were separated by dams (with decreasing altitude) in the following order: extirpation area, charr-dominant area, and salmon-dominant area. Within the uppermost sympatric section (i.e., situated between the dams), salmon congregated in the largest uppermost pool just below the dam; despite these conditions, salmon frequency increased in the downstream direction at the stream scale. The results suggest that habitat fragmentation threatens the coexistence of stream-dwelling charr and salmon at both the basin and stream scales. We believe that exclusion of one species by another is likely in extremely fragmented habitats with minimal gradients and little variation in physical conditions (through reduced stream gradient and increased sand sedimentation caused by damming). In addition, multiple sites of damming ensure that there are no salmonid refuges from the collapse of metapopulation structure. In such fragmented habitats, even small tributaries serve important roles, as they are used mainly by salmonid fry and juveniles. We propose that habitats of native salmonids should be maximized by reconnecting fragmented habitats as part of a broader management plan.     

Predictability of multispecies competitive interactions in three populations of Atlantic salmon Salmo salar

Juvenile Atlantic salmon Salmo salar from three allopatric populations (LaHave, Sebago and Saint‐Jean) were placed into artificial streams with combinations of four non‐native salmonids: brown trout Salmo trutta, rainbow trout Oncorhynchus mykiss, Chinook salmon Oncorhynchus tshawytscha and coho salmon Oncorhynchus kisutch. Non‐additive effects, as evidenced by lower performance than predicted from weighted summed two‐species competition trials, were detected for S. salar fork length (L_F) and mass, but not for survival, condition factor or riffle use. These data support emerging theory on niche overlap and species richness as factors that can lead to non‐additive competition effects.     

The most variable vertebrate on Earth

In the search for the most variable non-human vertebrate on Earth, intraspecific variation of ten variable traits was compared among ten highly variable species. Mammals, birds and many reptiles, amphibians and fishes were excluded because most of the variation is among, and not within species. The focus was on northern fishes, where high intraspecific variation is well documented. The ten selected species were European whitefish Coregonus lavaretus, chinook salmon Oncorhyncus tshawytscha, sockeye salmon O. nerka, rainbow trout O. mykiss, atlantic salmon Salmo salar, brown trout S. trutta, arctic charr Salvelinus alpinus, brook charr S. fontinalis, dolly varden charr S. malma and threespine stickleback Gasterosteus aculeatus. Variation included not only size and phenotype, but also ecology, behaviour and life history. The traits were geographic range, migration, habitat, adult size, colour, body form, polymorphism, diet, reproduction and genetics. Arctic charr came on top in the final ranking, followed by dolly varden charr and rainbow trout. The two least variable were chinook salmon and threespine stickleback. It is proposed that arctic charr, which is also the northernmost fish on Earth, has evolved its unique variability in range, size, phenotype, ecology and life history by adapting to the extreme and highly unpredictable ecological conditions of arctic and other northern lakes for many glacial periods.     

Food habit divergence between white-spotted charr and masu salmon in Japanese mountain streams: circumstantial evidence for competition

 The diet and foraging microhabitat of white-spotted charr, Salvelinus leucomaenis, were compared between mountain stream reaches where it occurs with (sympatric) and without (allopatric) masu salmon, Oncorhynchus masou masou, a potential competitor, to examine the evidence for interspecific competition between these fish, which commonly co-occur in Japanese mountain streams. In three streams examined, the similarity between the diets of allopatric charr and salmon was much greater than that between the diets of sympatric charr and salmon. Both allopatric charr and sympatric salmon intensively utilized terrestrial invertebrates among stream drifts (52%–65% of the diet), whereas this prey category made up only an intermediate portion of sympatric charr diets (11%–29%). Examination of available prey composition in stream drifts showed that the consumption of terrestrial invertebrates by allopatric charr and sympatric salmon was approximately twice as much as that by sympatric charr. The presence of salmon, a potential competitor, may alter the diet of charr in the sympatric reaches. Charr holding focal points closer to the streambed were considered less efficient than sympatric salmon in their utilization of terrestrial invertebrates drifting primarily on the stream surface, although the foraging microhabitat of the charr was not influenced by the salmon. The mechanisms responsible for the dietary divergence between sympatric charr and salmon are probably the consequence of scramble competition over terrestrial invertebrates drifting on the stream surface. Received: January 21, 2002 / Accepted: November 19, 2002 Acknowledgments We thank Y. Tokuda, T. Takasu, Y. Kaneda, H. Jyoya, and H. Aoe for their assistance. This work was partly supported by funding through the Takara Harmonist Fund by the Takara Syuzo Co. Ltd. and the Japan Ministry of Education, Science, Sport and Culture (grants 09NP1501 and 11440224). Correspondence to:H. Miyasaka     

Occurrence of a hybrid between endemic Miyabe charr Salvelinus malma miyabei and introduced masu salmon Oncorhynchus masou masou in the Lake Shikaribetsu system, Hokkaido, Japan

A hybrid specimen between endemic Miyabe charr Salvelinus malma miyabei and introduced masu salmon Oncorhynchus masou masou was collected in an inlet stream (Yamada Creek) of the Lake Shikaribetsu. This specimen showed intermediate external characteristics between two species and was also confirmed as a hybrid by DNA markers. The mtDNA of the specimen was identical with that of S. malma miyabei, suggesting that the mating occurred between a female Miyabe charr and a male masu salmon. Hybridization with introduced masu salmon may cause deleterious effects on S. malma miyabei.     

Discrimination between farmed and free-living invasive salmonids in Chilean Patagonia using stable isotope analysis

In Chilean Patagonia relatively pristine aquatic environments are being modified by the introduction of exotic salmonids, initially through their deliberate release for sport fishing since the early twentieth century, and more recently via the accidental escape from fish farms. There is therefore a need to reliably distinguish between naturally reproducing and fugitive salmonids associated with the Chilean salmonid farming industry, the second largest in the world. We tested the ability of stable isotope analysis (SIA) and analysis of scale growth profiles to discriminate between farmed and free-living salmonids sampled around the Island of Chiloé. Juvenile Atlantic salmon (Salmo salar) and rainbow trout (Oncorhynchus mykiss) from aquaculture facilities were significantly more enriched in δ¹⁵N and lipid-corrected δ¹³C than river-caught individuals. Scale growth slopes during the first year in freshwater were significantly higher in farmed than in wild-caught rainbow trout, indicating faster somatic growth under hatchery conditions. Stable isotopes analysis classified 94% of juvenile Atlantic salmon and rainbow trout to their correct farm or free-living groups. Our results, therefore, can help to elucidate the origin and spread of exotic invasive salmonids in Chile, and address one of the biggest threats to native freshwater fishes in Patagonia and other temperate zones of the Southern Hemisphere.     

The effects of fisheries management on harvest rates of native and non-native salmonid fish species

In central Europe, both brown trout Salmo trutta and European grayling Thymallus thymallus are threatened native salmonid species with high value in recreational angling and nature conservation. On the other hand, rainbow trout Oncorhynchus mykiss and brook trout Salvelinus fontinalis are intensively stocked non-native species of high angling value but no value for nature conservation. This study tested if harvest rates of native salmonids are negatively correlated to intensive stocking and harvest rates of non-native salmonids in inland freshwater recreational fisheries. Data were collected from 250 fishing sites (river and stream stretches) over 13 years using mandatory angling logbooks. Logbooks were collected from individual anglers by the Czech Fishing Union in the regions of Prague and Central Bohemia, Czechia (central Europe) and processed by the author of this study. In result, anglers harvested 200,000 salmonids with total weight of 80 tons over 13 years. Intensive stocking of multiple salmonid species lead to slightly lower harvests of native salmonids. Inversely, intensive harvests of multiple salmonid species lead to slightly higher harvest of native salmonids. Recapture rates of stocked salmonids were relatively low (0.6%–3.7%), proving fish stocking moderately ineffective. Since the effects of non-native salmonid stocking and harvest rates on native salmonid harvest were significant but not strong, it is suggested that rivers and streams that support fishing for non-native salmonids still support fishing for native salmonids. However, this idea does not apply for fishing sites with really high intensity of non-native salmonid stocking – harvest rates of natives were very low on these fishing sites.     

The diversity of juvenile salmonids does not affect their competitive impact on a native galaxiid

We used an invaded stream fish community in southern Chile to experimentally test whether the diversity of exotic species affects their competitive impact on a native species. In artificial enclosures an established invasive, rainbow trout, Oncorhynchus mykiss, and a potential invader, Atlantic salmon, Salmo salar, reduced the growth rate of native peladilla, Aplochiton zebra, by the same amount. In enclosures with both exotic salmonids, the growth rates of all three species were the same as in single exotic treatments. While neither species identity nor diversity appeared to affect competitive interactions in this experiment, the impact of salmonid diversity may vary with the type of interspecific interaction and/or the species identity of the exotics. Our experiment links two prominent concepts in invasion biology by testing whether the result of invasional meltdown, an increase in the diversity of exotic species, affects their impact through interspecific competition, the mechanism invoked by the biotic resistance hypothesis.     

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.     

Genomic arrangement of salinity tolerance QTLs in salmonids: A comparative analysis of Atlantic salmon (Salmo salar) with Arctic charr (Salvelinus alpinus) and rainbow trout (

ABSTRACT: BACKGROUND: Quantitative trait locus (QTL) studies show that variation in salinity tolerance in Arctic charr and rainbow trout has a genetic basis, even though both these species have low to moderate salinity tolerance capacities. QTL were observed to localize to homologous linkage group segments within putative chromosomal regions possessing multiple candidate genes. We compared salinity tolerance QTL in rainbow trout and Arctic charr to those detected in a higher salinity tolerant species, Atlantic salmon. The highly derived karyotype of Atlantic salmon allows for the assessment of whether disparity in salinity tolerance in salmonids is associated with differences in genetic architecture. To facilitate these comparisons, we examined the genomic synteny patterns of key candidate genes in the other model teleost fishes that have experienced three whole-genome duplication (3R) events which preceded a fourth (4R) whole genome duplication event common to all salmonid species. RESULTS: Nine linkage groups contained chromosome-wide significant QTL (AS-2, -4p, -4q, -5, -9, -12p, -12q, -14q -17q, -22, and [MINUS SIGN]23), while a single genome-wide significant QTL was located on AS-4q. Salmonid genomes shared the greatest marker homology with the genome of three-spined stickleback. All linkage group arms in Atlantic salmon were syntenic with at least one stickleback chromosome, while 18 arms had multiple affinities. Arm fusions in Atlantic salmon were often between multiple regions bearing salinity tolerance QTL. Nine linkage groups in Arctic charr and six linkage group arms in rainbow trout currently have no synteny alignments with stickleback chromosomes, while eight rainbow trout linkage group arms were syntenic with multiple stickleback chromosomes. Rearrangements in the stickleback lineage involving fusions of ancestral arm segments could account for the 21 chromosome pairs observed in the stickleback karyotype. CONCLUSIONS: Salinity tolerance in salmonids from three genera is to some extent controlled by the same loci. Synteny between QTL in salmonids and candidate genes in stickleback suggests genetic variation at candidate gene loci could affect salinity tolerance in all three salmonids investigated. Candidate genes often occur in pairs on chromosomes, and synteny patterns indicate these pairs are generally conserved in 2R, 3R, and 4R genomes. Synteny maps also suggest that the Atlantic salmon genome contains three larger syntenic combinations of candidate genes that are not evident in any of the other 2R, 3R, or 4R genomes examined. These larger synteny tracts appear to have resulted from ancestral arm fusions that occurred in the Atlantic salmon ancestor. We hypothesize that the superior hypo-osmoregulatory efficiency that is characteristic of Atlantic salmon may be related to these clusters.     

Marine food and diet overlap of co-occurring Arctic charr Salvelinus alpinus (L.), brown trout Salmo trutta L. and Atlantic salmon S. salar L. off Senja,N. Norway

Summary Stomach contents analyses and other biological information of Arctic charr (Savelinus alpinus (L.)), brown trout (Salmo trutta L.) and small Atlantic salmon (S. salar L.) caught 1982–85 close to the Åelv estuary (69°N) on the island of Senja, N. Norway are presented, and extracts of a 1975–85 fishing log given. this appears to be the first case study of the feeding habits of all three European anadromous salmonids in marine sympatry, and also one of very few reports on the marine food of the Arctic charr from Europe. The general feeding habits of the charr were similar to that found in N. Canada. Pelagic fish (herring, sand-eel) seem to be preferred. Plankton (crab megalopae, krill) and hyperbenthos (amphipods, mysids) are also taken, especially when suitable fish are scarce. In 1985 high herring densities provided superabundant food, and diet overlap between charr, trout and salmon was high. Salmonid nursery rivers are abundant in N. Norway and during summer the three species coexist in a near-shore, surface-oriented pelagic guild of fishes. The salmon seems to be a relatively specialized piscivore, while the trout takes a wider range of fish and also invertebrate prey. The charr probably is the most euryphagous of the three, being able to exploit the more marginal parts of the prey resources of their common habitat.     

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.     

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.