Terrestrial prey inputs to streams bordered by deciduous broadleaved forests,conifer plantations and clear‐cut sites in southwestern Japan: effects on the abundance of red‐spotted masu salmon

Terrestrial invertebrates falling from the riparian canopy are a major energy source for fishes in headwater streams. Because quantity and quality of such allochthonous resources can vary depending on riparian conditions, conversion of riparian forests to conifer plantations may affect stream productivity. We compared falling and drifting invertebrate abundances and the diet of masu salmon (Oncorhynchus masou) among stream reaches bordered by deciduous broadleaved forests, conifer plantations (Cryptomeria japonica), and clear‐cut sites in southwestern Japan. We also examined whether among‐reach variation in salmon abundance was related to the riparian vegetation types. The results indicated that, on an annual basis, falling inputs of terrestrial invertebrates at the broadleaved reaches were 2–4 times higher than those at the plantation and clear‐cut reaches. In nonwinter seasons, terrestrial invertebrates made up 40–60% and 30–90% of drift and masu salmon diets, respectively, and drifting invertebrate abundance was higher in the broadleaved reaches than in the plantation reaches. Furthermore, a multivariate analysis of salmon abundance showed that variation in salmon biomass was explained primarily by riparian vegetation type, with broadleaved and clear‐cut reaches having higher biomass than the plantation reaches. These results indicate that terrestrial invertebrates are an important resource for masu salmon, and suggest that streams bordered by conifer plantations receive lower terrestrial prey inputs, which results in lower salmon abundance. In regions where natural forests have been extensively converted to conifer plantations, forest management that allows and facilitates recovery of natural riparian stands is important.     

Effects of habitat and internal prey subsidies on juvenile coho salmon growth: implications for stream productive capacity

Abstract –  To evaluate the effects of habitat, foraging strategy (drift vs. limnetic feeding) and internal prey subsidies (downstream transport of invertebrate drift between habitats) on fish production, we measured the growth of juvenile coho salmon confined to enclosures in flowing (pond inlets and outlets) or standing water (centre of pond) habitats in a constructed river side-channel. The effects of habitat and foraging strategy on fish growth were mediated primarily through habitat effects on prey abundance. Invertebrate drift biomass was nearly an order of magnitude higher at pond inlets relative to outlets. Drift-feeding coho in inlet enclosures grew 50% faster than drift-feeding coho at pond outlets or limnetic feeding coho in the centre of ponds, suggesting that elevated drift at inlets was sufficient to account for higher inlet growth rates. Forty per cent of prey biomass in stomachs was terrestrial in origin. These results indicate that, in addition to dependence on external terrestrial subsidies, streams with alternating slow and fast water (i.e., pool-riffle) sequences are also characterised by internal prey subsidies based on transport of drifting invertebrates from refuge habitats (high velocity riffles) to habitats more suitable for drift-feeding predators (e.g., pools), which may result in higher maximum fish growth in systems where internal subsidises are large. Restoration of small streams to maximise productive capacity for pool-rearing salmonids will require a better understanding of the length and interspersion of habitats that maximises both internal prey subsidies and available rearing habitat for juvenile salmon.     

Relationships between juvenile salmon, Salmo salar L., and invertebrate densities in the River Tana, Norway

Abstract –  Juvenile salmon density was related to invertebrate density in 13 streams within the River Tana, northern Norway. There were only small, nonsignificant, differences in benthic density between streams with and without juvenile salmon. All streams with a high density of juvenile salmon had low benthic densities at the stream mouth. Juvenile salmon were not found, or were in very low densities, in streams where the benthic density at the stream mouth was as high or higher than that in the stream. A multiple regression model showed that parr density was related negatively to benthic density at the stream mouth, water velocity and pH, and positively to benthic density within the stream and the proportion of the substratum covered by moss. The amount of overhanging cover in the different streams explained 93% of the variation in the drift density of terrestrial invertebrates in August. The highest densities of juvenile salmon were found in streams with riparian vegetation, and were thus associated with an abundant supply of drift food, especially terrestrial invertebrates.     

Conserving fluxes of terrestrial invertebrates to trout in streams: A first field experiment on the effects of cattle grazing

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Trophic pathways supporting Arctic grayling in a small stream on the Arctic Coastal Plain,Alaska

Beaded streams are prominent across the Arctic Coastal Plain (ACP) of Alaska, yet prey flow and food web dynamics supporting fish inhabiting these streams are poorly understood. Arctic grayling (Thymallus arcticus) are a widely distributed upper‐level consumer on the ACP and migrate into beaded streams to forage during the short 3‐month open‐water season. We investigated energy pathways and key prey resources that support grayling in a representative beaded stream, Crea Creek. We measured terrestrial invertebrates entering the stream from predominant riparian vegetation types, prey types supporting a range of fish size classes, and how riparian plants and fish size influenced foraging habits. We found that riparian plants influenced the quantity of terrestrial invertebrates entering Crea Creek; however, these differences were not reflected in fish diets. Prey type and size ingested varied with grayling size and season. Small grayling (<15 cm fork length (FL)) consumed mostly aquatic invertebrates early in the summer, and terrestrial invertebrates later in summer, while larger fish (>15 cm FL) foraged most heavily on ninespine stickleback (Pungitius pungitius) throughout the summer, indicating that grayling can be insectivorous and piscivorous, depending on size. These findings underscore the potential importance of small streams in Arctic ecosystems as key summer foraging habitats for fish. Understanding trophic pathways supporting stream fishes in these systems will help interpret whether and how petroleum development and climate change may affect energy flow and stream productivity, terrestrial–aquatic linkages and fishes in Arctic ecosystems.     

A comparative study of juvenile salmon density in 20 streams throughout a very large river system in northern Norway

Abstract – The objective was to compare juvenile salmon density in 20 streams throughout the very large River Tana, northern Norway, and to relate variation in density to a suite of environmental factors. Four sampling sites were electrofished in each stream (one at the mouth of the stream and three within the stream) in August and October 2000, 2001, 2002. 0+ salmon parr were absent from seven streams, present at the mouth of 11 streams, and present within only two streams, both of which were probably spawning streams. Older parr migrated upstream into most streams and their highest densities were usually found in streams flowing directly into the spawning habitat in the three largest tributaries of the Tana or the river itself. Juvenile salmon were sparse or absent in streams flowing into smaller tributaries. Most streams with high parr densities were those of dense riparian vegetation that provided terrestrial invertebrates as drift food for the salmon parr, cover for fish, cooler stream temperatures in summer, and food for benthic stream invertebrates that were also a source of food for the parr.     

Prey selectivity and ontogenetic diet shift of the globally invasive western mosquitofish (Gambusia affinis) in agriculturally impacted streams

The dietary breadth of invaders can influence their success, and having a wide dietary niche can facilitate the spread and survival of invaders under a variety of resource scenarios. The western mosquitofish (Gambusia affinis) is a globally distributed freshwater invasive fish. The spread of G. affinis is associated with agricultural land use, although the trophic role it plays in degraded systems is not well understood. We analysed the invertebrate community in 11 stream reaches in the North Island of New Zealand, in catchments spanning a range (45%–90%) of agricultural land use to determine how prey availability changes with land use. We then analysed the gut contents of 400 G. affinis from the 11 sites to determine how diet varied with prey availability and ontogeny. Invertebrate communities varied along the agricultural land‐use gradient, both in regard to taxonomic richness and community composition. G. affinis consumed a wide variety of food items with invertebrates being the most dominant, in particular Culicidae, Copepods and amphipods were the most commonly consumed invertebrates. There was also an ontogenetic diet shift from microinvertebrates (Cladocera, Copepods and diatoms) to larger invertebrates, including Culicidae, amphipods and terrestrial invertebrates. G. affinis are capable of consuming a wide variety of prey in agricultural streams; their preferred prey are generally pollution‐tolerant taxa commonly found in degraded streams. Having a large level of dietary plasticity coupled with preferring prey that are often associated with degraded systems likely facilitates to the spread of one of the most widely distributed freshwater invasive fish.     

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.     

Riparian vegetation management for water temperature regulation: implications for the production of macroinvertebrate prey of salmonids

The effect of riparian vegetation on the seasonal availability of macroinvertebrate prey for salmonids was examined. Although prey composition changed significantly between sampling seasons, overall prey density remained similar. It is often assumed that there is greater benthic production within streams without riparian vegetation, but this assumption was found to be dependent on stream size. The presence of riparian vegetation increased benthic macroinvertebrate density, but only in larger streams. Greater densities of prey of terrestrial origin were found in the drift of streams with riparian cover but only in spring. An unforeseen contributor to potential prey items for salmonids in streams with riparian vegetation was the increased presence of drifting prey of aquatic origin. The results further support riparian management as a viable tool for regulating instream temperatures without impacting on macroinvertebrate density. The presence of riparian vegetation can even increase salmonid prey availability in some instances.     

Direct and indirect effects of riparian canopy on juvenile Atlantic salmon,Salmo salar,and brown trout,Salmo trutta,in south‐west Ireland

Abstract Dense riparian tree canopy is generally found to have a negative effect on salmonid populations. Canopy can affect fish both directly via trophic impacts and its effects as cover, and indirectly via its effects on the distribution of instream vegetation. This study examined the impact of riparian canopy on the ecology of juvenile Atlantic salmon, Salmo salar L., and brown trout, Salmo trutta L., in south‐west Ireland. Riparian canopy reduced the density, length and gut contents of juvenile salmon, but not brown trout. The negative relationship between canopy cover and fish size and feeding was strongly influenced by the abundance of instream macrophytes, which in turn varied as a function of catchment water chemistry. As a management strategy, the reduction of dense riparian canopy is unlikely to have the same effect on juvenile salmonids on all streams within an ecoregion because of differences in catchment‐wide factors.     

Salmonid colonization of new streams in Glacier Bay National Park, Alaska

Abstract. Following the rapid recession of a neoglacial ice sheet within the last 250 years, colonization of recently deglaciated streams by salmonid fishes was investigated in Glacier Bay National Park, south-eastern Alaska. The primary factors governing the establishment, species diversity composition and abundance of salmonids in Glacier Bay streams were water temperature, sediment loading and stream discharge. No salmonids were found in the turbid meltwater streams emerging from retreating ice. Coho, Oncorhynchus kisutch (Walbaum), and sockeye, Oncorhynchus nerka (Walbaum). salmon and Dolly Vardcn, Salvelinus malma (Walbaum), charr were the first salmonids to colonize the youngest clearwater stream. Juvenile Dolly Varden were more abundant than juvenile coho salmon in the most recently formed clearwater stream because of the characteristic absence of pool habital. Densities of juvenile coho salmon were six times greater in a stream with a series of lakes compared with a stream of similar age without lakes. Future advancement of salmonid stocks will probably depend upon the rate and extent of the development of riparian vegetation and inputs of large woody debris from the developing forest to provide further instream cover, habitat variation and channel stabilization.     

Competitive interactions among multiple non‐native salmonids and two populations of Atlantic salmon

Competitive interactions with non‐native species can have negative impacts on the conservation of native species, resulting in chronic stress and reduced survival. Here, juvenile Atlantic salmon (Salmo salar) from two allopatric populations (Sebago and LaHave) that are being used for reintroduction into Lake Ontario were placed into semi‐natural stream tanks with four non‐native salmonid competitors that are established in Ontario streams: brown trout (S. trutta), rainbow trout (Oncorhynchus mykiss), Chinook salmon (O. tshawytscha) and coho salmon (O. kisutch). Brown trout and rainbow trout reduced the survival and fitness‐related traits of Atlantic salmon, whereas Chinook salmon and coho salmon had no impact on these traits. These data support theories on ecological niche overlap and link differences in observed aggression levels with competitive outcomes. Measurements of circulating hormones indicated that the Atlantic salmon were not chronically stressed nor had a change in social status at the 10‐month time point in the semi‐natural stream tanks. Additionally, the Sebago population was better able to coexist with the non‐native salmonids than the LaHave population. Certain populations of Atlantic salmon may thus be more suitable for some environments of the juvenile stream phase for the reintroduction into Lake Ontario.     

Juvenile coho salmon movement,growth and survival in a coastal basin of southern Oregon

Juvenile salmonids display highly variable spatial and temporal patterns of early dispersal that are influenced by density‐dependent and density‐independent factors. Although juvenile coho salmon (Oncorhynchus kisutch) movement patterns in streams and their relationship with body mass and growth have been examined in previous studies, most observations were limited to one season or one stream section. In this study, we monitored the movement of juvenile coho salmon throughout their period of residence in a coastal basin to identify prevalent dispersal strategies and their relationships with body mass, growth rates and survival. Our results revealed seasonally and spatially variable movement patterns. Juvenile coho salmon that dispersed to tidally affected reaches soon after emergence remained more mobile and expressed lower site fidelity than those individuals that remained in upper riverine reaches. We did not detect significantly different growth rates between sedentary and mobile individuals. Although a greater proportion of sedentary than mobile fish survived winter to emigrate from the creek in the spring, reach of residence at the onset of winter influenced these survival estimates. Hence, apparent summer‐to‐smolt survival for mobile individuals was greater than for sedentary fish in tidally influenced reaches, whereas in riverine reaches the sedentary strategy seemed to be favoured. Our research identified complex movement patterns that reflect phenotypic and life history variation, and underscores the importance of maintaining diverse freshwater and estuarine habitats that support juvenile coho salmon before marine migration.     

Effect of fine wood on juvenile brown trout behaviour in experimental stream channels

In‐stream wood can increase shelter availability and prey abundance for stream‐living fish such as brown trout, Salmo trutta, but the input of wood to streams has decreased in recent years due to harvesting of riparian vegetation. During the last decades, fine wood (FW) has been increasingly used for biofuel, and the input of FW to streams may therefore decrease. Although effects of in‐stream FW have not been studied as extensively as those of large wood (LW), it is probably important as shelter for small‐sized trout. In a laboratory stream experiment, we tested the behavioural response of young‐of‐the‐year wild brown trout to three densities of FW, with trout tested alone and in groups of four. Video recordings were used to measure the proportion of time allocated to sheltering, cruising and foraging, as well as the number of aggressive interactions and prey attacks. Cruising activity increased with decreasing FW density and was higher in the four‐fish groups than when fish were alone. Foraging decreased and time spent sheltering in FW increased with increasing FW density. Our study shows that juvenile trout activity is higher in higher fish densities and that trout response to FW is related to FW density and differs from the response to LW as reported by others.     

Do network relationships matter? Comparing network and instream habitat variables to explain densities of juvenile coho salmon (Oncorhynchus kisutch) in mid‐coastal Oregon,USA

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Food habits and marine survival of juvenile Chinook and coho salmon from marine waters of Southeast Alaska

Little is known about the food habits of juvenile Chinook (Oncorhynchus tshawytscha) and coho (Oncorhynchus kisutch) salmon in marine environments of Alaska, or whether their diets may have contributed to extremely high marine survival rates for coho salmon from Southeast Alaska and much more modest survival rates for Southeast Alaskan Chinook salmon. To address these issues, we documented the spatial and temporal variability of diets of both species collected from marine waters of Southeast Alaska during summers of 1997–2000. Food habits were similar: major prey items of both species included fishes, crab larvae, hyperiid amphipods, insects, and euphausiids. Multivariate analyses of diet composition indicated that the most distinct groups were formed at the smallest spatial and temporal scales (the haul), although groups also formed at larger scales, such as by month or habitat type. Our expectations for how food habits would influence survival were only partially supported. As predicted, Southeast Alaskan coho salmon had more prey in their stomachs overall [1.8% of body weight (BW)] and proportionally far fewer empty stomachs (0.7%) than either Alaskan Chinook (1.4% BW, 5.1% empty) or coho salmon from other regions. However, contrary to our expectations, coho salmon diets contained surprisingly few fish (49% by weight). Apparently, Alaskan coho salmon achieved extremely high marine survival rates despite a diet consisting largely of small, less energetically‐efficient crustacean prey. Our results suggest that diet quantity (how much is eaten) rather than diet quality (what is eaten) is important to marine survival.     

Episodic predation of mammals by stream fishes in a boreal river basin

Stream‐dwelling fish typically feed on small prey items, such as benthic invertebrates, but maintain the capacity to opportunistically feed on rare, large‐bodied prey when available. However, consumption of particularly large prey is typically viewed as isolated events that are not organised spatially or temporally across watersheds. We assessed the occurrence of small mammals in the stomach contents of rainbow trout (Oncorhynchus mykiss) and Arctic grayling (Thymallus arcticus) over 13 years in the Wood River basin, Alaska (59°34′N, 158°48′W). Shrews (Sorex spp.) were the dominant mammals observed episodically every 2–3 years in the stomach contents of fish. Notably, shrew consumption was correlated in both Arctic grayling and rainbow trout within individual streams, and across the river basin in several subwatersheds. Predators of shrews were usually the largest individuals within each population, suggesting that smaller fish are gape‐limited and that dominant fish monopolised mammal prey. On average, 24% (11–38%) of Arctic grayling >298 mm (fork length) and rainbow trout >290 mm contained mammal prey during peak years. Small mammal populations often cycle every 2–5 years with well‐known functional and numerical effects for terrestrial predators, a dynamic that may be reflected in our 13 years data set of diet contents for aquatic predators. Although numerically infrequent when averaged over time, small mammal subsidies to streams may be episodically important to the energy budgets of long‐lived consumers in freshwater environments.     

Modelling the reestablishment of coho salmon (Oncorhynchus kisutch) in Klamath River tributaries after dam removal

The planned removal of four dams on the Klamath River (anticipated 2024) will be the largest river restoration effort ever undertaken on the planet. Dam removal will restore access to >50 km of the Klamath River mainstem for coho salmon, but mainstem habitat may not be suitable for rearing juvenile coho salmon. Instead, small tributaries may provide most rearing habitat for reestablishing coho salmon. We used four approaches to evaluate six Klamath River tributaries above existing dams to assess their potential to support juvenile coho salmon: (1) We measured summer temperature regimes and evaluated thermal suitability. (2) We applied an Intrinsic Potential (IP) model to evaluate large-scale geomorphological constraints on coho salmon habitat. (3) We used the Habitat Limiting Factors Model (HLFM) to estimate rearing capacity for juveniles given current habitat conditions. (4) We developed an occupancy model using data from reference tributaries to predict coho salmon rearing distribution. All six streams had summer temperatures cooler than the mainstem Klamath River. However, five of the streams have barriers that will restrict coho salmon to within 5 km of the confluence with the Klamath River and two were disconnected mid-summer. Despite these constraints, the tributaries will likely produce coho salmon. Most streams had high IP in their lower reaches, the HLFM model estimated a total capacity of 105,000 juvenile coho salmon, and the occupancy model predicted juvenile coho salmon will rear throughout the accessible reaches. Protection and habitat enhancement for these tributaries will be important for coho salmon reestablishment post-dam removal.     

The effects of stream canopy management on macroinvertebrate communities and juvenile salmonid production in a chalk stream

Abstract  The effects of changes in shading (through riparian canopy removal and re-growth) on juvenile salmon, Salmo salar L., trout, Salmo trutta L., and grayling, Thymallus thymallus (L.) populations, and macroinvertebrate biomass and species composition in a chalk stream in southern England were examined. Low levels of in-stream weed growth, because of shading by closed tree canopy, diminished macroinvertebrate production and diversity. 0+ salmon and trout had lower densities under closed canopy, relative to adjacent open sites with substantial weed cover, where fish were also found to be larger. Canopy removal positively affected the growth of aquatic macrophytes and the availability of potential prey for juvenile salmonids. The findings have implications for the management of chalk streams, in particular, that riparian tree canopy should be managed to prevent complete closure, and excessive cutting of weed should be avoided where salmon production is below sustainable levels.     

Cohort‐specific variation in juvenile coho salmon habitat use

In this study, we examined summer and fall freshwater rearing habitat use by juvenile coho salmon (Oncorhynchus kisutch) in the quickly urbanising Big Lake drainage in south‐central Alaska. Habitat use was assessed by regressing fish count data against habitat survey information across thirty study sites using generalised linear mixed models. Habitat associations were examined by age‐0 and age‐1+ cohorts separately, providing an opportunity to compare habitat use across different juvenile coho salmon life stages during freshwater rearing. Regression results indicated that the age‐0 cohorts were strongly associated with shallow, wide stream reaches with in‐stream vegetation, whereas age‐1+ cohorts were associated with deeper stream reaches. Furthermore, associations between fork length and habitat characteristics suggest cohort‐specific habitat use patterns are distinct from those attributable to fish size. Habitat use information generated from this study is being used to guide optimal fish passage restoration planning in the Big Lake drainage. Evidence for habitat use partitioning by age cohort during freshwater juvenile rearing indicates that pooling age cohorts into a single “juvenile” stage for the purposes of watershed management may mask important habitat use dynamics.