Seasonal influence of brook trout and mottled sculpin on lower trophic levels in an Appalachian stream

1. In some situations fish have strong top‐down effects in stream communities while in others they seem to be relatively unimportant. Differences in the impact of fish may depend on a variety of factors including the foraging mode of the fish, interactions among fish species and temporal variation in environmental conditions and species interactions. 2. We investigated the effect of brook trout (Salvelinus fontinalis) and mottled sculpin (Cottus bairdi) on lower trophic levels in Appalachian streams and whether or not interactions between these fish changed their influence. Mesocosms were placed in a headwater stream in a randomized complete block design. Within blocks, mesocosms were randomly assigned to one of the following treatments: (i) no fish; (ii) sculpin only; (iii) trout only and (iv) both sculpin and trout. Fish biomass was the same in all three fish treatments. Invertebrate density and algal biomass in mesocosms were determined after 3 weeks. We repeated the experiment in the autumn, spring and summer to test for seasonality of fish effects. 3. The effect of fish on invertebrate assemblages was seasonal and depended on prey identity. Sculpin strongly suppressed grazer abundance in spring while trout had little effect on grazers in any season. The influence of both fish on insect predators was similar and relatively constant across seasons. We found little evidence of an interaction between sculpin and trout that strongly influenced their effect on prey across seasons. 4. None of the fish treatments influenced algal biomass during any of the seasons. Algal growth was also seasonal, with a two‐ to four‐fold increase in algal biomass in spring compared to autumn and summer. 5. Our results indicate that benthic and drift feeding fish differ in their effects on some, but not all prey. Furthermore, fish effects on prey were strongly seasonal for some, but not all prey types. While the temporal context is not commonly considered, our results indicate seasonality can be an important component of predator–prey interactions in streams.     

Food partitioning between coexisting Atlantic salmon and brook trout in the Sainte‐Marguerite River ecosystem, Quebec

Food resource partitioning between similar‐sized, sympatric Atlantic salmon Salmo salar and brook trout Salvelinus fontinalis was examined as a possible mechanism enabling their coexistence in a stream (Allaire) of the Sainte‐Marguerite River ecosystem, Quebec, Canada. Fish stomach contents and invertebrate drift were collected concurrently during three diel cycles in August to September 1996. The food and feeding habits of an allopatric brook trout population in a nearby stream (Epinette) were studied for comparison. The diel feeding rhythms of the two coexisting fish species were similar. The composition of their diet, however, showed significant differences. Atlantic salmon predominantly (60–90%) fed on aquatic insects, mainly Ephemeroptera (35–60% of the diet). The brook trout mostly (50–80%) fed upon the allochthonous terrestrial insects (mainly adults of Coleoptera, Hymenoptera and Diptera) which comprised 5–40% of the stream drift. The allopatric brook trout fed opportunistically on the more abundant aquatic insects and terrestrial insects rarely formed 25% of its diet. The allopatric trout fed nearly twice as much as the sympatric brook trout during a day. The results suggest that the differences in feeding by brook trout in the two streams (with and without Atlantic salmon) are the result of inter‐specific interaction with Atlantic salmon and are not related to the differences in food availability between the two streams. Food resource partitioning between Atlantic salmon and brook trout may be viewed as an adaptive response resulting in a greater exploitation of available resources and coexistence.     

Food selection by brook trout in a subalpine stream

The diet and food selection of brook trout (Salvelinus fontinalis) and macroinvertebrate drift are described for a high altitude stream (3 205 m above mean sea level) adjacent to the alpine zone on the Snowy Range in the Medicine Bow National Forest, Wyoming. Diet composition differed between brook trout of two length classes, 150 mm and > 150 mm in total length. The number of prey organisms per stomach generally declined and the proportion of terrestrial prey items increased from July to September, 1985. Fish of both length classes selected for Trichoptera and Coleoptera over the three sampling months, and fish > 150 mm also selected for terrestrial insects in August and September.The Unit is jointly supported by the University of Wyoming, the Wyoming Game and Fish Department, and the US Fish and Wildlife ServiceThe Unit is jointly supported by the University of Wyoming, the Wyoming Game and Fish Department, and the US Fish and Wildlife Service     

Pelagic food web interactions among benthic invertebrates and trout in mountain lakes

1. Benthic chironomid larvae and the amphipod Gammarus lacustris have been observed in the pelagic habitats of many mountain lakes. The main goal of this study was to determine if chironomid larvae and gammarids potentially affect predator–prey and nutrient dynamics in pelagic food webs of mountain lakes. 2. Eighty‐six mountain lakes were surveyed in Alberta and eastern British Columbia during the years 1965–1984, 1991–2004 and 2005–2007. Pelagic chironomid larvae were found in 86% of these lakes, and pelagic gammarids were found in 29% of lakes. Densities of pelagic chironomid larvae were 92% lower in lakes with pelagic gammarids and 76% lower in lakes with trout (P < 0.05). Intraguild predation of trout on gammarids appeared to reduce predation pressure on chironomid larvae. Gammarids consumed in vitro about 1 chironomid per gammarid per day or about 20% of their body mass in chironomid biomass per day. 3. Concentrations of total dissolved P and N, particulate C, and chlorophyll‐a increased with increasing densities of pelagic gammarids and chironomid larvae in situ (R² = 0.14 ± 0.19 SD, P < 0.1) and in vitro (P < 0.001). 4. Our findings suggest that gammarids and chironomid larvae are linked as predators and prey in pelagic food webs, possibly stimulating phytoplankton abundance via nutrient release.     

Shifts in abundance and growth of slimy sculpin in response to changes in the predator population in an arctic Alaskan lake

Lake trout (Salvelinus namaycush) in Toolik Lake are tightly coupled to the benthos, since they have no pelagic forage fishes. Slimy sculpins (Cottus cognatus) are a prey of lake trout and the soft sediment chironomids are an important prey for the sculpin. Our previous work showed that the median size of lake trout in Toolik Lake had decreased significantly between 1977 and 1986, and smaller lake trout are likely to be less effective as sculpin predators. Using our historic data on the slimy sculpin population from 1978, we took advantage of the recent change in the predator community to examine for subsequent changes in the sculpin community. Between 1978 and 1987, the percentage of slimy sculpin caught in the soft sediments has increased (25% to 39.5%). In 1987 there was a significant difference in the mean weight of sculpin caught on different substrates. The mean individual weight of sculpins increased from the nearshore rock area to the rock/soft-sediment interface to the soft sediments. There was no difference in mean individual weight with habitat in 1978. The mean total length at age for slimy sculpins during this time has also increased significantly. We suggest that the risk of predation while foraging in the soft sediments has declined. The increased use of the soft sediment area appears to have contributed to their increased growth, likely due to greater food abundance.     

The role of trout in stream food webs: integrating evidence from field surveys and experiments

1. We evaluated the effects of brown trout on boreal stream food webs using field surveys and enclosure/exclosure experiments. Experimental results were related to prey preference of uncaged trout in the same stream, as well as to a survey of macroinvertebrate densities in streams with vs. without trout. Finally, we assessed the generality of our findings by examining salmonid predation on three groups of macroinvertebrate prey (chironomid midges, epibenthic grazers, invertebrate predators) in a meta-analysis. 2. In a preliminary experiment, invertebrate predators showed a strong negative response to trout, whereas chironomids benefited from trout presence. In the main experiment, trout impact increased with prey size. Trout had the strongest effect on invertebrate predators and cased caddis larvae, whereas Baetis mayfly and chironomid larvae were unaffected. Trout impact on the largest prey seemed mainly consumptive, because prey emigration rates were low and independent of fish presence. Despite strong effects on macroinvertebrates, trout did not induce a trophic cascade on periphyton. Uncaged trout showed a strong preference for the largest prey items (predatory invertebrates and aerial prey), whereas Baetis mayflies and chironomids were avoided by trout. 3. Densities of invertebrate predators were significantly higher in troutless streams. Baetis mayflies also were less abundant in trout streams, whereas densities of chironomids were positively, although non-significantly, related to trout presence. Meta-analysis showed a strong negative impact of trout on invertebrate predators, a negative but variable impact on mobile grazers (mainly mayfly larvae) and a slightly positive impact on chironomid larvae. 4. Being size-selective predators, salmonid fishes have a strong impact on the largest prey types available, and this effect spans several domains of scale. Discrepancies between our experimental findings and those from the field survey and meta-analysis show, however, that for most lotic prey, small-scale experiments do not reflect fish impact reliably at stream-wide scales. 5. Our findings suggest that small-scale experiments will be useful only if the experimental results are evaluated carefully against natural history information about the experimental system and interacting species across a wide array of spatial scales.     

Biotic interactions modify multiple‐stressor effects on juvenile brown trout in an experimental stream food web

Agricultural land use results in multiple stressors affecting stream ecosystems. Flow reduction due to water abstraction, elevated levels of nutrients and chemical contaminants are common agricultural stressors worldwide. Concurrently, stream ecosystems are also increasingly affected by climate change. Interactions among multiple co‐occurring stressors result in biological responses that cannot be predicted from single‐stressor effects (i.e. synergisms and antagonisms). At the ecosystem level, multiple‐stressor effects can be further modified by biotic interactions (e.g. trophic interactions). We conducted a field experiment using 128 flow‐through stream mesocosms to examine the individual and combined effects of water abstraction, nutrient enrichment and elevated levels of the nitrification inhibitor dicyandiamide (DCD) on survival, condition and gut content of juvenile brown trout and on benthic abundance of their invertebrate prey. Flow velocity reduction decreased fish survival (?12% compared to controls) and condition (?8% compared to initial condition), whereas effects of nutrient and DCD additions and interactions among these stressors were not significant. Negative effects of flow velocity reduction on fish survival and condition were consistent with effects on fish gut content (?25% compared to controls) and abundance of dominant invertebrate prey (?30% compared to controls), suggesting a negative metabolic balance driving fish mortality and condition decline, which was confirmed by structural equation modelling. Fish mortality under reduced flow velocity increased as maximal daily water temperatures approached the upper limit of their tolerance range, reflecting synergistic interactions between these stressors. Our study highlights the importance of indirect stressor effects such as those transferred through trophic interactions, which need to be considered when assessing and managing fish populations and stream food webs in multiple‐stressor situations. However, in real streams, compensatory mechanisms and behavioural responses, as well as seasonal and spatial variation, may alter the intensity of stressor effects and the sensitivity of trout populations.     

Importance of terrestrial subsidies for native brook trout in Appalachian intermittent streams

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Influence of growth rate on white muscle dynamics in rainbow trout and brook trout

In trout, fast growth stimulated white muscle fibre hypertrophy ( P 0·001) and hyperplasia ( P <0·01) in outer fibres but not in deep fibres. Glycogen was most prevalent in outer fibres ( P <0·01) and in brook trout ( P <0·01) that on average had three to four times larger fibres than rainbow trout.     

Aquatic and terrestrial invertebrate drift in southern Appalachian Mountain streams: implications for trout food resources

1. We characterised aquatic and terrestrial invertebrate drift in six south‐western North Carolina streams and their implications for trout production. Streams of this region typically have low standing stock and production of trout because of low benthic productivity. However, little is known about the contribution of terrestrial invertebrates entering drift, the factors that affect these inputs (including season, diel period and riparian cover type), or the energetic contribution of drift to trout. 2. Eight sites were sampled in streams with four riparian cover types. Drift samples were collected at sunrise, midday and sunset; and in spring, early summer, late summer and autumn. The importance of drift for trout production was assessed using literature estimates of annual benthic production in the southern Appalachians, ecotrophic coefficients and food conversion efficiencies. 3. Abundance and biomass of terrestrial invertebrate inputs and drifting aquatic larvae were typically highest in spring and early summer. Aquatic larval abundances were greater than terrestrial invertebrates during these seasons and terrestrial invertebrate biomass was greater than aquatic larval biomass in the autumn. Drift rates of aquatic larval abundance and biomass were greatest at sunset. Inputs of terrestrial invertebrate biomass were greater than aquatic larvae at midday. Terrestrial invertebrate abundances were highest in streams with open canopies and streams adjacent to pasture with limited forest canopy. 4. We estimate the combination of benthic invertebrate production and terrestrial invertebrate inputs can support 3.3–18.2 g (wet weight) m⁻² year⁻¹ of trout, which is generally lower than values considered productive [10.0–30.0 g (wet weight) m⁻² year⁻¹]. 5. Our results indicate terrestrial invertebrates can be an important energy source for trout in these streams, but trout production is still low. Any management activities that attempt to increase trout production should assess trout food resources and ensure their availability.     

Stable isotope evidence of trophic interactions between introduced brook trout Salvelinus fontinalis and native brown trout Salmo trutta in a mountain stream of south-west France

The potential trophic impact of introduced brook trout Salvelinus fontinalis on native brown trout Salmo trutta in a mountain stream (south-west France) was investigated using stable isotope analysis (SIA). The isotopic signatures (δ¹³C and δ¹⁵N) of S. fontinalis were similar regardless of the absence or presence of S. trutta , and SIA mixing models revealed that S. fontinalis diet consisted mainly of terrestrial invertebrates. Conversely, a significant shift in S. trutta isotopic signatures (depletion of 1·6‰δ¹³C and enrichment of 0·6‰δ¹⁵N) was observed in sympatry with S. fontinalis ; this may be due to a dietary shift towards terrestrial invertebrates. Contrary to an expected dietary divergence in sympatry, an elevated level of dietary overlap was observed between the non-native and native salmonids when in co-occurrence. This dietary convergence is more likely to be due to behavioural interactions than to variations in food availability or fish displacements.     

Diet of brown trout in relation to variation in abundance and size of pelagic fish prey

The seasonal diet of a predator, brown trout Salmo trutta [total length ( L _T) 17–69 cm] and simultaneous density and size‐structure of prey populations, vendace Coregonus albula and smelt Osmerus eperlanus (4–16 cm L _T), in a large boreal lake were analysed and compared in 2001 and 2002. The upper L _T limit for consumed prey was c . 40% of the predator L _T. All brown trout, however, preferred small (<10 cm L _T) and avoided large (≥10 cm L _T) prey. The results also suggested that equal densities of similar‐sized (4–10 cm L _T) fish of the two prey species led to random foraging on these species by brown trout, but if either one of the prey species predominated (>50%) in the lake, brown trout shifted to foraging on this species almost exclusively. Brown trout diets thus reflected the density dynamics of the two alternative prey species.     

Climate‐change winners and losers: stream macroinvertebrates of a submontane region in Central Europe

1. Freshwater ecosystems will be profoundly affected by global climate change, especially those in mountainous areas, which are known to be particularly vulnerable to warming temperatures. We modelled impacts of climate change on the distribution ranges of 38 species of benthic stream macroinvertebrates from nine macroinvertebrate orders covering all river zones from the headwaters to large river reaches. 2. Species altitudinal shifts as well as range changes up to the year 2080 were simulated using the A2a and B2a Intergovernmental Panel on Climate Change climate‐warming scenarios. Presence‐only species distribution models were constructed for a stream network in Germany’s lower mountain ranges by means of consensus projections of four algorithms, as implemented in the BIOMOD package in R (GLM, GAM, GBM and ANN). 3. Species were predicted to shift an average of 122 and 83 m up in altitude along the river continuum by the year 2080 under the A2a and B2a climate‐warming scenarios, respectively. No correlation between altitudinal shifts and mean annual air temperature of species’ occurrence could be detected. 4. Depending on the climate‐warming scenario, most or all (97% for A2a and 100% for B2a) of the macroinvertebrate species investigated were predicted to survive under climate change in the study area. Ranges were predicted to contract for species that currently occur in streams with low annual mean air temperatures but expand for species that inhabit rivers where air temperatures are higher. 5. Our models predict that novel climate conditions will reorganise species composition and community structure along the river continuum. Possible effects are discussed, including significant reductions in population size of headwater species, eventually leading to a loss of genetic diversity. A shift in river species composition is likely to enhance the establishment of non‐native macroinvertebrates in the lower reaches of the river continuum.     

Selective foraging on terrestrial invertebrates by rainbow trout in a forested headwater stream in northern Japan

The important contribution of terrestrial invertebrates to the energy budget of drift-foraging fishes has been well documented in many forested headwater streams. However, relatively little attention has been focused on the behavioral mechanisms behind such intensive exploitation. We tested for the hypothesis that active prey selection by fishes would be an important determinant of terrestrial invertebrates contribution to fish diets in a forested headwater stream in northern Japan. Rainbow trout, Oncorhynchus mykiss, were estimated to consume 57.12 mg m^–2 day^–1 (dry mass) terrestrial invertebrates, 77% of their total input (73.89 mg m^–2 day^–1), there being high selectivity for the former from stream drift. Both the falling input and drift of terrestrial invertebrates peaked at around dusk, decreasing dramatically toward midnight. In contrast, both aquatic insect adults and benthic invertebrates showed pronounced nocturnal drift. Because the prey consumption rates of rainbow trout were high at dawn and dusk, decreasing around midnight, the greater contribution of terrestrial invertebrates to trout diet was regarded as being partly influenced by the difference in diel periodicity of availability among prey categories. In addition, selectivity also depended upon differences in individual prey size among aquatic insect adults, and benthic and terrestrial invertebrates, the last category being largest in both the stream drift and the trout diets. We concluded that differences in both the timing of supplies and prey size among the three prey categories were the primary factors behind the selective foraging on terrestrial invertebrates by rainbow trout.     

Detection and apparent survival of PIT‐tagged stream fish in winter

Environmental fluctuations exert strong control on behavior, survival, and fitness of stream biota. Technical improvements increasingly allow for tracking the response of large numbers of individuals to environmental fluctuations, for instance, by remote detection of animals equipped with PIT (passive integrated transponder) tags. PIT tags were implanted into 393 juvenile and adult brown trout Salmo trutta L. and European sculpin Cottus gobio L. in a boreal stream subjected to considerable ice formation. With weekly trackings over 6 months, we quantified apparent survival and detection probability in relation to biological, environmental, and methodological factors. Individuals with a higher physical condition in autumn showed a higher apparent survival; this pattern was consistent across all species and age classes. Detection probability decreased with increasing thickness of the surface ice layer; this effect was most pronounced for juvenile trout and benthic‐living sculpin, both tagged with smaller‐sized tags. Detection probability was reduced in structurally complex habitats. Our study demonstrates that apparent survival and particularly detection probability may show pronounced spatiotemporal variation. In order to compare results from different sampling occasions and sites, a good knowledge of the study site and of the regulating factors is crucial.     

The St. Lawrence River in winter: population structure,biomass, and pattern of its primary and secondary food web components

Plankton and benthic invertebrate populations were examined along a 160 km section of the St. Lawrence River between Lake Ontario and Waddington, New York during a period of ice cover. Algal biomass was low in winter with diatoms and cryptomonads the most prominent forms. Zooplankton biomass was about one-tenth the algal standing crop and cyclopoid copepods predominated. Phytoplankton and zooplankton biomass declined downriver providing evidence that most plankton was imported from Lake Ontario. Benthic invertebrate standing crop was greatest near Lake Ontario where molluscs were abundant. The downriver macroinvertebrate community shifted to coarse particle feeders, namely annelids and chironomids. Beds of accumulated detrital material derived primarily from dieback of macrophytes were particularly rich in benthic invertebrates.     

Brown trout Salmo trutta redd superimposition by spawning Lampetra species in a lowland stream

Reproductive interaction between sympatric lampreys and salmonids was studied. The superimposition of brown trout Salmo trutta redds by spring-spawning river lamprey Lampetra fluviatilis and brook lamprey Lampetra planeri was examined in a small lowland stream of western Lithuania. A high superimposition rate of S. trutta redds by both L. fluviatilis (up to 83%) and L. planeri (up to 48%) was found, when the spawning intensity of Lampetra spp. was high. The occurrence of this phenomenon is the result of the overlap in the spawning habitat preferences at the reach-scale and at the microhabitat scale for the three species. One of the main requirements for Lampetra spp. spawning site selection was the negative streambed slope, an essential trait of the pool-riffle transitional zone. The structure of the salmonid redd created a considerable negative microhabitat slope suitable for Lampetra spp. spawning, which put the redds under a higher susceptibility to be superimposed. The timing of Lampetra spp. spawning overlapped closely with the emergence of S. trutta fry, suggesting a probable ecological effect of superimposition on S. trutta in the pre-emergent and emerging stages.     

Diel feeding periodicity, daily ration and prey selectivity in juvenile brown trout in a subarctic river

Feeding of age-1 brown trout Salmo trutta in a third-order river in northern Finland was usually highest in the twilight hours and lowest around midday. Diel periodicity in food intake was less distinct and rarely significant for age-0 trout. Daily rations declined seasonally, being lowest in October, and highest in June (age-1 trout) or early August (age-0 trout). Prey selection did not differ between day and night, but differences between age classes and sampling dates were distinct. Age-0 trout preferred Ephemerella nymphs in summer and Micrasema larvae later in the season. Age-1 trout fed selectively on caddis larvae on all sample dates. Aerial insects and Baetis nymphs were avoided by both age classes on most occasions. These patterns of preference are probably explained by a trend towards epibenthic feeding, which may be a predominant foraging mode in some trout populations. Nevertheless, repuscular peaks in feeding periodicity suggest that trout were able to capitalize on the increased availability of drifting prey in the twilight, especially in the early summer months.     

Stabilizing mechanisms in a food web with an introduced omnivore

Intraguild predation (IGP) is an omnivorous food web configuration in which the top predator consumes both a competitor (consumer) and a second prey that it shares with the competitor. This omnivorous configuration occurs frequently in food webs, but theory suggests that it is unstable unless stabilizing mechanisms exist that can decrease the strength of the omnivore and consumer interaction. Although these mechanisms have been documented in native food webs, little is known about whether they operate in the context of an introduced species. Here, we study a marine mussel aquaculture system where the introduction of omnivorous mussels should generate an unstable food web that favors the extinction of the consumer, yet it persists. Using field and laboratory approaches, we searched for stabilizing mechanisms that could reduce interaction strengths in the food web. While field zooplankton counts suggested that mussels influence the composition and abundance of copepods, stable isotope results indicated that life‐history omnivory and cannibalism facilitated the availability of prey refugia, and reduced competition and the interaction strength between the mussel omnivore and zooplankton consumers. In laboratory experiments, however, we found no evidence of adaptive feeding which could weaken predator–consumer interactions. Our food web study suggests that the impact of an introduced omnivore may not only depend on its interaction with native species but also on the availability of stabilizing mechanisms that alter the strength of those interactions.