Metapopulations and salmonids: a synthesis of life history patterns and empirical observations

Abstract – Metapopulation theory has attracted considerable interest with reference to the salmonids. There has been little empirical evidence, however, to guide the evaluation or application of metapopulation concepts. From knowledge of salmonid life histories and our own work with bull trout ( Salvelinus confluentus ), Lahontan cutthroat trout ( Oncorhynchus clarki henshawi ) and westslope cutthroat trout ( Oncorhynchus clarki lewisi ), we suggest that simple generalizations of salmonid metapopulations are inappropriate. Although spatial structuring and dispersal mechanisms are evident, the relevance of extinction and colonization processes are likely to vary with life history, species, scale, and landscape. Understanding dispersal, the role of suitable but unoccupied habitats, and the potential for extinction debts in non-equilibrium metapopulations are key issues. With regard to conservation of salmonids, we suggest that efforts to understand and conserve key processes likely to influence the persistence of populations or metapopulations will be more successful than efforts to design minimal habitat reserves based on metapopulation theory. ^NOTE     

A riverscape perspective of Pacific salmonids and aquatic habitats prior to large‐scale dam removal in the Elwha River,Washington, USA

Abstract Dam removal has been increasingly proposed as a river restoration technique. In 2011, two large hydroelectric dams will be removed from Washington State’s Elwha River. Ten anadromous fish populations are expected to recolonise historical habitats after dam removal. A key to understanding watershed recolonisation is the collection of spatially continuous information on fish and aquatic habitats. A riverscape approach with an emphasis on biological data has rarely been applied in mid‐sized, wilderness rivers, particularly in consecutive years prior to dam removal. Concurrent snorkel and habitat surveys were conducted from the headwaters to the mouth (rkm 65–0) of the Elwha River in 2007 and 2008. This riverscape approach characterised the spatial extent, assemblage structure and patterns of relative density of Pacific salmonids. The presence of dams influenced the longitudinal patterns of fish assemblages, and species richness was the highest downstream of the dams, where anadromous salmonids still have access. The percent composition of salmonids was similar in both years for rainbow trout, Oncorhynchus mykiss (Walbaum), coastal cutthroat trout, Oncorhynchus clarkii clarkii (Richardson) (89%; 88%), Chinook salmon, Oncorhynchus tshawytscha (Walbaum) (8%; 9%), and bull trout, Salvelinus confluentus (Suckley) (3% in both years). Spatial patterns of abundance for rainbow and cutthroat trout (r = 0.76) and bull trout (r = 0.70) were also consistent between years. Multivariate and univariate methods detected differences in habitat structure along the river profile caused by natural and anthropogenic factors. The riverscape view highlighted species‐specific biological hotspots and revealed that 60–69% of federally threatened bull trout occurred near or below the dams. Spatially continuous surveys will be vital in evaluating the effectiveness of upcoming dam removal projects at restoring anadromous salmonids.     

A case study of surrogate species in aquatic conservation planning

• 1. The use of surrogate species (i.e. keystones, indicators, umbrellas) has been advocated for the conservation of target taxa and communities.
• 2. A recent Habitat Conservation Plan, which provided conservation measures intended to protect multiple aquatic species of concern over a large area, established an important precedent for surrogate species in aquatic conservation pursuant to the US Endangered Species Act.
• 3. The Habitat Conservation Plan's application of federally threatened bull trout was evaluated as an umbrella species for westslope cutthroat trout, which is in decline but not listed under the Act. Approximately 75% of known westslope cutthroat trout strongholds are not captured within bull trout strongholds west of the continental divide. The Habitat Conservation Plan failed to evaluate the suitability of this umbrella species and consequently failed to cover important priority areas for westslope cutthroat trout conservation.
• 4. This case study highlights the feasibility and importance of formally validating assumptions of surrogate species utility in multi‐species conservation planning.

Copyright © 2004 John Wiley & Sons, Ltd.     

Stable isotopes and gut content show diet overlap among native and introduced piscivores in a large oligotrophic lake

Abstract – In past dietary studies kokanee Oncorhynchus nerka were prominent in the diet of Pend Oreille Lake's large piscivores: native bull trout Salvelinus confluentus, cutthroat trout O. clarki and northern pikeminnow Ptychocheilus oregonensis, and introduced lake trout S. namaycush and Kamloops rainbow trout O. mykiss gairdneri. However, kokanee have declined to 10–20% of their former abundance. We therefore initiated this study to understand current predation demands on kokanee and diet overlap among piscivores, using gut content samples and analysis of stable nitrogen (δ¹⁵N) and carbon (δ¹³C) isotopes from the lake's fish and invertebrate community. In gut content samples, kokanee were the main prey item of large [i.e., ≥400 mm total length (TL)] bull and lake trout; a conclusion that was affirmed by stable isotope analysis. Rainbow trout >500 mm TL consumed mostly kokanee, thus there was a high degree of diet overlap among large bull, lake and rainbow trout. Small (i.e., <400 mm TL) rainbow and cutthroat trout diets overlapped, and were composed mostly of littoral benthic invertebrates. However, gut content and stable isotope analysis did not accord for 400–500 mm TL rainbow trout, small lake trout, and large cutthroat trout. In these instances, a linear mixing model using stable isotope results predicted kokanee consumption for each species, but no kokanee were identified in rainbow or lake trout gut content. Gut content and stable isotope analysis of native northern pikeminnow indicated a diet of mostly littoral benthic invertebrates at smaller (100–150 mm TL) lengths, with kokanee becoming more prominent in the diet of individuals >300 mm TL. Percent of kokanee in the diet of northern pikeminnow has declined from a prior study; otherwise piscivore diets have apparently remained unchanged. In this study, judgments as to the feeding of some piscvores, based on gut content alone, would be tenuous because of small sample sizes, but stable isotope analysis provided an efficient means for confirming diets.     

Are native brook charr and introduced rainbow trout differentially adapted to upstream and downstream reaches?

Abstract— Due to species introductions, brook charr (Salvelinus fontinalis) and rainbow trout (Oncorhynchus mykiss) occur together in many North American streams and typically exhibit a pattern of distribution in which brook charr numerically dominate headwaters and rainbow trout dominate downstream reaches. It has been suggested that 1) the two species compete or 2) the two species do not compete because they are differentially adapted to environmental conditions found in upstream and downstream zones. We assessed whether there were differences in growth and macrohabitat (pool, run and riffle) selection of brook charr and rainbow trout in upper, middle and lower stream zones of a small Pennsylvania stream. Brook charr and rainbow trout placed in replicate paired enclosures set in upstream and downstream reaches showed no significant differences in growth and survival rates upstream, but brook charr had significantly greater growth rates than rainbow trout downstream. Enclosed fish and free-ranging fish both had negative growth rates during the summer. Enclosed fish lost significantly less weight than free-ranging fish. Instantaneous growth rates of free-ranging adult brook charr and rainbow trout from May to August were negative for both species in all stream zones. Underwater observations of adult brook charr and rainbow trout showed both species occurred significantly more often in pool macrohabitats than expected on the basis of macrohabitat availability, except for rainbow trout in the upstream zone. The proportion of pool macrohabitat was not significantly different among stream zones. Brook charr do not appear to be better adapted to upstream environments in Powdermill Run based on growth, survival and macrohabitat selection during summer. Negative biotic interactions acting along with differential environmental adaptations may explain the pattern of distribution of brook charr and rainbow trout in streams, but long-term transplant experiments with additional life stages will be necessary to examine this hypothesis.     

Growth in Arctic charr and rainbow trout fed temporally concentrated or spaced daily meals

The effect on growth of distributing feed over a few hours compared tomore frequent meals was tested on 1+ Arctic charr (Salvelinus alpinus L.) and 1+ rainbow trout (Oncorhynchus mykiss Walbaum). Triplicate hatchery groups for each treatment were fed at a ration level of 1%/dayeither with few meals (8 times per day divided into morning and evening)or with frequent meals (32 meals equally distributed during the day). Wefound an opposite effect of meal frequency on growth in the two species.Low feeding intensity (8 meals per day) had a significantly positive effecton growth in rainbow trout but a significantly negative effect on growth inArctic charr when compared to feeding the fish frequent meals. Theopposite response to meal frequency is likely to be an effect of thedifferences in activity during feeding. Rainbow trout feed much moreaggressively than charr which can result in feeding being a more stressfulevent. In this experiment, the specific growth rate was lower and the feedconversion ratio higher for Arctic charr compared to rainbow trout.     

Temporal segregation in spawning between native Yellowstone cutthroat trout and introduced rainbow trout

Hybridisation with introduced taxa poses a threat to native fish populations. Mechanisms of reproductive isolation can limit or prevent hybridisation between closely related species. Understanding how these mechanisms interact between the same species across geographically distinct occurrences of secondary contact, and how regional factors influence them, can inform our understanding of hybridisation as a threat and management actions to mitigate this threat. We used data collected on adult fish migration timing and approximate emergence timing of subsequent juvenile fish paired with genomic data to assess whether temporal isolation in the timing of spawning exists between Yellowstone cutthroat trout, rainbow trout and hybrids in the North Fork Shoshone River drainage in northwest Wyoming. We found evidence that Yellowstone cutthroat trout spawn, on average, two to four weeks later than rainbow trout and hybrids and two environmental covariates related to water temperature and discharge were associated with differences in spawning migration timing. Despite statistical support for Yellowstone cutthroat trout spawning later, disproportionately high numbers of rainbow trout and hybrids, paired with extended spawning seasons, lead to substantial overlap between all genotypes. Our results provide further evidence of temporal segregation in the timing of spawning as a mechanism of reproductive isolation between closely related species, but substantial spawning overlap suggests temporal segregation alone will not be enough to curtail hybridisation in conservation populations.     

Effects of oxygen supersaturation on the culture of cutthroat trout, Oncorhynchus clarki Richardson, and rainbow trout, Oncorhynchus mykiss Richardson

Abstract. Two studies were conducted to determine the effects of rearing Snake River cutthroat trout, Oncorhynchus clarki Richardson, and Eagle Lake rainbow trout, Oncorhynchus mykiss Richardson, in oxygen supersaturated water. The performance of cutthroat trout held at oxygen saturation as high as 172% was compared with that of control fish held in water at or below saturation. At an oxygen saturation level of 172%, total gas pressure reached 117% in late afternoon, and nitrogen saturation was reduced to 93%. The rainbow trout were held at a maximum of 150% oxygen saturation; total gas pressure did not exceed 112% in late afternoon and nitrogen saturation was reduced to 99%. Fish growth, fin quality and feed conversions were not significantly affected in either species. At the termination of the study gas bubble disease was observed in 94% of the cutthroat trout held in oxygen supersaturated water. Gas bubble disease was not observed in rainbow trout.     

Survival, growth and dressing traits of triploid hybrids between rainbow trout and three charr species

Triploid hybrids between female rainbow trout Oncorhynchus mykiss and male brook charr Salvelinus fontinalis, Arctic charr S. alpinus and lake charr S. namaycush, together with diploid and triploid rainbow trout controls from the same dams, were tested in freshwater farming up to their fourth year of life. All hybrids displayed lower survival rates than the controls, the weakest genotype being the Arctic charr hybrid. Mortalities were mostly observed at the embryonic and larval stages and at the adult stage as a consequence of male sexual maturation. Growth of all hybrids was hindered (compared with controls) during the first year, but only moderate differences remained after 3 years. Sexual maturation resulted in a weight inferiority of males in all genotypes. As to carcass traits, female hybrids displayed a slightly higher dressing percentage than female triploid rainbow trout, as a result of lower visceral losses. These results are discussed with reference to hybrid resistance to rhabdoviruses from the viewpoint of fish farming improvement.     

Invasions of rainbow trout and brown trout in Japan: A comparison of invasiveness and impact on native species

Many species of salmonids have been stocked into waters outside of their native range. The invasiveness and impact of these species on native species varies depending on their biological traits, and on environmental conditions, such as climate. In Japan, rainbow trout and brown trout, both listed in 100 of the world's worst invasive alien species by the International Union for Conservation of Nature, occur as non-native species. The invasiveness of these two species is thought to be related to seasonal flooding, given flood waters can physically damage fry and prevent population establishment. Rainbow trout have successfully invaded waters in Hokkaido, northern Japan, where the likelihood of flooding is low between June and July, when their fry emerge, but successful invasions are rare in regions south of Hokkaido. Brown trout, however, have successfully invaded waters not only in Hokkaido, but also other regions. Since brown trout have a similar life history to the native white-spotted charr and masu salmon, with fry emerging before the flood season, they are more suited to the Japanese climate than Rainbow trout. Rainbow and brown trout interact with native species in various ways, but a common outcome of these interactions is the displacement of native charr species. Legal regulations of non-native salmonids should be based on understandings of the ecological traits of each invasive species and regional impacts on native species. Given the ongoing nature of climate change, the nature and extent of the effects of rainbow and brown trout on native species might also change.     

Are invasive and native trout functionally equivalent predators? Results and lessons from a field experiment

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Projected impacts of climate change on stream salmonids with implications for resilience‐based management

The sustainability of freshwater fisheries is increasingly affected by climate warming, habitat alteration, invasive species and other drivers of global change. The State of Michigan, USA, contains ecologically, socioeconomically valuable coldwater stream salmonid fisheries that are highly susceptible to these ecological alterations. Thus, there is a need for future management approaches that promote resilient stream ecosystems that absorb change amidst disturbances. Fisheries professionals in Michigan are responding to this need by designing a comprehensive management plan for stream brook charr (Salvelinus fontinalis), brown trout (Salmo trutta) and rainbow trout (Oncorhynchus mykiss) populations. To assist in developing such a plan, we used stream‐specific regression models to forecast thermal habitat suitability in streams throughout Michigan from 2006 to 2056 under different predicted climate change scenarios. As baseflow index (i.e., relative groundwater input) increased, stream thermal sensitivity (i.e., relative susceptibility to temperature change) decreased. Thus, the magnitude of temperature warming and frequency of thermal habitat degradation were lowest in streams with the highest baseflow indices. Thermal habitats were most suitable in rainbow trout streams as this species has a wider temperature range for growth (12.0–22.5 °C) compared to brook charr (11.0–20.5 °C) and brown trout (12.0–20.0 °C). Our study promotes resilience‐based salmonid management by providing a methodology for stream temperature and thermal habitat suitability prediction. Fisheries professionals can use this approach to protect coldwater habitats and drivers of stream cooling and ultimately conserve resilient salmonid populations amidst global change.     

Effect of hook type and hook setting method on deep-hooking rates when bait fishing for trout in lentic waters

Fishing with circle hooks along with prescribed hook-setting methods can reduce deep-hooking rates in some fisheries, but baited circle hooks have not been evaluated in stillwater trout fisheries. Deep-hooking rates and catch probabilities were compared for cutthroat trout Oncorhynchus clarkii​ (Richardson), rainbow trout Oncorhyncus mykiss (Walbaum), cutthroat × rainbow trout hybrids, and brook trout Salvelinus fontinalis (MItchill) caught using three types of baited hooks (circle, intermediate circle and traditional J) and two hook-setting methods (active and passive) in several lakes containing naturally reared trout. Hook type had a negligible effect on deep-hooking rates, which averaged 20%. Actively setting the hook and fishing with a bobber both reduced deep-hooking rates, regardless of hook type. Larger fish were deeply hooked more frequently regardless of hook type. Catch probabilities were higher when active rather than passive hook-setting was used, particularly in combination with intermediate circle hooks. The present results, which indicate that circle hook use did not reduce deep hooking in lentic settings, contrast earlier studies done in lotic waters. Therefore, before proposing hook-type regulations intended to reduce deep hooking, fisheries managers should consider waterbody type in addition to fishing methods, fish size and species.     

A Survey of Trout Fin Erosion, Water Quality, and Rearing Conditions at State Fish Hatcheries in Utah

A total of 600 hatchery trout were examined for signs of fin erosion including rainbow trout Oncorhynchus mykiss , cutthroat trout O. clarki , brook trout Salvelinus fontinalis and brown trout Salmo trutta . A scoring system was used to evaluate erosion on all fins from 20 fish samples. Water quality and hatchery rearing variables were also determined for corresponding raceways or ponds. For rainbow trout groups ( N = 24), stepwise multiple linear regression was used to interpret the relationship between fin erosion and the other variables. These fish groups averaged between 92 and 243 mm in total length and no significant correlation was observed between length and a fin erosion index ( r = 0.045). The best-fit regression model (adjusted R² = 0.689) suggested that fin erosion was correlated with lower alkalinities, unnatural bottom substrates (concrete or steel), higher unionized ammonia levels, and higher fish densities. Despite significant variation between hatcheries, fin condition was significantly better in rainbow trout than in cutthroat trout in three of four hatcheries containing both species and the same substrate. Fin erosion in rainbow trout occurred primarily on dorsal fins, followed in order of decreasing severity, by pectoral, caudal, anal, and ventral fins.     

Stable isotope evaluation of population‐ and individual‐level diet variability in a large,oligotrophic lake with non‐native lake trout

Non‐native piscivores can alter food web dynamics; therefore, evaluating interspecific relationships is vital for conservation and management of ecosystems with introduced fishes. Priest Lake, Idaho, supports a number of introduced species, including lake trout Salvelinus namaycush, brook trout S. fontinalis and opossum shrimp Mysis diluviana. In this study, we used stable isotopes (δ¹³C and δ¹⁵N) to describe the food web structure of Priest Lake and to test hypotheses about apparent patterns in lake trout growth. We found that isotopic niches of species using pelagic‐origin carbon did not overlap with those using more littoral‐origin carbon. Species using more littoral‐origin carbon, such as brook trout and westslope cutthroat trout Oncorhynchus clarki lewisi, exhibited a high degree of isotopic niche overlap and high intrapopulation variability in resource use. Although we hypothesised that lake trout would experience an ontogenetic diet shift, no such patterns were apparent in isotopic signatures. Lake trout growth rates were not associated with patterns in δ¹⁵N, indicating that variation in adult body composition may not be related to adult diet. Understanding trophic relationships at both the individual and species levels provides a more complete understanding of food webs altered by non‐native species.     

Selective predation of threespine stickleback by rainbow trout

Abstract – Threespine stickleback (Gasterosteus aculeatus) with divergent pelvic phenotypes from Wallace Lake, Alaska, were exposed to rainbow trout (Oncorhynchus mykiss) to test for selective predation. Trout more often successfully preyed upon individuals with pelvic reduction, which supports the hypothesis that robust pelvic girdles are maintained in stickleback populations because of selection by fish predators. Stickleback with complete pelvic girdles experienced a higher incidence of wounds from unsuccessful predation attempts, although this finding was not statistically significant, possibly because of low frequencies of wounded individuals. Spines likely facilitate postcapture defence, although stickleback with spines may also be less targeted by trout. This study supports the hypothesis that stocked rainbow trout pose a conservation threat to pelvic‐reduced stickleback populations.     

Habitat niche separation of the nonnative rainbow trout and native masu salmon in the Atsuta River,Hokkaido, Japan

The mechanisms by which nonnative species establish populations can be classified into two broad categories: they usurp the niches of native species through interspecific competition, or they avoid this intense interspecific competition by making use of minimal niche overlap with the native species. In this study, we considered how a nonnative salmonid species, the rainbow trout Oncorhynchus mykiss, established a population in the presence of the native salmonid species, the masu salmon O. masou, in Hokkaido, Japan. Circumstantial field evidence shows that the masu salmon exceeds the rainbow trout in abundance and suggests that these species use different types of cover habitat (rainbow trout abundance increases with increasing abundance of large woody debris aggregates, whereas masu salmon abundance increases with increasing abundance of undercut banks). These results imply that the rainbow trout established a population due to minimal niche overlap with the masu salmon, and not by competitive exclusion of the native species.     

Brook charr, Salvelinus fontinalis (Mitchill), and cryptobiosis: a potential salmonid reservoir host for Cryptobia salmositica Katz, 1951

Abstract. Laboratory-raised Cryptobia -susceptible brook charr, Salvelinus fontinalis (Mitchill), and rainbow trout, Oncorhynchus mykiss (Walbaum), were vaccinated intraperitoneally with a live Cryptobia salmositica vaccine (250000 parasites per fish), and 4 weeks later were challenged with the pathogen (250000 parasites per fish). Unvaccinated and infected brook charr had high parasitaemias but no clinical signs of disease, while unvaccinated and infected rainbow trout had anaemia and general oedema. Vaccinated and challenged fish had very low parasitaemias compared to unvaccinated and infected brook charr and rainbow trout. Complement fixing antibodies were detected in vaccinated and challenged fish 2 weeks after challenge. Unvaccinated and infected brook charr had consistently higher litres of complement fixing antibody than unvaccinated and infected rainbow trout. Parasitaemias were lower in all fish in which titres of complement fixing antibody were high. In a second experiment, brook charr inoculated intraperitoneally or intramuscularly with 100000 C. salmositica per fish had high parasitaemias but no anaemia or other clinical signs. The results show that susceptible brook charr do not suffer from cryptobiosis and may serve as reservoir hosts for C. salmositica in areas where the disease is prevalent. Vaccination to reduce the parasitaemia when fish become infected may be a control strategy in these areas.     

Competitive effects between rainbow trout and Atlantic salmon in natural and artificial streams

Competition with non‐native species may impede the restoration of native species, but differences in competitive abilities among intraspecific native populations may make some populations more suitable for reintroduction than others. Here, juvenile Atlantic salmon (Salmo salar) from two allopatric populations (LaHave and Sebago) being used for reintroduction into Lake Ontario were placed into two natural stream sites differing in the presence of ecologically similar rainbow trout (Oncorhynchus mykiss). We assessed the effects of competition in the natural streams on fitness‐related traits and habitat use of the Atlantic salmon. We then compared these effects to those observed in artificial streams from a previous study. Atlantic salmon in natural streams had reduced fitness‐related traits that were associated with suboptimal microhabitats in the presence of rainbow trout, but utilised optimal microhabitats in their absence. In the presence of rainbow trout, the two Atlantic salmon populations exhibited comparable recapture proportions to each other, but the individuals from the Sebago population had better performance (body size and condition) than those from the LaHave population. Responses of both Atlantic salmon populations to competition with rainbow trout were generally similar in both direction and magnitude when compared to results from the artificial stream study. The combined results suggest that native species restoration efforts should be focused on candidate populations that are ecologically suitable to reintroduction environments, as well as on suitable habitats that do not contain exotic competitors. Moreover, this study highlights the value of controlled experiments in artificial environments for predicting fitness‐related performance in natural environments.     

Controls for trout and char migratory/resident behaviour mainly in stream systems above and below waterfalls/barriers: a multidecadal and broad geographical review

Northcote TG. Controls for trout and char migratory/resident behaviour mainly in stream systems above and below waterfalls/barriers: a multidecadal and broad geographical review. Ecology of Freshwater Fish 2010: 19: 487–509. © 2010 John Wiley & Sons A/S Abstract – Controls for trout and char migratory and resident behaviour in rivers and streams for above and below natural and man‐made waterfalls/barriers are covered in a multi‐decadal (1950–2000) and wide geographical review (North America, UK, Europe, Japan) that includes reference to over 380 publications. These note 53 for rainbow trout, 61 for cutthroat trout, 104 for brown trout, 47 for bull trout, 41 for brook trout, 35 for Dolly Varden and 42 for white‐spotted char, plus a few general contributions of relevance on some. For each of these species, there has been a major increase in relevant decadal publications since the early 1980s, no doubt in large part because of the upsurge in micro‐genetic methodology for DNA and related technology, coupled with a broadening of interest in stream migratory behaviour of salmonids. Main mechanisms for the control of stream migratory versus resident behaviour appear to cover an interplay among both genetic and environmental factors; in some populations and locations, genetic controls seem to be more important than environmental ones, but in others the reverse. Habitat degradation by various human activities and their introductions of non‐native fish species are becoming causes of reduction in abundance of above and below waterfall stream populations for several of these salmonid species.