Comparison of competitive ability between native and introduced salmonids: evidence from pairwise contests

Brown trout, Salmo trutta, and rainbow trout, Oncorhynchus mykiss, have been introduced to freshwaters in Hokkaido, Japan. Today, it is recognized that these introduced salmonids have negative impacts on native salmonids such as white-spotted charr, Salvelinus leucomaenis, and masu salmon, O. masou. In particular, interspecific competition may be an important mechanism that could contribute to the exclusion for native salmonids. In this study, experimental pairwise contests were conducted to compare interference competitive ability between native and introduced salmonids. We demonstrated that brown trout were competitively superior to white-spotted charr and masu salmon whereas rainbow trout were superior to white-spotted charr. We suggest that introduced brown trout negatively impact both white-spotted charr and masu salmon, and introduced rainbow trout negatively impact white-spotted charr.     

Invading rainbow trout usurp a terrestrial prey subsidy from native charr and reduce their growth and abundance

Movements of prey organisms across ecosystem boundaries often subsidize consumer populations in adjacent habitats. Human disturbances such as habitat degradation or non-native species invasions may alter the characteristics or fate of these prey subsidies, but few studies have measured the direct effects of this disruption on the growth and local abundance of predators in recipient habitats. Here we present evidence, obtained from a combined experimental and comparative study in northern Japan, that an invading stream fish usurped the flux of allochthonous prey to a native fish, consequently altering the diet and reducing the growth and abundance of the native species. A large-scale field experiment showed that excluding terrestrial invertebrates that fell into the stream with a mesh greenhouse reduced terrestrial prey in diets of native Dolly Varden charr (Salvelinus malma) by 46–70%, and reduced their growth by 25% over six weeks. However, when nonnative rainbow trout (Oncorhynchus mykiss) were introduced, they monopolized these prey and caused an even greater reduction of terrestrial prey in charr diets of 82–93%, and reduced charr growth by 31% over the same period. Adding both greenhouse and rainbow trout treatments together produced similar results to adding either alone. Results from a comparative field study of six other stream sites in the region corroborated the experimental findings, showing that at invaded sites rainbow trout usurped the terrestrial prey subsidy, causing a more than 75% decrease in the biomass of terrestrial invertebrates in Dolly Varden diets and forcing them to shift their foraging to insects on the stream bottom. Moreover, at sites with even low densities of rainbow trout, biomass of Dolly Varden was more than 75% lower than at sites without rainbow trout. Disruption of resource fluxes between habitats may be a common, but unidentified, consequence of invasions, and an additional mechanism contributing to the loss of native species Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

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.     

Asymmetric competition drives lake use of coexisting salmonids

To what degree are population differences in resource use caused by competition and the occupation of adjacent positions along environmental gradients evidence of competition? Habitat use may be the result of a competitive lottery, or restricted by competition. We tested to what extent population differences in habitat use of two salmonids, cutthroat trout (Oncorhynchus clarki) and Dolly Varden charr (Salvelinus malma) were influenced by interspecific competition. We hypothesized that the depth distribution of Dolly Varden charr would be affected by competition from the more littoral and surface-oriented cutthroat trout, and that the depth distribution of cutthroat trout would be little affected by competition from Dolly Varden charr. Sympatric populations of cutthroat trout and Dolly Varden charr were created by reciprocal transfers of previously allopatric populations in two experimental lakes. We found evidence of asymmetric competition, as Dolly Varden charr were displaced from littoral habitats when sympatric with cutthroat trout, whereas cutthroat trout remained unaffected by the presence of Dolly Varden charr. Evolved differences between the species, and differences between experimental lakes, also contributed to population differences in habitat use, but asymmetric competition remained as the main driver of different depth distributions in sympatry.     

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.     

Potential fragmentation and loss of thermal habitats for charrs in the Japanese archipelago due to climatic warming

1. The upper thermal limits of the present distributions of two charr species, Dolly Varden, Salvelinus malma , and white-spotted charr, S. leucomaenis , in streams of the Japanese archipelago were examined using groundwater temperature as an index of thermal condition. The lower limits of the altitudinal distributions of Dolly Varden and white-spotted charr were delineated, respectively, by 8 and 16 °C groundwater isotherms.
2. The potential impact of future climatic warming on the geographical distribution, habitat extent and population fragmentation of each species was predicted at both the full archipelago and individual catchment levels.
3. For Dolly Varden, analysis at the full archipelago level indicated a loss of 27.6, 67.2, 79.6 and 89.6% of the current geographical range, respectively, for a 1, 2, 3 and 4 °C increase in mean annual air temperature. The present distribution area of white-spotted charr would likewise reduce by 4.1, 20.5, 33.8 and 45.6%, respectively.
4. Based on the analyses of three individual catchments, one for Dolly Varden and two for white-spotted charr, the lower habitat boundaries for the two charr species could be expected to rise increasingly to higher elevations in each catchment as warming proceeded. As a consequence, there would be large reductions in mean habitat area, with increasing habitat fragmentation followed by localized extinctions of the two species.     

Potential fragmentation and loss of thermal habitats for charrs in the Japanese archipelago due to climatic warming

1. The upper thermal limits of the present distributions of two charr species, Dolly Varden, Salvelinus malma , and white-spotted charr, S. leucomaenis , in streams of the Japanese archipelago were examined using groundwater temperature as an index of thermal condition. The lower limits of the altitudinal distributions of Dolly Varden and white-spotted charr were delineated, respectively, by 8 and 16 °C groundwater isotherms.
2. The potential impact of future climatic warming on the geographical distribution, habitat extent and population fragmentation of each species was predicted at both the full archipelago and individual catchment levels.
3. For Dolly Varden, analysis at the full archipelago level indicated a loss of 27.6, 67.2, 79.6 and 89.6% of the current geographical range, respectively, for a 1, 2, 3 and 4 °C increase in mean annual air temperature. The present distribution area of white-spotted charr would likewise reduce by 4.1, 20.5, 33.8 and 45.6%, respectively.
4. Based on the analyses of three individual catchments, one for Dolly Varden and two for white-spotted charr, the lower habitat boundaries for the two charr species could be expected to rise increasingly to higher elevations in each catchment as warming proceeded. As a consequence, there would be large reductions in mean habitat area, with increasing habitat fragmentation followed by localized extinctions of the two species.     

Genetic Differentiation of Three Sympatric Charr Species from the Genus SalvelinusInferred from PCR–RFLP Analysis of Mitochondrial DNA

We examined differentiation of three sympatric charr species Dolly Varden charr Salvelinus malmaWalbaum, white-spotted charrS.leucomaenisPallas, and Levanidov charrS. levanidovi Chereshnev, Scopetz, Gudkov. Using restriction fragment length polymorphism analysis (RFLP), three mitochondrial DNA fragments (ND1/ND2, ND5/ND6, and Cytb/D-loop) amplified in PCR were compared. The divergence of mtDNA nucleotide sequences betweenS. levanidovi and S. leucomaenis was about 9.7%, between S. levanidovi and S. malma, 8.7%, and between S. malma andS. laecomaenis, 7.5%. These results indicate genetic isolation of the species examined and support the earlier suggestion on closeness of Levanidov charr to the common ancestor of the genus Salvelinus.     

Distribution of two congeneric charrs in streams of Hokkaido Island,Japan: considering multiple factors across scales

Salvelinus leucomaenis (white-spotted charr) and S. malma (Dolly Varden) are distributed throughout Hokkaido Island, Japan, but sites where they occur in sympatry are rare. In general, S. malma inhabit upstream reaches and S. leucomaenis extend downstream to the ocean. Factors influencing their distribution were analyzed at four spatial scales ranging from the whole island to individual stream pools. At the island scale, S. leucomaenis were found in the warmer south-west region and at lower altitudes elsewhere, whereas S. malma were found in the colder north-east and at higher altitudes. At a regional scale, the downstream limit of S. malma and upstream limit of S. leucomaenis shifted to lower altitude from south-west to north-east across the island, coincident with the decrease in temperature. Further analysis showed that transition points from S. leucomaenis or sympatry to S. malma in individual watersheds were closely related to an index of cumulative mean monthly temperatures exceeding 5°C. However, at the scale of a single watershed, the transition occurred at different altitudes, gradients, and temperatures in two tributaries, apparently because stream discharge, habitat, and disturbances from floods interacted with these abiotic factors to limit distribution. The two charr species developed interspecific dominance hierarchies in individual pools, and there was strong complementary density compensation among stream pools that could be explained by interspecific competition but not by differences in habitat. However, patterns at watershed and regional scales suggested that interspecific competition interacts with temperature in complex ways. We conclude that the importance of various abiotic and biotic factors in shaping Hokkaido charr distributions depends on the scale at which they are viewed.     

Temperature‐dependent competition between juvenile salmonids in small streams

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Intra- and interspecific dominance hierarchies and variation in foraging tactics of two species of stream-dwelling chars

Aggressive interactions, foraging behavior, habitat use and diet were studied in sympatric populations of white-sported char,Salvelinus leucomaenis, and Dolly Varden,Salvelinus malma, in a Japanese mountain stream. Underwater observations on individuals of both species revealed two distinct behavioral regimes: aggressive drift foragers and non-aggressive benthos foragers. Aggressive drift foragers defended partial territories around focal points from which they made forays to capture invertebrates drifting in the water column. Non-aggressive benthos foragers cruised around and beneath cobble in large foraging ranges that overlapped each other. Intra- and interspecific, size-dependent dominance hierarchies were recognized among aggressive drift foragers, whereas non-aggressive benthos foragers showed no such relationships. Terrestrial invertebrates were the most abundant prey in the diets of drift foragers, whereas a very small proportion of the diet of benthos foragers was made up of these taxa. Benthos foragers showed more complex diet composition than drift foragers. These results suggest that non-aggressive benthos foragers may avoid not only interference but also exploitative competition by using alternative foraging tactics. The proportion of drift foragers to benthos foragers among white-spotted char was more than 35 times that among Dolly Varden. The significant difference in the proportion of each species using the two types of foraging strategy results in interspecific food segregation in sympatric populations.     

Introgressive hybridization between Dolly Varden Salvelinus malma and white‐spotted charr Salvelinus leucomaenis on Hokkaido Island, Japan

Intra‐ and interspecific phylogenetic analysis of Dolly Varden Salvelinus malma and white‐spotted charr Salvelinus leucomaenis throughout Hokkaido Island was conducted using nucleotide sequences of the mitochondrial DNA control region and three nuclear DNA markers [ribosomal DNA (ITS‐1), microsatellite ( u ‐85), SINE ( Fok ‐223)]. A total of 19 mtDNA haplotypes from 271 individuals of Dolly Varden, nine haplotypes from 134 white‐spotted charr were defined, and identified two well‐supported monophyletic clades for each species. Two haplotypes of Dolly Varden, however, were clustered together in the white‐spotted charr clade. Those haplotypes were found only from several rivers in the Shiretoko Peninsula, and a neighbouring river of that region. Analyses of ribosomal DNA and Fok ‐223 loci revealed that both species are characterized by having completely species‐specific diagnostic sequence and fragment patterns. Dolly Varden of the Shiretoko populations are typically allopatric in distribution where white‐spotted charr do not currently occur. In addition, incongruence in genetic relationships between mtDNA and nuclear DNA markers give strong evidence of historical mtDNA introgression between Dolly Varden and white‐spotted charr. Some white‐spotted charr diagnostic alleles in a nuclear microsatellite locus ( u ‐85) were found in some Shiretoko Dolly Varden populations, suggesting that introgressive hybridization might have also occurred in relatively recent contact and potentially ongoing evolutionary event. The present study presents an example of historical hybridization and introgression at the southernmost distribution limits of Dolly Varden.     

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.     

Effects of forestry on the thermal habitat of Dolly Varden (<Emphasis Type="Italic">Salvelinus malma</Emphasis>)

Forestry activities in riparian areas are known to affect stream communities considerably. Not only do riparian deforestation resulting from agriculture or urbanization developments affect stream communities but extensive commercial plantation and forestry practices can alter stream environments adjacent to remaining, intact or secondary forests. Because forestry often includes the construction of logging roads through the riparian zone, this can directly degrade stream environments. Twelve streams in the Shiretoko Peninsula, Hokkaido were investigated so as to determine the effects of forestry practices on stream temperature, periphyton biomass, grazer (benthic invertebrates) biomass and Dolly Varden (Salvelinus malma Walbaum) biomass. The greater the proportion of planted area in the catchment, the higher the stream temperature. Stream temperature directly affects periphyton biomass and Dolly Varden biomass negatively. Neither stream temperature nor periphyton biomass predicted grazer biomass, whereas a positive correlation was found between grazer biomass and Dolly Varden biomass that forage on invertebrates. The overall results indicated that Dolly Varden in the Shiretoko streams were negatively affected by forestry practices and the resultant stream temperature increases. Without effective future riparian forest management, the complex effects of both riparian disturbance and ongoing global warming could further reduce Dolly Varden populations in the region.     

Karyological Differentiation of Northern Dolly Varden and Sympatric Chars of the Genus <Emphasis Type="Italic">Salvelinus</Emphasis> in Northeastern Russia

In many cases the taxonomic status of sympatric chars (with exception of white-spotted char Salvelinus leucomaenis) is not clear and is actively debated. We karyotyped three pairs of sympatric chars inhabiting the Russian Far East-northern Dolly Varden, S. malma malma, and Lavanidov's char, S. levanidovi, from the Yama River (northern coast of the Sea of Okhotsk, Magadan Region), northern Dolly Varden and Taranetz's char, S. taranetzi, from Lake Achchen (east Chukotka), and northern Dolly Varden and white char, S. albus, from the Kamchatka River (Kamchatka Peninsula). Three of them had similar chromosome numbers. But all chars studied had an individual and discrete set of karyotypic characters, which enabled reliable identification each of them by chromosome number, chromosome arm number, and number and location of active nuclear organizer regions.     

Metapopulation structure of stream-dwelling Dolly Varden charr inferred from patterns of occurrence in the Sorachi River basin, Hokkaido, Japan

1. We investigated the occurrence (presence or absence) of stream‐dwelling Dolly Varden (Salmonidae: Salvelinus malma) in the 78 tributaries of the Sorachi River system, Hokkaido, Japan to examine evidence for metapopulation structure. 2. We hypothesised (i) that the occurrence of Dolly Varden would be less likely in small tributaries because of the higher extinction risk and (ii) that occurrence would be less likely in isolated tributaries because of reduced rates of dispersal and colonisation. In addition to these two general predictions, we hypothesised (iii) that the occurrence would be more likely in spring‐fed tributaries, because Dolly Varden prefer cold water as present in these systems. 3. When all the tributaries were included in a multiple logistic regression analysis, habitat size and habitat type (spring‐fed versus non‐spring‐fed tributaries) were selected for the best model, whereas isolation was not important. However, when tributaries in the lower Sorachi basin (an area likely influenced by human activities) were excluded from the analysis, the effect of isolation became significant. 4. Our results indicate there is a potential metapopulation structure of the Dolly Varden in the Sorachi River basin. Furthermore, it is possible that metapopulation dynamics follow a clinal pattern. In the upper and middle reaches of the Sorachi River basin, an equilibrium between extinction and colonisation of Dolly Varden populations is a possibility. In the lower portion of the basin, isolation appeared to be less important, and extinction rate may be driving patterns of occurrence toward systematic extinction of local populations. 5. Our results illustrate the possibility of spatial variation in the processes influencing occurrence of Dolly Varden within a single river basin, and the potential influences of habitat alteration on extinction and colonisation processes.     

Genetic differentiation of three sympatric charr species from genus Salvelinus inferred from PCR-RFLP analysis of mitochondrial DNA

We examined differentiation of three sympatric charr species Dolly Varden charr Salvelinus malma Walbaum, white-spotted charr S. leucomaenis Pallas, and Levanidov charr S. levanidovi Chereshnev, Scopetz, Gudkov. Using restriction fragment length polymorphism analysis (RFLP), three mitochondrial DNA fragments (ND1/ND2, ND5/ND6, and Cytb/D-loop) amplified in PCR were compared. The divergence of mtDNA nucleotide sequences between S. levanidovi and S. leucomaenis was about 9.7%, between S. levanidovi and S. malma, 8.7%, and between S. malma and S. laecomaenis, 7.5%. These results indicate genetic isolation of the species examined and support the earlier suggestion on closeness of Levanidov charr to the common ancestor of the genus Salvelinus.     

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     

Habitat use by Nonnative Rainbow Trout, Oncorhynchus mykiss, and Native Little Colorado spinedace, Lepidomeda vittata

We evaluated overlap in microhabitat use between nonnative rainbow trout, Oncorhynchus mykiss, and native Little Colorado spinedace, Lepidomeda vittata, a federally threatened cyprinid, in natural and experimental settings. In natural settings, we also examined occurrence and microhabitat use of two other native fishes, speckled dace, Rhinichthys osculus, and bluehead sucker, Catostomus discobolus. Native species co-occurred, as did rainbow trout and bluehead sucker. However, occurrences of Little Colorado spinedace and speckled dace were not significantly correlated with occurrence of rainbow trout. Total lengths of all three native species were significantly smaller at allopatric sites than at sites sympatric with rainbow trout. Microhabitat characteristics at sites with rainbow trout did not differ from those where the other three species were found, but did differ among the native species. In laboratory experiments with Little Colorado spinedace and rainbow trout, rainbow trout used the lower depth zone most, and spinedace increased use of the lower depth zone upon addition of rainbow trout. In addition, species tended to co-occur in zones, but used cover independently of one-another, suggesting a low level of agonistic interactions. However, after addition of a high density of rainbow trout, spinedace tended to use cover less than before. We suggest that the species can coexist at low rainbow trout densities. Potential negative effects of rainbow trout on Little Colorado spinedace likely increase with increasing densities of rainbow trout, and rainbow trout likely affect smaller size classes of Little Colorado spinedace more than larger ones.     

Ongoing localized extinctions of stream-dwelling white-spotted charr populations in small dammed-off habitats of Hokkaido Island,Japan

Habitat fragmentation caused by damming can greatly reduce the population viability of aquatic organisms, with smaller fragmented populations at higher risk of extinction due to increased demographic, genetic, and environmental stochasticity. However, empirical evidence demonstrating that smaller natural populations are more vulnerable to extinction is limited. We studied the vulnerability to extinction of white-spotted charr (Salvelinus leucomaenis) populations in 30 dammed-off streams in Oshima Peninsula, southwestern Hokkaido Island, Japan, by comparing the incidence of charr populations in streams between 1999 and 2014. Using electrofishing and environmental DNA surveys, we identified three localized extinctions, with the probability of extinction increasing with decreasing watershed area (our surrogate for habitat size). We also found a new population in one dammed-off stream in which white-spotted charr were previously unknown, after installation of a fish ladder, indicating the capacity of white-spotted charr to recolonize reconnected habitat in a short period. Our results suggest that localized extinction of white-spotted charr in small dammed-off streams is ongoing, but that appropriate fish migration corridors can reduce localized extinction risk and increase the probability of species persistence.