Subsidies of energy and material from the riparian zone have large impacts on recipient stream habitats. Human-induced changes, such as deforestation, may profoundly affect these pathways. However, the strength of individual factors on stream ecosystems is poorly understood since the factors involved often interact in complex ways. We isolated two of these factors, manipulating the flux of terrestrial input and the intensity of light in a 2×2 factorial design, where we followed the growth and diet of two size-classes of brown trout (Salmo trutta) and the development of periphyton, grazer macroinvertebrates, terrestrial invertebrate inputs, and drift in twelve 20 m long enclosed stream reaches in a five-month-long experiment in a boreal coniferous forest stream. We found that light intensity, which was artificially increased 2.5 times above ambient levels, had an effect on grazer density, but no detectable effect on chlorophyll a biomass. We also found a seasonal effect on the amount of drift and that the reduction of terrestrial prey input, accomplished by covering enclosures with transparent plastic, had a negative impact on the amount of terrestrial invertebrates in the drift. Further, trout growth was strongly seasonal and followed the same pattern as drift biomass, and the reduction of terrestrial prey input had a negative effect on trout growth. Diet analysis was consistent with growth differences, showing that trout in open enclosures consumed relatively more terrestrial prey in summer than trout living in covered enclosures. We also predicted ontogenetic differences in the diet and growth of old and young trout, where we expected old fish to be more affected by the terrestrial prey reduction, but we found little evidence of ontogenetic differences. Overall, our results showed that reduced terrestrial prey inputs, as would be expected from forest harvesting, shaped differences in the growth and diet of the top predator, brown trout.
Terrestrial invertebrate subsidies are believed to be important energy sources for drift-feeding salmonids. Despite this, size-specific use of and efficiency in procuring this resource have not been studied to any great extent. Therefore, we measured the functional responses of three size classes of wild brown trout Salmo trutta (0+, 1+ and ‡2+) when fed either benthic-(Gammarus sp.) or surface-drifting prey (Musca domestica) in laboratory experiments. To test for size-specific prey preferences, both benthic and surface prey were presented simultaneously by presenting the fish with a constant density of benthic prey and a variable density of surface prey. The results showed that the functional response of 0+ trout differed significantly from the larger size classes, with 0+ fish having the lowest capture rates. Capture rates did not differ significantly between prey types. In experiments when both prey items were presented simultaneously, capture rate differed significantly between size classes, with larger trout having higher capture rates than smaller trout. However, capture rates within each size class did not change with prey density or prey composition. The two-prey experiments also showed that 1+ trout ate significantly more surface-drifting prey than 0+ trout. In contrast, there was no difference between 0+ and ‡2+ trout. Analyses of the vertical position of the fish in the water column corroborated size-specific foraging results: larger trout remained in the upper part of the water column between attacks on surface prey more often than smaller trout, which tended to seek refuge at the bottom between attacks. These size-specific differences in foraging and vertical position suggest that larger trout may be able to use surface-drifting prey to a greater extent than smaller conspecifics.
Summary 1. Changes in riparian vegetation owing to forest harvesting may affect the input of large wood, a major structural element, to streams. Studies of large wood impacts on stream fish have focused on population‐level responses, whereas little attention has been given to how wood affects fish behaviour. 2. In a laboratory stream experiment, we tested how two size classes of brown trout, Salmo trutta, (mean size of 85 and 125 mm), alone and together, responded to a gradient of large wood in terms of activity, foraging on terrestrial drift and interactions between conspecifics. 3. The results showed that the presence of large wood significantly reduced the overall activity of the fish, the number of agonistic interactions between individuals and the proportion of captured prey. However, activity decreased relatively more than the proportion of captured prey, resulting in a significant positive net effect of wood on the number of prey captures per time spent active (PTA). This indicates that trout living in habitats with high wood density may have a higher net energy gain than trout living in habitats with less wood. 4. There were no observable size‐class differences in the benefits of large wood or in the utilisation of surface‐drifting terrestrial prey. 5. These results suggest that the presence of large wood may be an important factor shaping stream communities and that a lack of structural complexity may decrease energy gain, increase agonistic interactions and, consequently, lower the production of brown trout.
Summary Changes to the riparian vegetation of forest streams during timber harvesting may have considerable impacts on stream biota, but few studies have attempted to separate the effects of individual factors that are altered during clear‐felling operations. We studied the effects of large wood and terrestrial invertebrate supply, two factors affected by forest harvesting, on the growth and diet of two size classes of brown trout (Salmo trutta) during a two‐month (June–August) field enclosure experiment. Twelve 20‐m‐long enclosed stream reaches were used in a 2 × 2 factorial design, with large wood either absent or added to mimic pre‐modern forestry conditions, and terrestrial invertebrate inputs either reduced or maintained at ambient levels. The addition of large wood had a positive effect on the growth of large trout but no effect on small trout, whereas terrestrial invertebrate input had no effect on the growth of either size class. Growth rates were highest in the treatment with ambient terrestrial invertebrate inputs and added wood, were lowest in the treatment with reduced terrestrial invertebrate inputs and no added wood and were intermediate in the other two treatments. Dietary analyses showed no difference in treatments with and without added wood, perhaps because instream wood influences growth by producing profitable stream positions for trout, rather than by acting as a source of prey. Terrestrial invertebrate inputs affected the diet, as trout in enclosures with reduced inputs had a lower proportion of terrestrial invertebrate biomass in the diet than trout in enclosures with ambient terrestrial inputs. Our results suggest that leaving woody debris in streams when harvesting forests may enhance trout growth and that this is probably due to the physical changes in depth and current velocity caused by the wood rather than to changes in dietary prey composition.
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