Quantifying the direct and indirect effects of flow‐related disturbance on stream fish assemblages

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Comparing stream-specific to generalized temperature models to guide salmonid management in a changing climate

Global climate change is predicted to increase air and stream temperatures and alter thermal habitat suitability for growth and survival of coldwater fishes, including brook charr (Salvelinus fontinalis), brown trout (Salmo trutta), and rainbow trout (Oncorhynchus mykiss). In a changing climate, accurate stream temperature modeling is increasingly important for sustainable salmonid management throughout the world. However, finite resource availability (e.g. funding, personnel) drives a tradeoff between thermal model accuracy and efficiency (i.e. cost-effective applicability at management-relevant spatial extents). Using different projected climate change scenarios, we compared the accuracy and efficiency of stream-specific and generalized (i.e. region-specific) temperature models for coldwater salmonids within and outside the State of Michigan, USA, a region with long-term stream temperature data and productive coldwater fisheries. Projected stream temperature warming between 2016 and 2056 ranged from 0.1 to 3.8 °C in groundwater-dominated streams and 0.2–6.8 °C in surface-runoff dominated systems in the State of Michigan. Despite their generally lower accuracy in predicting exact stream temperatures, generalized models accurately projected salmonid thermal habitat suitability in 82% of groundwater-dominated streams, including those with brook charr (80% accuracy), brown trout (89% accuracy), and rainbow trout (75% accuracy). In contrast, generalized models predicted thermal habitat suitability in runoff-dominated streams with much lower accuracy (54%). These results suggest that, amidst climate change and constraints in resource availability, generalized models are appropriate to forecast thermal conditions in groundwater-dominated streams within and outside Michigan and inform regional-level salmonid management strategies that are practical for coldwater fisheries managers, policy makers, and the public. We recommend fisheries professionals reserve resource-intensive stream-specific models for runoff-dominated systems containing high-priority fisheries resources (e.g. trophy individuals, endangered species) that will be directly impacted by projected stream warming.     

Effects of Large Woody Debris Addition on Stream Habitat and Brook Trout Populations in Appalachian Streams

Large woody debris (LWD) was added to eight streams in the central Appalachians of West Virginia to determine if stream habitat could be enhanced and brook trout (Salvelinus fontinalis) populations increased. Brook trout populations were assessed one year prior to habitat manipulation and 3 years post-habitat manipulation. LWD was added by felling approximately 15 trees per 300 m stream reach. Four of the streams had LWD added to one 300 m reach with 300 m unmanipulated reaches upstream and downstream of the manipulated reach to observe within-stream effects of LWD additions on brook trout density. The remaining four streams had LWD added to three 300 m reaches and these streams were compared to those with only a single 300 m manipulated reach to observe the effects of the extent of habitat manipulation on brook trout density. New pools were formed by the addition of LWD, but overall pool area did not increase significantly in reaches where LWD was added. The relatively high gradient and coarse substrate of these streams may have precluded the added LWD from having a significant influence on stream channel morphology and habitat complexity. No pools were formed in the highest gradient stream, while the stream with the most pools formed had the lowest gradient. Brook trout populations fluctuated following habitat manipulations, and there was no overall effect of the LWD additions on within-stream variability in brook trout density. When there were significant differences among-streams with different extents of LWD additions, those streams receiving LWD additions over a large extent had the greatest brook trout densities. The full potential of added LWD to change stream habitat and influence on brook trout populations may take more time to develop than the 3 years post-manipulation period of this study.     

Influences of logging history and riparian forest characteristics on macroinvertebrates and brook trout (Salvelinus fontinalis) in headwater streams (New Hampshire, U.S.A.)

1. Logging can strongly affect stream macroinvertebrate communities, but the direction and magnitude of these effects and their implications for trout abundance are frequently region‐specific and difficult to predict. 2. In first‐order streams in northern New England (U.S.A.) representing a chronosequence of logging history (<2 to >80 years since logging), we measured riparian forest conditions, stream macroinvertebrate community characteristics and brook trout (Salvelinus fontinalis) abundance. Principal component analysis was used to collapse forest data into two independent variables representing variation in logging history, riparian forest structure and canopy cover. We used these data to test whether logging history and associated forest conditions were significant predictors of macroinvertebrate abundance and functional feeding group composition, and whether brook trout abundance was related to logging‐associated variation in invertebrate communities. 3. Catchments with high PC1 scores (recently logged, high‐density stands with low mean tree diameter) and low PC2 scores (low canopy cover) had significantly higher total macroinvertebrate abundance, particularly with respect to chironomid larvae (low PC2 scores) and invertebrates in the grazer functional feeding group (high PC1 scores). In contrast, proportional representation of macroinvertebrates in the shredder functional feeding group increased with time since logging and canopy cover (high PC2 scores). Brook trout density and biomass was significantly greater in young, recently logged stands (high PC1 scores) and was positively related to overall macroinvertebrate abundance. In addition, three variables – trout density, invertebrate abundance and shredder abundance – successfully discriminated between streams that were less‐impacted versus more‐impacted by forestry. 4. These results indicate that timber harvest in northern New England headwater streams may shift shredder‐dominated macroinvertebrate communities supporting low trout abundance to a grazer/chironomid‐dominated macroinvertebrate community supporting higher trout abundance. However, while local effects on brook trout abundance may be positive, these benefits may be outweighed by negative effects of brook trout on co‐occurring species, as well as impairment of habitat quality downstream. Research testing the generality of these patterns will improve understanding of how aquatic ecosystems respond to anthropogenic and natural trajectories of forest change.     

Grazing management influences the subsidy of terrestrial prey to trout in central Rocky Mountain streams (USA)

1. Research in forest and grassland ecosystems indicates that terrestrial invertebrates that fall into streams can be an important prey resource for fish, providing about 50% of their annual energy and having strong effects on growth and abundance. However, the indirect effects of land uses like cattle grazing on this important prey subsidy for stream salmonids are unclear. 2. During summer 2007, we compared the effects of three commonly used grazing systems on terrestrial invertebrate inputs to streams in northern Colorado and their use by trout. Cattle graze individual pastures for about 120 days under traditional season‐long grazing (SLG), about 35–45 days under simple rotational grazing and 10–20 days under intensive rotational grazing in this region. We also compared these effects to a fourth group of sites grazed only by wildlife (i.e. no livestock use). 3. Overall, rotational grazing management (either simple or intensive), resulted in more riparian vegetation, greater inputs of terrestrial invertebrates, greater biomass of terrestrial invertebrate prey in trout diets, a higher input compared to trout metabolic demand and more trout biomass than SLG. However, these differences were frequently not statistically significant owing to high variability, especially for trout diets and biomass. 4. Despite the inherent variability, riparian vegetation and terrestrial invertebrates entering streams and in trout diets at sites managed for rotational grazing were similar to sites managed for wildlife grazing only. 5. These results indicate that rotational grazing systems can be effective for maintaining levels of terrestrial invertebrate subsidies to streams necessary to support robust trout populations. However, factors influencing the effect of riparian grazing on stream subsidies are both spatially variable and complex, owing to differences in microclimate, invertebrate and plant populations and the efforts of ranchers to tailor grazing systems to specific riparian pastures.     

What predicts the use by brook trout (Salvelinus fontinalis) of terrestrial invertebrate subsidies in headwater streams?

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Indirect effects of predatory trout on organic matter processing in detritus-based stream food webs

Indirect effects of predators on basal resources in allochthonous-based food webs are poorly understood. We investigated indirect effects of predatory brown trout ( Salmo trutta ) on detritus dynamics in southern beech ( Nothofagus spp.) forest streams in New Zealand through predation on the obligate detritivore, Zelandopsyche ingens (Trichoptera, Oeconesidae). Trout presence/absence and Z. ingens density were manipulated in flow-through tanks to investigate the lethal and sub-lethal effects of trout on litter processing by Z. ingens . An experiment that allowed trout access to Z. ingens showed trout predation reduced densities of Z. ingens resulting in slower breakdown of coarse particulate organic matter (CPOM) and reduced production of fine particulate organic matter (FPOM). A second experiment that prevented trout access to Z. ingens , but allowed the transmission of trout cues, resulted in no change in litter processing rates in the presence of trout. Litter processing rates were higher in high Z. ingens density treatments compared to low density treatments. Thus, trout effects on litter processing were due to reduced Z. ingens densities, not trout-induced modifications to Z. ingens feeding behaviour. Field assays of litter processing rates using artificial leaf packs in natural streams showed significant reductions in CPOM loss in trout streams compared to fishless streams. Z. ingens dominated biomass in fishless stream leaf packs, but a facultative shredder, Olinga feredayi , dominated trout stream leaf packs. Thus, the absence of Z. ingens drove differences in processing rates between trout and fishless streams and the indirect effects of trout on litter processing observed in mesocosms were evident in complex, natural food webs. Overall our study provides evidence that predators can influence resource dynamics in donor-controlled food webs through their effects on consumers.     

Contribution of woody debris to trout habitat modification in small streams in secondary deciduous forest, northern Japan

Field studies to examine the influence of woody debris on rainbow trout (Oncorhynchus mykiss) abundance through habitat modification were conducted in two small streams, the Horonai and Uenae streams, running through secondary deciduous forest in south-western Hokkaido, northern Japan. Reach-based woody debris volume (total woody debris volume per 100 m² of study reach) was significantly correlated with the total basal area of riparian stands along the margins of the Horonai stream, but no significant relationship was evident between them for the Uenae stream. This inconsistency between the streams was considered to be a result of the difference in stream size (width, depth and discharge). Woody debris was the principal agent for pool formation, although it had a far smaller volume than that found in streams draining old-growth coniferous forest in North America, where most of the previous studies have been carried out. Untransported debris pieces of larger volume more effectively contributed to pool formation than smaller transported pieces. The volume of individual debris scour pools was positively correlated with the volume of woody debris associated with each. Similarly, reach-based pool volume increased with total woody debris volume, but the relationship was less clear in the Uenae stream, having more abundant transported woody debris than the Horonai stream. The biomass of rainbow trout in individual pools, which were regarded as the most preferred habitat type for stream salmonids, was correlated with pool volume. A positive relationship also existed between reach-based standing crop and pool volume. These results revealed that secondary deciduous forest, like old-growth coniferous forest, plays an important role in enhancing the carrying capacity for rainbow trout by supplying woody debris which promoted preferred habitat formation.     

Increases in disturbance and reductions in habitat size interact to suppress predator body size

Food webs are strongly size‐structured so will be vulnerable to changes in environmental factors that affect large predators. However, mechanistic understanding of environmental controls of top predator size is poorly developed. We used streams to investigate how predator body size is altered by three fundamental climate change stressors: reductions in habitat size, increases in disturbance and warmer temperatures. Using new survey data from 74 streams, we showed that habitat size and disturbance were the most important stressors influencing predator body size. A synergistic interaction between that habitat size and disturbance due to flooding meant the sizes of predatory fishes peaked in large, benign habitats and their body size decreased as habitats became either smaller or harsher. These patterns were supported by experiments indicating that habitat‐size reductions and increased flood disturbance decreased both the abundance and biomass of large predators. This research indicates that interacting climate change stressors can influence predator body size, resulting in smaller predators than would be predicted from examining an environmental factor in isolation. Thus, climate‐induced changes to key interacting environmental factors are likely to have synergistic impacts on predator body size which, because of their influence on the strength of biological interactions, will have far‐reaching effects on food‐web responses to global environmental change.     

Future climate change effects on US forest composition may offset benefits of reduced atmospheric deposition of N and S

Climate change and atmospheric deposition of nitrogen (N) and sulfur (S) are important drivers of forest demography. Here we apply previously derived growth and survival responses for 94 tree species, representing >90% of the contiguous US forest basal area, to project how changes in mean annual temperature, precipitation, and N and S deposition from 20 different future scenarios may affect forest composition to 2100. We find that under the low climate change scenario (RCP 4.5), reductions in aboveground tree biomass from higher temperatures are roughly offset by increases in aboveground tree biomass from reductions in N and S deposition. However, under the higher climate change scenario (RCP 8.5) the decreases from climate change overwhelm increases from reductions in N and S deposition. These broad trends underlie wide variation among species. We found averaged across temperature scenarios the relative abundance of 60 species were projected to decrease more than 5% and 20 species were projected to increase more than 5%; and reductions of N and S deposition led to a decrease for 13 species and an increase for 40 species. This suggests large shifts in the composition of US forests in the future. Negative climate effects were mostly from elevated temperature and were not offset by scenarios with wetter conditions. We found that by 2100 an estimated 1 billion trees under the RCP 4.5 scenario and 20 billion trees under the RCP 8.5 scenario may be pushed outside the temperature record upon which these relationships were derived. These results may not fully capture future changes in forest composition as several other factors were not included. Overall efforts to reduce atmospheric deposition of N and S will likely be insufficient to overcome climate change impacts on forest demography across much of the United States unless we adhere to the low climate change scenario.     

The impact of trout on galaxiid fishes in New Zealand

Compared with the effect of invaders on the native terrestrial fauna of New Zealand, interactions between native fishes and introduced trout (sports fish in the genera Salmo, Oncorhynchus and Salvelinus) are less well known and there have been fewer efforts to remedy their effects. Trout have caused widespread reductions in the distribution and abundance of native galaxiid fishes, a family dominated by threatened species. The effects have been most severe on non-diadromous species (those lacking a marine migratory stage), which are commonly eliminated from streams by trout. Galaxiid populations in lakes, and those with migratory ?whitebait? stages, have also been affected, but the extent of the impacts are less understood. The mechanisms controlling negative interactions between trout and native fish, and how the environment modifies those interactions, will be important for future management. Experiments and field comparisons indicate size-specific predation by trout is the main driver of negative interactions. Large trout (>150 mm long) do the greatest damage and small galaxiids (those with adult sizes <150 mm long) are the most at risk. The fry stage of non-diadromous galaxiids is particularly vulnerable. Despite galaxiid fry production in some trout-invaded reaches, often no fry survive making them population ?sinks? that must be sustained by adult dispersal. Trout are also associated with changes in galaxiid behaviour and alterations to stream benthic communities. However, effects on galaxiid growth and fecundity have been little studied. Recent work also indicates that habitat conditions, especially floods, low flows and natural acidity, can mediate trout?galaxiid interactions. We argue that managers should be more proactive in their response to the plight of galaxiids, and we identify avenues of research that will benefit native fish conservation activities in the future.     

Top-down and bottom-up influences on populations of a stream detritivore

1. Knowledge of the influence of predatory fish in detritus‐based stream food webs is poor. We tested whether larval abundance of the New Zealand leaf‐shredding caddisfly, Zelandopsyche ingens (family Oeconesidae), was affected by the presence of predatory brown trout, Salmo trutta and the abundance of their primary detrital resource (Nothofagus leaves). 2. The density of Z. ingens and the biomass of leaves were determined in seven fishless streams and four trout streams in the Cass region, central South Island, on four occasions spanning 5 years. 3. Physicochemical conditions were similar in trout and fishless streams, but ancova indicated that Z. ingens numbers were positively related to leaf biomass and that caddisfly numbers were significantly greater in fishless streams than trout streams for any given biomass of leaf. The cases of trout stream larvae were also heavier per unit length than those in fishless streams. 4. Our results provide evidence for both top‐down and bottom‐up influences on a detritus‐based stream food web. Although stream detritivores may benefit from a habitat that provides both food and a degree of protection from predators, top‐down effects of predators on detritivore population abundance were still important. Thus, detrital resource availability may determine maximum attainable population size, whereas predation is likely to reduce the population to a level below that.     

Adapting streams for climate change using riparian broadleaf trees and its consequences for stream salmonids

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The emergence period of sea trout fry in a Lake District stream correlates with the North Atlantic Oscillation

The date of fry emergence over 30 years in a sea trout nursery stream, predicted by an individual-based model, correlated significantly ( r =0·660, P <0·001) with an index of the North Atlantic Oscillation. Water temperature is the main driving variable in the model and stream temperature also correlated significantly ( r =0·662, P <0·001) with the index, providing a probable causal link. Therefore, the inter-annual variations in emergence may not be unique to this one stream, but may be typical of other trout streams with similar climatic conditions.     

Linking climate change projections for an Alaskan watershed to future coho salmon production

Climate change is predicted to dramatically change hydrologic processes across Alaska, but estimates of how these impacts will influence specific watersheds and aquatic species are lacking. Here, we linked climate, hydrology, and habitat models within a coho salmon (Oncorhynchus kisutch) population model to assess how projected climate change could affect survival at each freshwater life stage and, in turn, production of coho salmon smolts in three subwatersheds of the Chuitna (Chuit) River watershed, Alaska. Based on future climate scenarios and projections from a three‐dimensional hydrology model, we simulated coho smolt production over a 20‐year span at the end of the century (2080–2100). The direction (i.e., positive vs. negative) and magnitude of changes in smolt production varied substantially by climate scenario and subwatershed. Projected smolt production decreased in all three subwatersheds under the minimum air temperature and maximum precipitation scenario due to elevated peak flows and a resulting 98% reduction in egg‐to‐fry survival. In contrast, the maximum air temperature and minimum precipitation scenario led to an increase in smolt production in all three subwatersheds through an increase in fry survival. Other climate change scenarios led to mixed responses, with projected smolt production increasing and decreasing in different subwatersheds. Our analysis highlights the complexity inherent in predicting climate‐change‐related impacts to salmon populations and demonstrates that population effects may depend on interactions between the relative magnitude of hydrologic and thermal changes and their interactions with features of the local habitat.     

Evaluating ecosystem effects of climate change on tropical island streams using high spatial and temporal resolution sampling regimes

Climate change is expected to alter precipitation patterns worldwide, which will affect streamflow in riverine ecosystems. It is vital to understand the impacts of projected flow variations, especially in tropical regions where the effects of climate change are expected to be one of the earliest to emerge. Space‐for‐time substitutions have been successful at predicting effects of climate change in terrestrial systems by using a spatial gradient to mimic the projected temporal change. However, concerns have been raised that the spatial variability in these models might not reflect the temporal variability. We utilized a well‐constrained rainfall gradient on Hawaii Island to determine (a) how predicted decreases in flow and increases in flow variability affect stream food resources and consumers and (b) if using a high temporal (monthly, four streams) or a high spatial (annual, eight streams) resolution sampling scheme would alter the results of a space‐for‐time substitution. Declines in benthic and suspended resource quantity (10‐ to 40‐fold) and quality (shift from macrophyte to leaf litter dominated) contributed to 35‐fold decreases in macroinvertebrate biomass with predicted changes in the magnitude and variability in the flow. Invertebrate composition switched from caddisflies and damselflies to taxa with faster turnover rates (mosquitoes, copepods). Changes in resource and consumer composition patterns were stronger with high temporal resolution sampling. However, trends and ranges of results did not differ between the two sampling regimes, indicating that a suitable, well‐constrained spatial gradient is an appropriate tool for examining temporal change. Our study is the first to investigate resource to community wide effects of climate change on tropical streams on a spatial and temporal scale. We determined that predicted flow alterations would decrease stream resource and consumer quantity and quality, which can alter stream function, as well as biomass and habitat for freshwater, marine, and terrestrial consumers dependent on these resources.     

The effects of intra- and inter-specific competition on the distribution of stocked juvenile Atlantic salmon, Salmo salar L., in relation to depth and gradient in an upland trout, Salmo trutta L., stream

The relative effects of inter- and intra-specific competition on the distribution of stocked salmon in relation to depth and gradient were investigated in an upland stream during two summer sampling periods. The stream was divided into two areas by an impassable fish barrier, and trout were removed from the upstream section prior to 2 years of salmon stocking. A small amount of trout re-immigration to the cleared area occurred. Under sympatric conditions in the downstream section the fry of both species were significantly more abundant in shallow water, whereas the yearling and older fish tended to inhabit the deeper, slow-flowing areas. Under near allopatric conditions in the cleared section salmon fry changed their distribution to include deeper areas, with shallow, fast-flowing water becoming the least preferred habitat. This trend for a wide distribution of salmon fry over all the available habitat in the upstream section was maintained in the second year when parr were present but trout remained at very low densities. It was therefore concluded that high inter-specific competition from trout was responsible for restricting the distribution of salmon fry to shallow habitat in the control area. Intra-specific competition from older salmon apparently only affected the growth and survival of salmon fry. The regulating mechanisms involved in these inter- and intra-specific effects are discussed in terms of competition for stream resources and predation. Recommendations based on the findings are made for stocking and habitat management of salmonid waters.     

The influence of introduced trout on the benthic communities of paired headwater streams in the Sierra Nevada of California

1. Non‐native trout have been stocked in streams and lakes worldwide largely without knowledge of the consequences for native ecosystems. Although trout have been introduced widely throughout the Sierra Nevada of California, U.S.A., fishless streams and their communities of native invertebrates persist in some high elevation areas, providing an opportunity to study the effects of trout introductions on natural fishless stream communities. 2. We compared algal biomass and cover, organic matter levels and invertebrate assemblages in 21 natural fishless headwater streams with 21 paired nearby streams containing stocked trout in Yosemite National Park. 3. Although environmental conditions and particulate organic matter levels did not differ between the fishless and trout streams, algal biomass (as chlorophyll a concentration) and macroalgal cover were, on average, approximately two times and five times higher, respectively, in streams containing trout. 4. There were no differences in the overall densities of invertebrates in fishless versus paired trout streams; however, invertebrate richness (after rarefaction), evenness, and Simpson and Shannon diversities were 10–20% higher in fishless than in trout streams. 5. The densities of invertebrates belonging to the scraper‐algivore and predator functional feeding guilds were higher, and those for the collector‐gatherer guild lower, in fishless than trout streams, but there was considerable variation in the effects of trout on specific taxa within functional feeding groups. 6. We found that the densities of 10 of 50 common native invertebrate taxa (found in more than half of the stream pairs) were reduced in trout compared to fishless streams. A similar number of rarer taxa also were absent or less abundant in the presence of trout. Many of the taxa that declined with trout were conspicuous forms (by size and behaviour) whose native habitats are primarily high elevation montane streams above the original range of trout. 7. Only a few taxa increased in the presence of trout, possibly benefiting from reductions in their competitors and predators by trout predation. 8. These field studies provide catchment‐scale evidence showing the selective influence of introduced trout on stream invertebrate and algal communities. Removal of trout from targeted headwater streams may promote the recovery of native taxa, community structure and trophic organisation.     

Climate change effects on upland stream macroinvertebrates over a 25-year period

Climate change effects on some ecosystems are still poorly known, particularly where they interact with other climatic phenomena or stressors. We used data spanning 25 years (1981–2005) from temperate headwaters at Llyn Brianne (UK) to test three hypotheses: (1) stream macroinvertebrates vary with winter climate; (2) ecological effects attributable to directional climate change and the North Atlantic Oscillation (NAO) are distinguishable and (3) climatic effects on macroinvertebrates depend on whether streams are impacted by acidification. Positive (i.e. warmer, wetter) NAO phases were accompanied by reduced interannual stability (=similarity) in macroinvertebrate assemblage in all streams, but associated variations in composition occurred only in acid moorland. The NAO and directional climate change together explained 70% of interannual variation in temperature, but forest and moorland streams warmed respectively by 1.4 and 1.7°C (P<0.001) between 1981 and 2005 after accounting for NAO effects. Significant responses among macroinvertebrates were confined to circumneutral streams, where future thermal projections (+1, +2, +3°C) suggested considerable change. Spring macroinvertebrate abundance might decline by 21% for every 1°C rise. Although many core species could persist if temperature gain reached 3°C, 4–10 mostly scarce taxa (5–12% of the species pool) would risk local extinction. Temperature increase in Wales approaches this magnitude by the 2050s under the Hadley HadCM3 scenarios. These results support all three hypotheses and illustrate how headwater stream ecosystems are sensitive to climate change. Altered composition and abundance could affect conservation and ecological function, with the NAO compounding climate change effects during positive phases. We suggest that acidification, in impacted streams, overrides climatic effects on macroinvertebrates by simplifying assemblages and reducing richness. Climatic processes might, nevertheless, exacerbate acidification or offset biological recovery.     

Population changes after two years of salmon (Salmo salar L.) stocking in upland trout (Salmo tmtta L.) streams

The survival of salmon stocked in upland trout streams in the presence of salmon parr was found to be only about half the value recorded when trout alone made up the resident stock. Changes in the trout population were also recorded following the two years of salmon stocking, and these suggested that the presence of salmon parr may also influence trout fry survival. The findings are discussed in the context of habitat competition and total stream holding capacity.