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

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

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Terrestrial and marine trophic pathways support young-of-year growth in a nearshore Arctic fish

River discharge supplies nearshore communities with a terrestrial carbon source that is often reflected in invertebrate and fish consumers. Recent studies in the Beaufort Sea have documented widespread terrestrial carbon use among invertebrates, but only limited use among nearshore fish consumers. Here, we examine the carbon source and diet of rapidly growing young-of-year Arctic cisco (Coregonus autumnalis) using stable isotope values (δ¹³C and δ¹⁵N) from muscle and diet analysis (stomach contents) during a critical and previously unsampled life stage. Stable isotope values (δ¹⁵N and δ¹³C) may differentiate between terrestrial and marine sources and integrate over longer time frames (weeks). Diet analysis provides species-specific information, but only from recent foraging (days). Average δ¹³C for all individuals was ?25.7 ‰, with the smallest individuals possessing significantly depleted δ¹³C values indicative of a stronger reliance of terrestrial carbon sources as compared to larger individuals. Average δ¹⁵N for all individuals was 10.4 ‰, with little variation among individuals. As fish length increased, the proportion of offshore Calanus prey and neritic Mysis prey increased. Rapid young-of-year growth in Arctic cisco appears to use terrestrial carbon sources obtained by consuming a mixture of neritic and offshore zooplankton. Shifts in the magnitude or phenology of river discharge and the delivery of terrestrial carbon may alter the ecology of nearshore fish consumers.     

Daily energy intake and growth of piscivorous brown trout,Salmo trutta

• 1 The chief objectives were to determine the daily energy intake and growth of piscivorous brown trout (Salmo trutta), and to compare the observed values with those expected from models developed previously for brown trout feeding on freshwater invertebrates. Energy budgets for individual fish were obtained from experiments with 40 trout (initial live weight 250–318 g) bred from wild parents, and kept at five constant temperatures (5, 10, 13, 15, 18 °C) and 100% oxygen saturation. Each trout was fed to satiation on freshly killed sticklebacks (Gasterosteus aculeatus) over a period of 42 days.
• 2 Energy intake (C_IN cal day^‐1) and growth (C_G cal day^‐1) were measured directly and energy losses (C_Q cal day^‐1) were estimated by difference (C_Q = C_IN ‐ C_G). All three variables increased with temperature. A model previously used to predict the daily energy intake (C_IN(INV)) of trout feeding to satiation on invertebrates was adapted, by changing only one parameter, to provide an excellent model (R² = 0.998) for predicting the mean daily energy intake (C_IN(ST)) for the piscivorous trout. Values of C_IN(ST) were 58% (range 48–67%) higher than those for C_IN(INV). A simple model was also developed to estimate mean daily energy losses for piscivorous trout (R² = 0.999). Both models were combined to provide excellent estimates of the daily energy gain (growth) of the piscivorous trout, and this was about three times that for trout feeding on invertebrates. The optimum temperature for maximum growth in energy terms increased from 13.9 °C for trout feeding on invertebrates to 17.0 °C (range 16.6–17.4 °C) for piscivorous trout.
• 3 The models are basically an extension of those developed for trout feeding on invertebrates. They show clearly how energy intake, growth, and the optimum temperature for growth increase markedly when trout change their diet from invertebrates to fish. The implications of this are discussed and it is shown that, in theory, these increases should continue if a more energy‐rich diet was utilised by the 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.     

Reliance of brown trout on terrestrial prey varies with season but not fish density

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Terrestrial contributions to Afrotropical aquatic food webs: The Congo River case

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Stable isotopes indicate that zebra mussels (Dreissena polymorpha) increase dependence of lake food webs on littoral energy sources

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Consumer‐resource stoichiometry as a predictor of trophic discrimination (Δ13C, Δ15N) in aquatic invertebrates

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The influence of water content and leaf anatomy on carbon isotope discrimination and photosynthesis in Sphagnum

The relative effect of diffusional resistance due to water films (r_wf) and leaf anatomy (r_p) on rates of net photosynthesis and on-line measures of carbon isotope discrimination (Δ=Δδ¹³C) was investigated in Sphagnum. Sphagnum species differ in the exposure of photosynthetic cells at the leaf surface. In S. affine, photosynthetic cells are widely exposed at the surface, whereas in S. magellanicum, photo-synthetic cells are enclosed within water-filled hyaline cells. This difference is expected to lead to variation in diffusive resistance within leaves (r_p). Net photosynthesis and on-line Δ were measured at two water contents: greenhouse water content (wet) and blotted dry (dry). Without correcting for respiration, on-line Δ values differed significantly between wet (23.7%_o) and dry (30.9%_o) plants. However, there was no significant difference between species means and no species × water content interaction. Corrections for respiration lowered Δ values by approximately 8.1%_o and reduced the mean difference to 3.1%_o, but did not alter the rank order of treatments. Net photosynthesis also decreased by 16% in wet plants, but there was no significant difference between the two species. In addition, five populations of S. affine and S. magellanicum grown in a common garden were analysed for their organic matter carbon isotope composition (δ¹³C). These values varied more within each species (0.9–1.2%_o) than between the two species (0.6%_o). Therefore, we conclude that variation in surface water films leads to a greater difference in resistance to CO₂ uptake and carbon isotope discrimination than that due to variation in leaf anatomical properties in Sphagnum.     

Effects of woody debris and the supply of terrestrial invertebrates on the diet and growth of brown trout (Salmo trutta) in a boreal stream

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Carbon sources and trophic structure in a macrophyte‐dominated polyculture pond assessed by stable‐isotope analysis

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Top-down and bottom-up processes in grassland and forested streams

The influence of predatory fish on the structure of stream food webs may be altered by the presence of forest canopy cover, and consequent differences in allochthonous inputs and primary production. Eight sites containing introduced brown trout (Salmo trutta) and eight sites that did not were sampled in the Cass region, South Island, New Zealand. For each predator category, half the sites were located in southern beech (Nothofagus) forest patches (range of canopy cover, 65–90%) and the other half were in tussock grassland. Food resources used by two dominant herbivores-detritivores were assessed using stable isotopes. ¹³C/¹²C ratios were obtained for coarse particulate organic matter (CPOM), fine particulate organic matter (FPOM), algal dominated biofilm from rocks, and larvae of Deleatidium (Ephemeroptera) and Olinga (Trichoptera). Total abundance and biomass of macroinvertebrates did not differ between streams with and without trout, but were significantly higher at grassland sites than forested sites. However, taxon richness and species composition differed substantially between trout and no-trout sites, irrespective of whether streams were located in forest or not. Trout streams typically contained more taxa, had low biomass of predatory invertebrates and large shredders, but a high proportion of consumers with cases or shells. The standing stock of CPOM was higher at forested sites, but there was less FPOM and more algae at sites with trout, regardless of the presence or absence of forest cover. The stable carbon isotope range for biofilm on rocks was broad and encompassed the narrow CPOM and FPOM ranges. At trout sites, carbon isotope ratios of Deleatidium, the most abundant invertebrate primary consumer, were closely related to biofilm values, but no relationship was found at no-trout sites where algal biomass was much lower. These results support a role for both bottom-up and top-down processes in controlling the structure of the stream communities studied, but indicate that predatory fish and forest cover had largely independent effects.     

Linking Intertidal and Subtidal Food Webs: Consumer-Mediated Transport of Intertidal Benthic Microalgal Carbon

We examined stable carbon and nitrogen isotope ratios for a large variety of consumers in intertidal and subtidal habitats, and their potential primary food sources [i.e., microphytobenthos (MPB), phytoplankton, and Phragmites australis] in a coastal bay system, Yeoja Bay of Korea, to test the hypothesis that the transfer of intertidal MPB-derived organic carbon to the subtidal food web can be mediated by motile consumers. Compared to a narrow δ¹³C range (−18 to −16‰) of offshore consumers, a broad δ¹³C range (−18 to −12‰) of both intertidal and subtidal consumers indicated that ¹³C-enriched sources of organic matter are an important trophic source to coastal consumers. In the intertidal areas, δ¹³C of most consumers overlapped with or was ¹³C-enriched relative to MPB. Despite the scarcity of MPB in the subtidal, highly motile consumers in subtidal habitat had nearly identical δ¹³C range with many intertidal foragers (including crustaceans and fish), overlapping with the range of MPB. In contrast, δ¹³C values of many sedentary benthic invertebrates in the subtidal areas were similar to those of offshore consumers and more ¹³C-depleted than motile foragers, indicating high dependence on phytoplankton-derived carbon. The isotopic mixing model calculation confirms that the majority of motile consumers and also some of subtidal sedentary ones depend on intertidal MPB for more than a half of their tissue carbon. Finally, although further quantitative estimates are needed, these results suggest that direct foraging by motile consumers on intertidal areas, and thereby biological transport of MPB-derived organic carbon to the subtidal areas, may provide important trophic connection between intertidal production and the nearshore shallow subtidal food webs.     

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.     

Artificial light as a disturbance to light‐naïve streams

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Resource use by salmonids in riverine,lacustrine and marine environments: Evidence from stable isotope analysis

Synopsis We measured stable isotope ratios (δ¹³C and δ¹⁵N) of invertebrates, Atlantic salmon, Salmo salar, and brook trout, Salvelinus fontinalis, in three distinct freshwater environments (headwater tributary, ultra-oligotrophic lake, and main-stem river) in the Western Brook system, Newfoundland, Canada. Large differences in the stable carbon signatures of invertebrates allowed the identification of organic matter assimilation from each environment by resident parr and migrating smolts. Brook trout captured in the headwater tributary in June had a carbon signature characteristic of the tributary, while those collected in August had enriched ¹³C (maximum = −15.6‰) and ¹⁵N (maximum = 12.8‰) values. These enriched carbon and nitrogen signatures were indicative of foraging at sea. There was a low correlation between δ¹³C and δ¹⁵N (r² = 0.198) for individual fish that was likely due to the confounding influence of trout feeding in the lake and the lower main-stem of the river, where δ¹³C of food sources was high but δ¹⁵N was low. Smolts emigrating from Western Brook Pond where they had been foraging (based on lacustrine carbon signatures) were significantly larger than those emigrating from a nursery brook and the main river in the same basin, despite having the same median age. These results suggest better growth opportunities in the lake environment. Trout fork length was positively correlated with δ¹³C and δ¹⁵N, demonstrating that larger individuals had been feeding outside the brook. These results support previous studies that found increased growth potential for salmonids in lacustrine and marine environments, and further, indicate possible adaptive advantages for salmonid movement away from natal brooks.     

A Comparative Study by δ13C-Analysis of Crassulacean Acid Metabolism (CAM) in Kalanchoë (Crassulaceae) Species of Africa and Madagascar

The carbon isotope ratios (δ¹³C values) of samples of Kalanchoë species collected in Africa were compared with previous data obtained with species from Madagascar. In contrast to the Malagasy species which cover the whole range of δ¹³C values from ?10 to ? 30%_o, indicating high inter- and intraspecific diversity of CAM performance, in the African species nearly all δ¹³C values were less negative than ?18%₀. Thus, in the African species the CAM behaviour is characterized by CO₂ uptake proceeding mainly during the night. The distribution of δ¹³C values among the species clearly mirrors the taxonomic groups and the three sections of the genus Kalanchoë sensu lato. The Kitchingia section comprises only groups having CAM with a high proportion of carbon acquisition by the C₃-pathway of photosynthesis. The same holds true for the first three groups of the Bryophyllum section, whereas in the following groups of the section CAM with CO₂ proceeding mainly during the night is common. The latter CAM mode is typical also for the majority of groups and species in the section Eukalanchoë. The African Kalanchoë species belong to the Eukalanchoë section, whereas in Madagascar all three sections are abundant. The data support the view that the centre of adaptive radiation of the genus is located in Madagascar. They also suggest that high CAM variability is abundant in the more primitive taxa of the genus, whereas the phylogenetically more derived taxa show a stereotype CAM with CO₂ uptake taking place only during the night.     

Impact of eutrophication on the carbon stable‐isotopic baseline of benthic invertebrates in two deep soft‐water lakes

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Effect of riparian land use on contributions of terrestrial invertebrates to streams

Since terrestrial invertebrates are often consumed by stream fishes, land-use practices that influence the input of terrestrial invertebrates to streams are predicted to have consequences for fish production. We studied the effect of riparian land-use regime on terrestrial invertebrate inputs by estimating the biomass, abundance and taxonomic richness of terrestrial invertebrate drift from 15 streams draining catchments with three different riparian land-use regimes and vegetation types: intensive grazing — exotic pasture grasses (4 streams), extensive grazing — native tussock grasses (6 streams), reserve — native forest (5 streams). Terrestrial invertebrate drift was sampled from replicated stream reaches enclosed by two 1 mm mesh drift nets that spanned the entire channel. The mean biomass of terrestrial invertebrates that entered tussock grassland (12 mg ash-free dry mass m^–2 d^–1) and forest streams (6 mg AFDM m^–2 d^–1) was not significantly different (p > 0.05). However, biomass estimated for tussock grassland and forest streams was significantly higher than biomass that entered pasture streams (1 mg AFDM m^–2 d^–1). Mean abundance and richness of drifting terrestrial invertebrates was not significantly different among land-use types. Winged insects contributed more biomass than wingless invertebrates to both pasture and tussock grassland streams. Winged and wingless invertebrates contributed equally to biomass entering forest streams. Land use was a useful variable explaining landscape-level patterns of terrestrial invertebrate input for New Zealand streams. Evidence from this study suggests that riparian land-use regime will have important influences on the availability of terrestrial invertebrates to stream fishes.