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.     

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.     

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.     

Spatial covariation of fish population vital rates in a stream network

Animal populations are spatially structured in heterogeneous landscapes, in which local patches with differing vital rates are connected by dispersal of individuals to varying degrees. Although there is evidence that vital rates differ among local populations, much less is understood about how vital rates covary among local patches in spatially heterogeneous landscapes. In this study, we conducted a nine-year annual mark–recapture survey to characterize spatial covariation of survival and growth for two Japanese native salmonids, white-spotted charr Salvelinus leucomaenis japonicus and red-spotted masu salmon Oncorhynchus masou ishikawae, in a headwater stream network composed of distinctly different tributary and mainstem habitats. Spatial structure of survival and growth differed by species and age class, but results provided support for negative covariation between vital rates, where survival was higher in the tributary habitat but growth was higher in the mainstem habitat. Thus, neither habitat was apparently more important than the other, and local habitats with complementary vital rates may make this spatially structured population less vulnerable to environmental change (i.e. portfolio effect). Despite the spatial structure of vital rates and possibilities that fish can exploit spatially distributed resources, movement of fish was limited due partly to a series of low-head dams that prevented upstream movement of fish in the study area. This study shows that spatial structure of vital rates can be complex and depend on species and age class, and this knowledge is likely paramount to elucidating dynamics of spatially structured populations.     

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.     

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.

     

Genetic structure of a native cyprinid in a reservoir‐altered stream network

1. Reservoirs modify riverine ecosystems worldwide, and often with deleterious impacts on native biota. The immediate effects of reservoirs on native fish species below dams and in impounded reaches have received considerable attention, but it is unclear how reservoirs may affect fish species at larger spatial and temporal scales. Documented declines of stream fish populations in direct tributaries of reservoirs suggest reservoir pools may reduce gene flow among historically connected populations. 2. Because of increased predator densities in reservoirs and the extent of habitat alteration in impounded reaches, I predicted reservoir habitats would reduce gene flow among small‐bodied fish populations separated by reservoir habitat. I used microsatellite markers to assess the spatial genetic structure of populations of the red shiner (Cyprinella lutrensis), in a reservoir‐fragmented stream network (Lake Texoma, U.S.A.). I also tested the prediction that populations in two direct tributaries that have experienced population declines would have low genetic diversity. Individuals were collected from six sites upstream of the reservoir, three sites in the reservoir and three sites in direct tributaries of the reservoir during 2008 and 2009. 3. Results indicate that most populations were isolated by distance with little divergence among populations. In one direct tributary population, however, there was substantial genetic divergence, and genetic diversity was significantly lower than in other populations. Gene flow also seemed to be lower in reservoir habitats than in intact stream habitats, suggesting reservoir habitats may be reducing gene flow among the reservoir‐separated populations. These results indicate that reservoirs may reduce gene flow among reservoir‐fragmented stream fish populations, altering the evolutionary trajectories of fragmented populations.     

Application of a bioenergetics model to estimate the influence of habitat degradation by check dams and potential recovery of masu salmon populations

Using a bioenergetics model, we examined how check dams negatively effect masu salmon (Oncorhynchus masou) populations by causing habitat loss in upstream areas and habitat degradation in downstream areas. The potential recovery of masu salmon populations in the upstream area was estimated based on the expected biomass and potential recovery area. We also determined if and how fish carrying capacity is affected by degradation of substrate conditions (armoring and compaction) in the downstream area. Recovery of upstream areas was considered to be effective in enhancing and conserving masu salmon populations. We demonstrated that the dam-induced altered substrate conditions and habitat degradation in the downstream area resulted in a considerable reduction of drifting prey. Simulation analysis revealed that a 40 % increase in the abundance of masu salmon juveniles in the downstream area could be expected if substrate conditions were restored. We concluded that both improvement of migration barriers and restoring the sediment regime would be important in enhancing and conserving wild masu salmon populations.     

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.     

Non‐native salmonids affect amphibian occupancy at multiple spatial scales

Aim The introduction of non‐native species into aquatic environments has been linked with local extinctions and altered distributions of native species. We investigated the effect of non‐native salmonids on the occupancy of two native amphibians, the long‐toed salamander (Ambystoma macrodactylum) and Columbia spotted frog (Rana luteiventris), across three spatial scales: water bodies, small catchments and large catchments. Location Mountain lakes at ≥ 1500 m elevation were surveyed across the northern Rocky Mountains, USA. Methods We surveyed 2267 water bodies for amphibian occupancy (based on evidence of reproduction) and fish presence between 1986 and 2002 and modelled the probability of amphibian occupancy at each spatial scale in relation to habitat availability and quality and fish presence. Results After accounting for habitat features, we estimated that A. macrodactylum was 2.3 times more likely to breed in fishless water bodies than in water bodies with fish. Ambystoma macrodactylum also was more likely to occupy small catchments where none of the water bodies contained fish than in catchments where at least one water body contained fish. However, the probability of salamander occupancy in small catchments was also influenced by habitat availability (i.e. the number of water bodies within a catchment) and suitability of remaining fishless water bodies. We found no relationship between fish presence and salamander occupancy at the large‐catchment scale, probably because of increased habitat availability. In contrast to A. macrodactylum, we found no relationship between fish presence and R. luteiventris occupancy at any scale. Main conclusions Our results suggest that the negative effects of non‐native salmonids can extend beyond the boundaries of individual water bodies and increase A. macrodactylum extinction risk at landscape scales. We suspect that niche overlap between non‐native fish and A. macrodactylum at higher elevations in the northern Rocky Mountains may lead to extinction in catchments with limited suitable habitat.     

Hierarchical genetic structure of native masu salmon populations in Hokkaido,Japan

Identification of the spatial extent of genetic structuring that may be influenced by evolutionary, ecological and historical factors is critical for effective conservation or management strategies. Masu salmon Oncorhynchus masou is commonly distributed in Far East, however, many local populations have been under threats of decline due to habitat destruction, overexploitation, and genetic introgression. To reveal the spatial genetic structure of native masu salmon populations in Hokkaido, masu salmon samples were collected from 16 rivers in which there was no official record of artificial releases of any masu salmon stock and were analyzed using 15 microsatellite loci. A Bayesian assignment test revealed that masu salmon populations were divided into two genetically distinct groups: the northeastern and southwestern groups. For within-group genetic structure, all populations, except for geographically proximate populations, were significantly different from each other. AMOVA revealed that genetic variation at among-group level based on groups identified assignment test was greater than that of groups based on geographic locations. There was no significant IBD for the 16 populations. However, the Mantel test revealed significant IBD for the northeastern group, but did not for the southwestern group. This study suggested that native masu salmon populations in Hokkaido exhibit a hierarchical genetic structure that is largely a result of their precise homing behavior. The results of this study also highlight the importance of defining populations by using genetic data rather than by using predefined populations based on geographic locations for the correct determination of genetic structure.     

Complementary distribution of non-native white-spotted charr and native red-spotted masu salmon in Shikoku Island,southwestern Japan: a consequence of interspecific interactions?

Ichthyological Research - White-spotted charr (Salvelinus leucomaenis subspp.) and masu salmon (Oncorhynchus masou subspp.) are representative stream salmonids in the Japanese Archipelago. Although...     

Habitat use and fish activity of landlocked Atlantic salmon and brook charr in a newly developed habitat compensation facility

Degradation and destruction of valuable spawning and rearing habitat due to anthropogenic changes (e.g., flow modification and channelisation) is known to have dramatic impacts on fish populations. To compensate for habitat losses due to hydropower development, an artificial fluvial habitat channel (‘Compensation Creek’) was constructed in south-central Newfoundland, Canada. The creek was designed to include appropriate habitat features for the two dominant salmonid fish species, landlocked Atlantic salmon (Salmo salar L.) and brook charr (Salvenius fontinalis Mitchell). The study examines the habitat use of landlocked Atlantic salmon and brook charr in the Compensation Creek using electromyogram (EMG) radio telemetry. Ten landlocked Atlantic salmon and eight brook charr were captured and tagged with EMG transmitters. In laboratory swimming experiments, the EMG values were calibrated against swimming speed. Fish were then released in the Compensation Creek and tracked on a daily basis. The results show that (1) during residence in the creek, both species used preferentially the habitat features designed to match their rearing habitat preferences, and (2) swimming speed did not vary among habitat types for either species.     

Ecological assessment criteria for restoring anadromous salmonid habitat in Pacific Northwest estuaries

Restoration of estuarine habitats is essential for the conservation and recovery of depressed Pacific salmon populations. However, assessing the functions of recently restored habitat poses a number of problems because of the transitory occurrence of salmonids in any one location. We propose assessment criteria and metrics that are based on the habitat’s capacity, opportunity, and realized function to enhance survivability of juvenile salmon. Because of the paucity of data relating capacity and opportunity attributes to realized function (e.g. growth, consumption rate, survival), there continues to be a need for manipulative experiments to assess the developmental status of restoration sites. Such a self-monitoring approach of letting the fish diagnose the ecological state of restoration would effectively address the small-scale, site-specific assessment goals and criteria, but ignores the larger-scale issues relating to the ability of diverse salmon species and life histories to occupy estuarine habitat landscapes. If coastal restoration is going to contribute the recovery of anadromous salmonid populations, a landscape perspective is fundamental to restoration planning, implementation, and particularly assessment.     

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.     

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.     

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.     

Persistence at distributional edges: Columbia spotted frog habitat in the arid Great Basin,USA

A common challenge in the conservation of broadly distributed, yet imperiled species is understanding which factors facilitate persistence at distributional edges, locations where populations are often vulnerable to extirpation due to changes in climate, land use, or distributions of other species. For Columbia spotted frogs (Rana luteiventris) in the Great Basin (USA), a genetically distinct population segment of conservation concern, we approached this problem by examining (1) landscape‐scale habitat availability and distribution, (2) water body‐scale habitat associations, and (3) resource management‐identified threats to persistence. We found that areas with perennial aquatic habitat and suitable climate are extremely limited in the southern portion of the species’ range. Within these suitable areas, native and non‐native predators (trout and American bullfrogs [Lithobates catesbeianus]) are widespread and may further limit habitat availability in upper‐ and lower‐elevation areas, respectively. At the water body scale, spotted frog occupancy was associated with deeper sites containing abundant emergent vegetation and nontrout fish species. Streams with American beaver (Castor canadensis) frequently had these structural characteristics and were significantly more likely to be occupied than ponds, lakes, streams without beaver, or streams with inactive beaver ponds, highlighting the importance of active manipulation of stream environments by beaver. Native and non‐native trout reduced the likelihood of spotted frog occupancy, especially where emergent vegetation cover was sparse. Intensive livestock grazing, low aquatic connectivity, and ephemeral hydroperiods were also negatively associated with spotted frog occupancy. We conclude that persistence of this species at the arid end of its range has been largely facilitated by habitat stability (i.e., permanent hydroperiod), connectivity, predator‐free refugia, and a commensalistic interaction with an ecosystem engineer. Beaver‐induced changes to habitat quality, stability, and connectivity may increase spotted frog population resistance and resilience to seasonal drought, grazing, non‐native predators, and climate change, factors which threaten local or regional persistence.     

Recent local adaptation of sockeye salmon to glacial spawning habitats

Salmonids spawn in highly diverse habitats, exhibit strong genetic population structuring, and can quickly colonize newly created habitats with few founders. Spawning traits often differ among populations, but it is largely unknown if these differences are adaptive or due to genetic drift. To test if sockeye salmon (Oncorhynchus nerka) populations are adapted to glacial, beach, and tributary spawning habitats, we examined variation in heritable phenotypic traits associated with spawning in 13 populations of wild sockeye salmon in Lake Clark, Alaska. These populations were commonly founded between 100 and 400 hundred sockeye salmon generations ago and exhibit low genetic divergence at 11 microsatellite loci (F _ST < 0.024) that is uncorrelated with spawning habitat type. We found that mean P _ST (phenotypic divergence among populations) exceeded neutral F _ST for most phenotypic traits measured, indicating that phenotypic differences among populations could not be explained by genetic drift alone. Phenotypic divergence among populations was associated with spawning habitat differences, but not with neutral genetic divergence. For example, female body color was lighter and egg color was darker in glacial than non-glacial habitats. This may be due to reduced sexual selection for red spawning color in glacial habitats and an apparent trade-off in carotenoid allocation to body and egg color in females. Phenotypic plasticity is an unlikely source of phenotypic differences because Lake Clark sockeye salmon spend nearly all their lives in a common environment. Our data suggest that Lake Clark sockeye salmon populations are adapted to spawning in glacial, beach and tributary habitats and provide the first evidence of a glacial spawning ecotype in salmonids. Glacial spawning habitats are often young (i.e., <200 years old) and ephemeral. Thus, local adaptation of sockeye salmon to glacial habitats appears to have occurred recently.