Density‐dependent effects of eastern Kamchatka pink salmon (Oncorhynchus gorbuscha) and Japanese chum salmon (O. keta) on age‐specific growth of western Alaska chum salmon

Hatcheries release >4.5 billion juvenile Pacific salmon (Oncorhynchus spp.) into the North Pacific Ocean annually, raising concerns about competition with wild salmon populations. We used retrospective scale analysis to investigate how the growth of chum salmon (O. keta) from western Alaska is affected by the abundance of chum salmon from Japanese hatcheries and wild pink salmon (O. gorbuscha) from the Russian Far East. Over nearly five decades, the growth of Kuskokwim River chum salmon was negatively correlated with the abundance of Japanese hatchery chum salmon after accounting for the effects of sex and spring/summer sea‐surface temperature in the Bering Sea. An effect of wild eastern Kamchatka pink salmon abundance on the growth of Kuskokwim River salmon was detectable but modest compared to the intraspecific competitive effect. A decrease in Japanese hatchery chum salmon releases in 2011–2013 was not associated with increased growth of Bering Sea chum salmon. However, the abundance of wild chum salmon from the Russian Far East increased during that time, possibly obscuring reduced competition with hatchery chum salmon. Our results support previous evidence that chum salmon are affected by intraspecific competition, and to a lesser extent interspecific competition, in the North Pacific, underscoring that the effects of salmon hatchery production transcend national boundaries.     

Spatial variations in feeding and condition of juvenile pink and chum salmon off Vancouver Island, British Columbia

Spatial variations in feeding and condition of juvenile pink (Oncorhynchus gorbuscha) and chum (Oncorhynchus keta) salmon, and their implications for growth, were examined on the Vancouver Island continental shelf in early summer 1992. Juvenile pink salmon off northern Vancouver Island had more material in their stomachs, were in better condition, and had higher potential growth rates (from a bioenergetics model) than pink salmon off southern Vancouver Island. These variations were consistent with spatial differences in zoo-plankton biomass, there being more plankton in the northern region. There was a significant positive relationship between condition of pinks and the amount of material in their stomachs, suggesting a positive feedback on feeding success. Juvenile chum in the north also had more material in their stomachs than chum to the south. However, condition factor was not significantly different between southern and northern regions nor was there a significant relationship between condition factor and the weight of stomach contents for chum on the southern shelf. A bioenergetics model suggests that chum in the south were food limited. Stable carbon isotope data also indicated different feeding histories for some chum in the southern region, which may have been recent migrants onto the continental shelf from near-shore areas, or possibly a nearby hatchery. Estimation of the energy required by juvenile salmon to migrate north in a continental shelf area with low zooplankton biomass and a weak northerly current (inner shelf), compared with an area with higher zooplankton biomass but a strong southerly current (outer shelf), indicated sufficient surplus energy only in the inner shelf, consistent with observations of northward migrations predominantly through this area. Spatial variations in current velocity and zooplankton biomass can affect feeding, condition, and potential growth of juvenile pink and chum salmon off Vancouver Island.     

Changing salmon: An analysis of body mass,abundance, survival,and productivity trends across 45 years in Puget Sound

Pacific salmon and trout (Oncorhynchus spp., Salmonidae) of the Puget Sound region of Washington State, USA, have experienced recent and longer‐term (multidecadal) variability in abundance while supporting robust fisheries. As part of the post‐season salmon management process, population‐specific estimates of total adult abundance to Puget Sound (Strait of Juan de Fuca) for pink (O. gorbuscha), chum (O. keta), coho (O. kisutch), sockeye (O. nerka), and Chinook (O. tshawytscha) salmon and steelhead trout (O. mykiss) are calculated annually. We compiled annual estimates of body mass, abundance and survival of hatchery‐ and naturally produced salmon from 1970 to 2015 to compare spatial and temporal patterns across species. Average weights of adult salmon and steelhead returning to Puget Sound, with the exception of coho salmon, have decreased since the 1970s. Temporal trends in abundance, survival and productivity varied by species and origin (hatchery vs. naturally produced). Generally, abundance and survival rates of natural‐origin species decreased whereas those of hatchery‐produced species did not, which is in contrast with other studies' general conclusions of decreasing survival among Puget Sound salmonids. Species diversity has decreased in recent years, with salmonids that rely on a short freshwater rearing phase in the natural environment (hatchery‐produced fish and naturally produced pink and chum) representing >90% of total returns in most years. This new information reveals patterns of body size, abundance, survival and productivity across species, life history and rearing type over the past 45 years and, in doing so, demonstrates the strength in multidecadal, multifactor time series to critically evaluate salmonid species.     

Multi‐timescale interactions between pink and chum salmon catch per unit effort in the Bering Sea

Based on generalized linear models, interspecific interactions were identified between chum and pink salmon. In addition, the effects of sea surface temperature and location on the variability of catch per unit effort (CPUE) of chum salmon from gill‐net surveys carried out between 1972 and 2010 were investigated. In the optimal model, interspecific interactions between CPUEs of chum and pink salmon on a year scale were positive for approximately half of all years in the central Bering Sea. In addition, interspecific interactions on a multi‐year scale were positive in even‐numbered years. The effects of location on the CPUE of chum salmon were significant variables in the optimal model. The CPUEs of chum salmon located near the continental shelf in the Bering Sea were higher than those of other locations. This study provides new evidence of positive interspecific interactions between the CPUEs of chum and pink salmon. The results also suggest that the standardized CPUE of chum salmon from the gill‐net surveys reflects relative chum salmon abundance in the North Pacific Ocean in the following year.     

Freshwater habitat associations between pink (Oncorhynchus gorbuscha), chum (O. keta) and Chinook salmon (O. tshawytscha) in a watershed dominated by sockeye salmon (O. nerka) abundance

To understand the interplay between habitat use and contemporary anadromous Pacific salmon, Oncorhynchus spp., distributions we explored the habitat associations of three species, pink (O. gorbuscha), chum (O. keta) and Chinook salmon (O. tshawytscha) in streams of the Wood River system of Bristol Bay, Alaska, where sockeye salmon (O. nerka) are numerically dominant. We developed models to investigate the occurrence of nondominant salmon in relation to habitat characteristics and sockeye salmon density, using four decades of salmon presence and abundance data. The frequency of occurrence and abundance of nondominant species increased with watershed drainage area and stream depth and decreased with sockeye salmon density. The range of occurrence varied from nonexistent to perennial for the other species in sockeye‐dominated streams. Increasing watershed area resulted in larger stream habitat area and deeper habitats, allowing for the sympatric occurrence and persistence of all salmon species. The relationships between habitat and the presence of these Pacific salmon help define their requirements but also remind us that the patterns of presence and absence, within the overall ranges of salmon species, have yet to be fully understood.     

Ecological processes influencing mortality of juvenile pink salmon (Oncorhynchus gorbuscha) in Prince William Sound,Alaska

Our collaborative work focused on understanding the system of mechanisms influencing the mortality of juvenile pink salmon (Oncorhynchus gorbuscha) in Prince William Sound, Alaska. Coordinated field studies, data analysis and numerical modelling projects were used to identify and explain the mechanisms and their roles in juvenile mortality. In particular, project studies addressed the identification of major fish and bird predators consuming juvenile salmon and the evaluation of three hypotheses linking these losses to (i) alternative prey for predators (prey‐switching hypothesis); (ii) salmon foraging behaviour (refuge‐dispersion hypothesis); and (iii) salmon size and growth (size‐refuge hypothesis). Two facultative planktivorous fishes, Pacific herring (Clupea pallasi) and walleye pollock (Theragra chalcogramma), probably consumed the most juvenile pink salmon each year, although other gadids were also important. Our prey‐switching hypothesis was supported by data indicating that herring and pollock switched to alternative nekton prey, including juvenile salmon, when the biomass of large copepods declined below about 0.2 g m^?3. Model simulations were consistent with these findings, but simulations suggested that a June pteropod bloom also sheltered juvenile salmon from predation. Our refuge‐dispersion hypothesis was supported by data indicating a five‐fold increase in predation losses of juvenile salmon when salmon dispersed from nearshore habitats as the biomass of large copepods declined. Our size‐refuge hypothesis was supported by data indicating that size‐ and growth‐dependent vulnerabilities of salmon to predators were a function of predator and prey sizes and the timing of predation events. Our model simulations offered support for the efficacy of representing ecological processes affecting juvenile fishes as systems of coupled evolution equations representing both spatial distribution and physiological status. Simulations wherein model dimensionality was limited through construction of composite trophic groups reproduced the dominant patterns in salmon survival data. In our study, these composite trophic groups were six key zooplankton taxonomic groups, two categories of adult pelagic fishes, and from six to 12 groups for tagged hatchery‐reared juvenile salmon. Model simulations also suggested the importance of salmon density and predator size as important factors modifying the predation process.     

Experimental infections with Lepeophtheirus salmonis (Krøyer) on threespine sticklebacks, Gasterosteus aculeatus L., and juvenile Pacific salmon, Oncorhynchus spp

Experimental infections with Lepeophtheirus salmonis (Krøyer) were established on threespine sticklebacks, Gasterosteus aculeatus L., juvenile pink, Oncorhynchus gorbuscha (Walbaum), and chum, Oncorhynchus keta (Walbaum), salmon. The prevalence and abundance of infections were initially higher on sticklebacks than on either salmon species. The initial prevalence and intensity of infections on chum salmon were higher than those on pink salmon, and declined on both species during louse development. The rate of parasite development to adult stages was similar on all species although development beyond the preadult stage was not observed on sticklebacks. These results confirm previous field observations on the occurrence and development of L. salmonis on threespine sticklebacks.     

Potential trophodynamic and environmental drivers of steelhead (Oncorhynchus mykiss) productivity in the North Pacific Ocean

Information on prey availability, diets, and trophic levels of fish predators and their prey provides a link between physical and biological changes in the ecosystem and subsequent productivity (growth and survival) of fish populations. In this study two long‐term data sets on summer diets of steelhead (Oncorhynchus mykiss) in international waters of the central North Pacific Ocean (CNP; 1991–2009) and Gulf of Alaska (GOA; 1993–2002) were evaluated to identify potential drivers of steelhead productivity in the North Pacific. Stable isotopes of steelhead muscle tissue were assessed to corroborate the results of stomach content analysis. We found the composition of steelhead diets varied by ocean age group, region, and year. In both the GOA and CNP, gonatid squid (Berryteuthis anonychus) were the most influential component of steelhead diets, leading to higher prey energy densities and stomach fullness. Stomach contents during an exceptionally warm year in the GOA and CNP (1997) were characterized by high diversity of prey with low energy density, few squid, and a large amount of potentially toxic debris (e.g., plastic). Indicators of good diets (high proportions of squid and high prey energy density) were negatively correlated with abundance of wild populations of eastern Kamchatka pink salmon (O. gorbuscha) in the CNP. In conclusion, interannual variations in climate, abundance of squid, and density‐dependent interactions with highly‐abundant stocks of pink salmon were identified as potential key drivers of steelhead productivity in these ecosystems. Additional research in genetic stock identification is needed to link these potential drivers of productivity to individual populations.     

Change in feeding ecology and trophic dynamics of Pacific salmon (Oncorhynchus spp.) in the central Gulf of Alaska in relation to climate events

The effects of climate events on the feeding ecology and trophic dynamics of Pacific salmon (Oncorhynchus spp.) in offshore waters of the central Gulf of Alaska were investigated during early summers (1994–2000), based on analyses of stomach contents, and carbon and nitrogen stable isotopes (δ¹³C and δ¹⁵N). Gonatid squids (mainly Berryteuthis anonychus) were the dominant prey of all salmon species except for chum salmon (O. keta). During the 1997 El Niño event and the 1999 La Niña event, squids decreased sharply in the diets of all Pacific salmon except coho salmon (O. kisutch) in the Subarctic Current, and chum salmon diets changed from gelatinous zooplankton (1995–97) to a more diverse array of zooplankton species. A δ¹³C and δ¹⁵N analysis indicated that all salmon species occupied the same branch of the food web in 1999–2000. We hypothesize that high‐seas salmon adapt to climate‐induced changes in their prey resources by switching their diets either within or between trophic levels. To understand the effects of climate change on Pacific salmon in the Gulf of Alaska, biological oceanographic research on B. anonychus and other important prey resources is needed.     

An evaluation of the potential influence of SST and currents on the oceanic migration of juvenile and immature chum salmon (Oncorhynchus keta) by a simulation model

Using a salmon migration model based on the assumption that swimming orientation is temperature dependent, we investigated the determining factors of the migration of juvenile and immature chum salmon (Oncorhynchus keta) in the North Pacific. We compared the predictions of the model with catch data of immature and juvenile chum salmon collected by Japanese research vessels from 1972 to 1999. The salmon migration model reproduced the observed distributions of immature chum salmon and indicates that passive transport by wind‐driven and geostrophic currents plays an important role in the eastward migration of Asian salmon. These factors result in a non‐symmetric distribution of Asian and North American chum salmon in the open ocean. The directional swimming component contributes to the northward migration in summer. The model results indicate that during the first winter Asian chum salmon swim northward against the southward wind‐driven currents to stay in the western North Pacific. This suggests that Asian chum salmon require more energy to migrate than other stocks during the first winter of their ocean life.     

Foraging behaviour of juvenile pink salmon (Oncorhynchus gorbuscha) and size‐dependent predation risk

Two hypotheses related to effects of juvenile pink salmon (Oncorhynchus gorbuscha) foraging behaviour and size on their predation risk were evaluated using field data collected in Prince William Sound, Alaska 1 995–97. My results supported the hypothesis that low macrozooplankton density leads to dispersion of juvenile salmon from shallow nearshore habitats and greater predation risk, but zooplankton type was an important factor. When the biomass of large copepods (primarily Neocalanus spp.) declined, salmon dispersed from shallow nearshore habitats, and mean daily individual predator consumption of salmon increased by a factor of 5. A concomitant five‐fold increase in the probability of occurrence of salmon in predator stomachs supported the notion that increased predation on salmon was caused by a greater overlap between predator and prey when salmon dispersed offshore, not an increase in the number of salmon consumed per feeding bout. The results also generally supported the hypothesis that the timing of predation events modifies the nature of size‐dependent predation losses of salmon to different predator groups (small and large planktivores and piscivores). Size‐dependent vulnerabilities of salmon to predators were a function of both predator and prey sizes. When simulated predation was shifted from May to June, the vulnerability of salmon became more dependent on their growth than initial size. But, the size‐ and growth‐dependent vulnerabilities of salmon differed more among predator groups than between May and June, suggesting that changes in the composition of predator fields could more strongly affect the nature of size‐dependent predation than changes in the timing of predation losses.     

Body size trends along vertical and thermal gradients of chum salmon in the Bering Sea during summer

Relationships between the vertical distribution and thermal habitat, and body size of chum salmon Oncorhynchus keta were studied in the Bering Sea in summer using trawl surveys at various depths. Chum salmon abundance decreased with increasing depth, but the patterns of decrease differed between size groups. The abundance of small salmon fell rapidly with depth, whereas that of large salmon decreased gradually to 40 m depth, and abruptly below that. The average fork length of chum salmon collected from each trawl correlated positively with trawl net depth and negatively with water temperature. Since the optimal temperature for growth decreases with body size in this species, the observed body size‐related vertical habitat use by chum salmon may indicate size‐dependent thermal preferences.     

Development of a Piscirickettsia salmonis immersion challenge model to investigate the comparative susceptibility of three salmon species

Piscirickettsia salmonis, the aetiological agent of salmonid rickettsial septicaemia (SRS), is a global pathogen of wild and cultured marine salmonids. Here, we describe the development and application of a reproducible, standardized immersion challenge model to induce clinical SRS in juvenile pink (Oncorhynchus gorbuscha), Atlantic (Salmo salar) and sockeye salmon (O. nerka). Following a 1‐hr immersion in 10⁵ colony‐forming units/ml, cumulative mortality in Atlantic salmon was 63.2% while mortality in sockeye salmon was 10%. Prevalence and levels of the bacterium in kidney prior to onset of mortality were lower in sockeye compared with Atlantic or pink salmon. The timing and magnitude of bacterial shedding were estimated from water samples collected during the exposure trials. Shedding was estimated to be 82‐fold higher in Atlantic salmon as compared to sockeye salmon and peaked in the Atlantic salmon trial at 36 d post‐immersion. These data suggest sockeye salmon are less susceptible to P. salmonis than Atlantic or pink salmon. Finally, skin lesions were observed on infected fish during all trials, often in the absence of detectable infection in kidney. As a result, we hypothesize that skin is the primary point of entry for P. salmonis during the immersion challenge.     

Introduced northern pike predation on salmonids in southcentral Alaska

Northern pike (Esox lucius) are opportunistic predators that can switch to alternative prey species after preferred prey have declined. This trophic adaptability allows invasive pike to have negative effects on aquatic food webs. In Southcentral Alaska, invasive pike are a substantial concern because they have spread to important spawning and rearing habitat for salmonids and are hypothesised to be responsible for recent salmonid declines. We described the relative importance of salmonids and other prey species to pike diets in the Deshka River and Alexander Creek in Southcentral Alaska. Salmonids were once abundant in both rivers, but they are now rare in Alexander Creek. In the Deshka River, we found that juvenile Chinook salmon (Oncorhynchus tshawytscha) and coho salmon (O. kisutch) dominated pike diets and that small pike consumed more of these salmonids than large pike. In Alexander Creek, pike diets reflected the distribution of spawning salmonids, which decrease with distance upstream. Although salmonids dominated pike diets in the lowest reach of the stream, Arctic lamprey (Lampetra camtschatica) and slimy sculpin (Cottus cognatus) dominated pike diets in the middle and upper reaches. In both rivers, pike density did not influence diet and pike consumed smaller prey items than predicted by their gape‐width. Our data suggest that (1) juvenile salmonids are a dominant prey item for pike, (2) small pike are the primary consumers of juvenile salmonids and (3) pike consume other native fish species when juvenile salmonids are less abundant. Implications of this trophic adaptability are that invasive pike can continue to increase while driving multiple species to low abundance.     

The vertical distribution of juvenile salmon (Oncorhynchus spp.) and associated fishes in the Columbia River plume

Simultaneous trawling at surface and at depth at one location off the Columbia River, Oregon, in June 2000 identified the depth distribution of juvenile salmonids and associated fishes. Juvenile salmon off the Columbia River were distributed primarily near the surface, within the upper 12 m. Highest densities of subyearling chinook salmon (Oncorhynchus tshawytscha) off the Columbia River were associated with high surface currents and decreasing tidal levels, with time of day possibly a co‐factor. Densities of yearling chinook salmon increased with higher turbidity. Pacific herring (Clupea pallasi) was the most abundant and commonly caught forage fish, with density increasing at night, probably related to diel vertical migration. Catches of juvenile salmonids were not associated with catches of forage fishes. Daytime surface trawling appears to be an appropriate method for assessing the distribution and abundance of juvenile salmonids in marine habitats.     

Escaped farmed Atlantic salmon, Salmo salar L., feeding in Scottish coastal waters

Escaped reared salmon, Salmo salar L., were distinguished from wild salmon in the catch of a coastal salmon fishery on the west coast of Scotland. The stomach contents of 54 escaped fish were examined to determine their recent feeding history and 19 (35%) were found to contain food. The predominant prey were juvenile whiting, Merlangius merlangus (L.), unidentified Gadidae and sandeels (Ammodytidae), although other fish and invertebrates, mainly post-larval hermit crabs (Paguridae), were recorded. All these prey are pelagic or semi-pelagic. These observations demonstrate that escaped salmon feed on natural prey in coastal waters and extend our knowledge of the diet of salmon in their marine phase.     

Introduced northern pike consumption of salmonids in Southcentral Alaska

The impacts of introduced northern pike (Esox lucius) on salmonid populations have attracted much attention because salmonids are popular subsistence, sport and commercial fish. Concern over the predatory effects of introduced pike on salmonids is especially high in Southcentral Alaska, where pike were illegally introduced to the Susitna River basin in the 1950s. We used pike abundance, growth, and diet estimates and bioenergetics models to characterise the realised and potential consumptive impacts that introduced pike (age 2 and older) have on salmonids in Alexander Creek, a tributary to the Susitna River. We found that juvenile salmonids were the dominant prey item in pike diets and that pike could consume up to 1.10 metric tons (realised consumption) and 1.66 metric tons (potential consumption) of juvenile salmonids in a summer. Age 3–4 pike had the highest per capita consumption of juvenile salmonids, and age 2 and age 3–4 pike had the highest overall consumption of juvenile salmonid biomass. Using historical data on Chinook salmon and pike potential consumption of juvenile salmonids, we found that pike consumption of juvenile salmonids may lead to collapsed salmon stocks in Alexander Creek. Taken together, our results indicate that pike consume a substantial biomass of juvenile salmonids in Alexander Creek and that coexistence of pike and salmon is unlikely without management actions to reduce or eliminate introduced pike.     

Competition between Asian pink salmon (Oncorhynchus gorbuscha) and Alaskan sockeye salmon (O. nerka) in the North Pacific Ocean

The importance of interspecific competition as a mechanism regulating population abundance in offshore marine communities is largely unknown. We evaluated offshore competition between Asian pink salmon and Bristol Bay (Alaska) sockeye salmon, which intermingle in the North Pacific Ocean and Bering Sea, using the unique biennial abundance cycle of Asian pink salmon from 1955 to 2000. Sockeye salmon growth during the second and third growing seasons at sea, as determined by scale measurements, declined significantly in odd‐numbered years, corresponding to years when Asian pink salmon are most abundant. Bristol Bay sockeye salmon do not interact with Asian pink salmon during their first summer and fall seasons and no difference in first year scale growth was detected. The interaction with odd‐year pink salmon led to significantly smaller size at age of adult sockeye salmon, especially among younger female salmon. Examination of sockeye salmon smolt to adult survival rates during 1977–97 indicated that smolts entering the ocean during even‐numbered years and interacting with abundant odd‐year pink salmon during the following year experienced 26% (age‐2 smolt) to 45% (age‐1 smolt) lower survival compared with smolts migrating during odd‐numbered years. Adult sockeye salmon returning to Bristol Bay from even‐year smolt migrations were 22% less abundant (reduced by 5.9 million fish per year) compared with returns from odd‐year migrations. The greatest reduction in adult returns occurred among adults spending 2 compared with 3 years at sea. Our new evidence for interspecific competition highlights the need for multispecies, international management of salmon production, including salmon released from hatcheries into the ocean.     

Spatiotemporal change in growth of two populations of Asian chum salmon in relation to intraspecific interaction

Spatiotemporal changes in growth patterns of chum salmon Oncorhynchus keta that returned to the Ishikari (Japan) and Namdae (Korea) rivers in 1984–1998 were investigated using scale analysis. Juvenile chum salmon from both populations left coastal marine areas after spring at a size of over 8 cm fork length (FL). In summer, juvenile salmon from the Namdae River entered the Okhotsk Sea at a larger FL than did Ishikari River juveniles. There were no significant differences in annual growth between populations of 1-, 2-, and 4-year-old fish. For 3-year-old fish, however, Namdae River salmon had significantly higher synchronous and sympatric growth than did Ishikari River salmon. Mean FL of adults was also larger in Namdae River salmon than in Ishikari River salmon. Analysis of covariance (ANCOVA) results showed (1) negative linear relations between FL and catch, (2) homogeneous slopes of those relations at regional and species levels, and (3) nonhomogeneous slopes at the population level, indicating that density-dependent effects on growth were most significant at this level. We concluded that growth of chum salmon was concurrently influenced by stronger effects of intrapopulation competition and weaker effects of inter- and intraspecific interactions in the Bering Sea.     

Above parr: Lowland river habitat characteristics associated with higher juvenile Atlantic salmon (Salmo salar) and brown trout (S. trutta) densities

Understanding juvenile salmonid habitat requirements is critical for their effective management, but little is known about these requirements in lowland rivers, which include important but unique salmonid habitats. We compared the relative influence of in-stream Ranunculus cover, water depth, prey abundance, distance upstream and two previously unexplored factors (water velocity heterogeneity and site colonisation potential) on summer densities of juvenile Atlantic salmon and brown trout. We applied electrofishing, habitat surveys and macroinvertebrate kick sampling, and calculated the site colonisation potential from salmon redd surveys across 18–22 sites in a lowland river in 2015–2017. Due to a recruitment crash in 2016, models including and excluding this unusual year were explored. Excluding 2016 data, juvenile salmon densities showed a positive association with Ranunculus cover and numbers of nearby upstream redds, and a negative association with distance upstream from the tidal limit. Trout densities were positively associated with velocity heterogeneity, indicating a potential indirect influence of Ranunculus mediated by water velocity. When including 2016, year had the largest effect on densities of both species, highlighting the impact of the recruitment failure. These findings uncover interspecific differences in the habitat requirements of juvenile salmonids in lowland rivers. Velocity heterogeneity and site colonisation potential had high explanatory power, highlighting that they should be considered in future studies of habitat use. These findings demonstrate that temporal replication and recruitment dynamics are important considerations when exploring species–habitat associations. We discuss potential management implications and argue that Ranunculus cover could be an important management tool in conservation of lowland salmonids.