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.     

Feeding ecology of a vertebrate assemblage inhabiting a stream of NW Spain (Riobo; Ulla basin)

The food resource use of a stream in NW Spain by fish (Salmo trutta L. and Anguilla anguilla L.), birds (Cinclus cinclus L. and Motacilla cinerea L.) and mammals (Galemys pyrenaicus G. and Neomys anomalus C.) was studied. Data on seasonal diets and stream benthos prey were used to determine prey selection patterns.Caddisfly larvae are the main resource for Cinclus and Galemys, but these predators also consumed other benthic prey. Salmo fed on a wide range of benthic invertebrates, emergent pupae and terrestrial prey, whereas Anguilla consumed primarily benthic invertebrates, especially Lumbricids. Neomys fed mainly on terrestrial prey (Gasteropods and Lumbricids), but also consumed aquatic prey. Motacilla captured aquatic insects both in larval and aerial stages, as well as terrestrial prey.Both prey availability and selection led to seasonal differences in the use of food resources. All species showed a marked prey selection of aquatic taxa. Prey size plays an important role in this selection, most species consuming the largest of available prey sizes. In spite of the fact that all species feed upon freshwater invertebrates, substantial resource partitioning was observed in all seasons. This partitioning may be attributable to morpholological and physiological differences. Nevertheless, Anguilla and Galemys, two quite different animals, did feed on the same prey much of the time.     

Resource seasonality and fish diets in an Illinois stream

Synopsis The purpose of this study was to evaluate the intensity of competition for food among 9 species of stream fishes that primarily eat aquatic invertebrates. The taxonomic and size composition, and numerical abundance of aquatic invertebrates were monitored for one year using drift and benthic samples. Diet data were obtained from stomachs of fishes captured at the same time and place that invertebrate sampling was done. Diet characteristics examined included taxonomic and size composition, number of prey per fish, and diet breadth. Drifting invertebrates were more abundant early in the year (March–June) than later (July–January). The summer-early fall scarcity of invertebrates was especially notable among those>3.6 mm long, which comprised the bulk of prey found in fish stomachs. Average prey size eaten by a fish species was positively correlated with fish mouth size, but interspecific overlap in prey size was extensive. Cyprinids as a group (5 species) ate proportionally fewer small (< 3.6 mm long) prey from July to January than did the centrarchids and stonecat. Taxonomic compositions of available invertebrates and fish diets varied markedly among sampling dates, but the use of prey taxa by fishes was not correlated with the availability of those taxa. Use of aquatic prey taxa was generally similar among fish species, but cyprinids as a group ate proportionally more terrestrial prey from July to January than did the centrarchids and stonecat. Diet breadths for all species increased as food levels declined, indicating that these fishes experienced resource depression. Food scarcity was evidently more severe for cyprinids since their stomachs contained few prey through the summer and fall relative to the centrarchids and stonecat. Though the fish species studied probably compete for food in the summer and fall, this competition did not account for the community structure observed.     

Invading rainbow trout usurp a terrestrial prey subsidy from native charr and reduce their growth and abundance

Movements of prey organisms across ecosystem boundaries often subsidize consumer populations in adjacent habitats. Human disturbances such as habitat degradation or non-native species invasions may alter the characteristics or fate of these prey subsidies, but few studies have measured the direct effects of this disruption on the growth and local abundance of predators in recipient habitats. Here we present evidence, obtained from a combined experimental and comparative study in northern Japan, that an invading stream fish usurped the flux of allochthonous prey to a native fish, consequently altering the diet and reducing the growth and abundance of the native species. A large-scale field experiment showed that excluding terrestrial invertebrates that fell into the stream with a mesh greenhouse reduced terrestrial prey in diets of native Dolly Varden charr (Salvelinus malma) by 46–70%, and reduced their growth by 25% over six weeks. However, when nonnative rainbow trout (Oncorhynchus mykiss) were introduced, they monopolized these prey and caused an even greater reduction of terrestrial prey in charr diets of 82–93%, and reduced charr growth by 31% over the same period. Adding both greenhouse and rainbow trout treatments together produced similar results to adding either alone. Results from a comparative field study of six other stream sites in the region corroborated the experimental findings, showing that at invaded sites rainbow trout usurped the terrestrial prey subsidy, causing a more than 75% decrease in the biomass of terrestrial invertebrates in Dolly Varden diets and forcing them to shift their foraging to insects on the stream bottom. Moreover, at sites with even low densities of rainbow trout, biomass of Dolly Varden was more than 75% lower than at sites without rainbow trout. Disruption of resource fluxes between habitats may be a common, but unidentified, consequence of invasions, and an additional mechanism contributing to the loss of native species Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Diet and condition of three fish species (Characidae) of the Andean foothills in relation to deforestation

Synopsis I studied effects of deforestation on diet and condition of three characid species; Astyanax zonatus, Knodus gamma, and Prionobrama filigera in small streams in the Ecuadorian Amazon. The best condition of K. gamma was found at sites with low canopy cover, while the best condition of P. filigera was found at sites with high canopy cover. Deforestation changed population structure of A. zonatus by decreasing mean size of individuals, but this was not associated with lower fish condition. A. zonatus was omnivorous, and fed equally on invertebrate and plant food. A. zonatus increased feeding on aquatic food items (filamentous algae and fish scales) in the deforested sites (43%) compared to the forested sites (16%). K. gamma was an omnivorous species with preference for invertebrates. K. gamma was mainly feeding on terrestrial food sources in the forested sites, but switched to equal feeding between terrestrial and aquatic food sources in the deforested sites. P. filigera was an terrestrial insectivore feeder, and no differences in the overall food categories was found between stream groups. In contrast, there were no pronounced difference in the composition of invertebrate taxa in the stomachs of K. gamma between stream groups. However, less empty stomachs were observed among K. gamma in the deforested sites. The stomach contents of P. filigera showed great changes in composition of the predominating prey taxa between stream groups and less empty stomachs were observed among P. filigera in the forested sites. Food availability and utilisation seemed to be of main importance for the condition of these fish. Deforestation seems to favour species with high availability to utilise aquatic food items. In contrast, deforestation seems to have negative effects on species that are specialised on terrestrial invertebrates, and have low capacity to switch between aquatic and terrestrial food items.     

Selective Feeding of age-0 Arctic Grayling in Lake-Outlet Streams of the Northwest Territories, Canada

A paucity of information exists on the diet of Arctic grayling, Thymallus arcticus, particularly for young-of-the-year (YOY). We examined the diet of YOY Arctic grayling in relation to food availability, in the Barrenlands region of the Northwest Territories, Canada, where lake-outlet streams serve as nursery habitat for these fish. Given the small size of YOY grayling and the abundance of lake-derived microcrustacea in the drift of these lake-outlet streams, we anticipated that these prey would make up a major component of the YOY's diet. Food selectivity by YOY grayling, however, was strongly sized-biased; although microcrustacea dominated the drift, YOY primarily consumed larger taxa, especially Chironomidae and Simuliidae. Even among these taxa, grayling tended to select the larger individuals. As they grew, YOY grayling took larger numbers of both large and small prey, particularly the larger invertebrates, although prey size range did not change after mid-July. Selection of pupae and avoidance of Ephemeroptera suggest that prey characteristics other than size also contribute to selectivity by YOY grayling. The relatively limited consumption of terrestrial invertebrates and other large prey may reflect the small sizes of fish in this arctic study, as well as differences in prey availability. Despite the abundance of lake-derived prey, instream production of invertebrates should largely determine the productive capacity of Barrenlands streams as fish habitat.     

Food habit divergence between white-spotted charr and masu salmon in Japanese mountain streams: circumstantial evidence for competition

 The diet and foraging microhabitat of white-spotted charr, Salvelinus leucomaenis, were compared between mountain stream reaches where it occurs with (sympatric) and without (allopatric) masu salmon, Oncorhynchus masou masou, a potential competitor, to examine the evidence for interspecific competition between these fish, which commonly co-occur in Japanese mountain streams. In three streams examined, the similarity between the diets of allopatric charr and salmon was much greater than that between the diets of sympatric charr and salmon. Both allopatric charr and sympatric salmon intensively utilized terrestrial invertebrates among stream drifts (52%–65% of the diet), whereas this prey category made up only an intermediate portion of sympatric charr diets (11%–29%). Examination of available prey composition in stream drifts showed that the consumption of terrestrial invertebrates by allopatric charr and sympatric salmon was approximately twice as much as that by sympatric charr. The presence of salmon, a potential competitor, may alter the diet of charr in the sympatric reaches. Charr holding focal points closer to the streambed were considered less efficient than sympatric salmon in their utilization of terrestrial invertebrates drifting primarily on the stream surface, although the foraging microhabitat of the charr was not influenced by the salmon. The mechanisms responsible for the dietary divergence between sympatric charr and salmon are probably the consequence of scramble competition over terrestrial invertebrates drifting on the stream surface. Received: January 21, 2002 / Accepted: November 19, 2002 Acknowledgments We thank Y. Tokuda, T. Takasu, Y. Kaneda, H. Jyoya, and H. Aoe for their assistance. This work was partly supported by funding through the Takara Harmonist Fund by the Takara Syuzo Co. Ltd. and the Japan Ministry of Education, Science, Sport and Culture (grants 09NP1501 and 11440224). Correspondence to:H. Miyasaka     

Effects of stream restoration and wastewater treatment plant effluent on fish communities in urban streams

1. Fish community characteristics, resource availability and resource use were assessed in three headwater urban streams in Piedmont North Carolina, U.S.A. Three site types were examined on each stream; two urban (restored and unrestored) and a forested site downstream of urbanisation, which was impacted by effluent from a wastewater treatment plant (WWTP). Stream basal resources, aquatic macroinvertebrates, terrestrial macroinvertebrates and fish were collected at each site. 2. The WWTPs affected isotope signatures in the biota. Basal resource, aquatic macroinvertebrate and fish δ¹⁵N showed significant enrichments in the downstream sites, although δ¹³C signatures were not greatly influenced by the WWTP. Fish were clearly deriving a significant part of their nutrition from sewage effluent‐derived sources. There was a trend towards lower richness and abundance of fish at sewage‐influenced sites compared with urban restored sites, although the difference was not significant. 3. Restored stream sites had significantly higher fish richness and a trend towards greater abundance compared with unrestored sites. Although significant differences did not exist between urban restored and unrestored areas for aquatic and terrestrial macroinvertebrate abundances and biotic indices of stream health, there appeared to be a trend towards improvements in restored sites for these parameters. Additional surveys of these sites on a regular basis, along with maintenance of restored features are vital to understanding and maximising restoration effectiveness. 4. A pattern of enriched δ¹³C in fish in restored and unrestored streams in conjunction with enriched δ¹³C of terrestrial invertebrates at these sites suggests that these terrestrial subsidies are important to the fish, a conclusion also supported by isotope cross plots. Furthermore, enriched δ¹³C observed for terrestrial invertebrates is consistent with some utilisation of the invasive C4 plants that occur in the urban riparian areas.     

Diet partitioning in sympatric Atlantic salmon and brown trout in streams with contrasting riparian vegetation

Prey intake by Atlantic salmon Salmo salar and brown trout Salmo trutta was measured across different riparian vegetation types: grassland, open canopy deciduous and closed canopy deciduous, in upland streams in County Mayo, Western Ireland. Fishes were collected by electrofishing while invertebrates were sampled from the benthos using a Surber sampler and drifting invertebrates collected in drift traps. Aquatic invertebrates dominated prey numbers in the diets of 0+ year Atlantic salmon and brown trout and 1+ year Atlantic salmon, whereas terrestrial invertebrates were of greater importance for diets of 1+ and 2+ year brown trout. Terrestrial prey biomass was generally greater than aquatic prey for 1+ and 2+ year brown trout across seasons and riparian types. Prey intake was greatest in spring and summer and least in autumn apart from 2+ year brown trout that sustained feeding into autumn. Total prey numbers captured tended to be greater for all age classes in streams with deciduous riparian canopy. Atlantic salmon consumed more aquatic prey and brown trout more terrestrial prey with an ontogenetic increase in prey species richness and diversity. Atlantic salmon and brown trout diets were most similar in summer. Terrestrial invertebrates provided an important energy subsidy particularly for brown trout. In grassland streams, each fish age class was strongly associated with aquatic, mainly benthic invertebrates. In streams with deciduous riparian canopy cover, diet composition partitioned between conspecifics with older brown trout associated with surface drifting terrestrial invertebrates and older Atlantic salmon associated with aquatic invertebrates with a high drift propensity in the water column and 0+ year fish feeding on benthic aquatic invertebrates. Deciduous riparian canopy cover may therefore facilitate vertical partitioning of feeding position within the water column between sympatric Atlantic salmon and brown trout. Implications for riparian management are discussed.     

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.     

Seasonal variation in the trophic structure of a spatial prey subsidy linking aquatic and terrestrial food webs: adult aquatic insects

Research over the past decade has established spatial resource subsidies as important determinants of food web dynamics. However, most empirical studies have considered the role of subsidies only in terms of magnitude, ignoring an important property of subsidies that may affect their impact in recipient food webs: the trophic structure of the subsidy relative to in situ resources. This may be especially important when subsidies are composed of organisms, as opposed to nutrient subsidies, because the trophic position of subsidy organisms may differ from in situ prey. I explored the relative magnitude and trophic structure of a cross-habitat prey subsidy, adult aquatic insects, in terrestrial habitats along three streams in the south–central United States. Overall, adult aquatic insects contributed more than one-third of potential insect prey abundance and biomass to the terrestrial habitat. This contribution peaked along a permanent spring stream, reaching as high as 94% of abundance and 86% of biomass in winter. Trophic structure of adult aquatic and terrestrial insects differed. Nearly all adult aquatic insects were non-consumers as adults, whereas all but one taxon of terrestrial insects were consumers. Such a difference created a strong relationship between the relative contribution of the prey subsidy and the trophic structure of the prey assemblage: as the proportion of adult aquatic insects increased, the proportion of consumers in the prey assemblage declined. Specific effects varied seasonally and with distance from the stream as the taxonomic composition of the subsidy changed, but general patterns were consistent. These findings show that adult aquatic insect subsidies to riparian food webs not only elevate prey availability, but also alter the trophic structure of the entire winged insect prey assemblage.     

Habitat overlap between predatory benthic fish and their invertebrate prey in streams: the relative influence of spatial and temporal factors on predation risk

1. The spatial heterogeneity of ecosystems as well as temporal activity patterns of organisms can have far‐reaching effects on predator–prey relationships. We hypothesised that spatiotemporal constraints in mesohabitat use by benthic fish predators would reduce habitat overlap with benthic invertebrates and lead to mesohabitat‐specific predation risks. 2. We analysed the spatiotemporal activity patterns of two small‐bodied benthivorous fishes, gudgeon (Gobio gobio) and stone loach (Barbatula barbatula), and of benthic invertebrates in a small temperate stream during three 24‐h field experiments. By applying a novel method of field video observation, we monitored the spatiotemporal foraging behaviour of the fish in their natural environment. A parallel analysis of invertebrate mesohabitat use by means of small area Hess sampling allowed a direct estimation of habitat overlap at a pool–riffle scale. 3. Gudgeon showed a dominant spatial activity pattern preferring pools at all times of day, whereas stone loach used both mesohabitats but with a distinct temporal (nocturnal) activity pattern. The patterns of residence were not identical with those of active foraging. Invertebrate community composition differed significantly between mesohabitats but not between times of day. More than half of the total dissimilarity between pools and riffles was accounted for by six invertebrate taxa. Five of these were subject to higher fish predation in pools than in riffles. The total prey consumption of the two fish species together in pools was about three times as high as in riffles. Trophic niche breadth of stone loach and thus its predation range was broader than that of gudgeon. 4. These results indicate that the potential predation risk for stream invertebrates depends on the combination of spatial and temporal patterns of both predator and prey. Given the distinct differences in predation risk found between pools and riffles, we conclude that spatial heterogeneity at the mesohabitat scale can influence mechanisms and consequences of selective predation. We also suggest that the analysis of spatiotemporal predator–prey relationships should not be based on the premise that the main residence habitat and active foraging habitat of a predator are identical.     

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.     

A Bioenergetic Analysis of Factors Limiting Brown Trout Growth in an Ozark Tailwater River

We examined prey utilization and energy consumption by brown trout, Salmo trutta, in a cold tailwater (Little Red River, Arkansas, USA; LRR) having low biodiversity and low availability of fish as prey. Stomach content analysis and age estimation were performed on thirty brown trout (10 each of three size classes for a total of 710 trout) collected monthly from an upstream and downstream site over a 1-year period. Diet diversity was low at both sites, as 80% and 70% of all prey consumed by upstream and downstream brown trout, respectively, were isopods. Piscivory (<0.5% of individuals sampled) and consumption of terrestrial invertebrates were rare. There was no relation between diet diversity and trout age, and a very small ontogenetic shift in brown trout diet. Second, we investigated brown trout growth rates relative to prey consumption and temperature. Temperatures and availability of prey were less than required for maximal trout growth. However, prey availability limited trout growth directly, but sub-optimal temperatures probably buffered the effect of this reduced energy consumption by reducing metabolic energy expenditures. Brown trout growth was 54.8–57.0% of the maximum predicted by a bioenergetics model. Instantaneous growth rates for age 1 and adult brown trout were slightly higher for those downstream (0.195) versus those upstream (0.152). Although isopods are abundant within this tailwater to serve as a forage base, the displacement of native fish fauna and subsequent lack of establishment of cold-tolerant forage fish species due to the thermal regime of hypolimnetic release from Greers Ferry Reservoir probably serves as a major barrier to brown trout growth.     

Ecosystem-level evidence for top-down and bottom-up control of production in a grassland stream system

Ecosystem-wide effects of introduced brown trout (Salmo trutta L.) and native river galaxias (Galaxiaseldoni McDowall) were studied by analysing ecosystem production budgets for two adjacent tributaries of a grassland stream-system in the South Island of New Zealand. One tributary was inhabited by brown trout, the other by river galaxias. No other fish species were present in either stream. The budget for the river galaxias stream indicated little top-down control of invertebrates by fish predation (river galaxias consumed ∼18% of available prey production). A large proportion of annual net primary production was required to support production by invertebrates (invertebrates consumed an average of ∼75% of available primary production), and mean surplus primary production (i.e. not consumed) was not significantly different from zero. Primary and secondary production were presumably mutually limiting in this system (i.e. controlled by simultaneous top-down and bottom-up mechanisms). In contrast, the budget for the brown trout stream indicated extreme top-down control of invertebrate populations by fish predation; essentially all invertebrate production (∼100%) was required to support trout production. Invertebrate production required only a minor portion of annual net primary production (∼21%) and primary production was presumably controlled by mechanisms other than grazing (e.g. sloughing, nutrient limitation). Predatory invertebrates had little quantitative effect on prey populations in either stream. Recent experimental studies of invertebrate behaviour, fish behaviour, and food-web structure in New Zealand streams with physically stable channels indicate that a trophic cascade should be observed in streams inhabited by brown trout, in contrast to those inhabited by native fish. The results reported here provide ecosystem-level evidence supporting this prediction. Received: 10 March 1997 / Accepted: 12 December 1997     

Forest succession and prey availability influence the strength and scale of terrestrial‐aquatic linkages in a headwater salamander system

1. Trophic linkages between terrestrial and aquatic ecosystems are common and sensitive to disruption. However, there is little information on what causes variation in the strength and spatial scale of these linkages. 2. In the highly aquatic adults of the headwater salamander Gyrinophilus porphyriticus (family Plethodontidae), use of terrestrial prey decreases along a gradient from early‐ to late‐successional riparian forests. To understand the cause of this relationship, we tested the predictions that (i) terrestrial prey abundance is lower in late‐successional forests, and (ii) G. porphyriticus adults cannot move as far from the stream to forage in late‐successional forests, thus limiting access to terrestrial prey. 3. We established 100‐m long study reaches on six headwater streams in the Hubbard Brook Experimental Forest, New Hampshire. Three reaches were in early‐successional forests and three were in late‐successional forests. We conducted pitfall trapping for invertebrate prey in June and July of 2005, with three traps at 0, 2, 5 and 10 m from the stream at each reach. In June, July and August of 2004 and 2005, nighttime salamander surveys were conducted at each reach along ten, 10‐m long by 2.5‐m wide transects perpendicular to the stream. 4. Abundance of terrestrial prey was consistently lower in late‐successional forests, suggesting that consumption of terrestrial prey by G. porphyriticus is affected by prey abundance. Contrary to our prediction, G. porphyriticus adults moved farther from the stream in late‐successional forests, suggesting that habitat conditions in late‐successional forests do not limit movement away from the stream, and that lower abundances of terrestrial prey in these forests may cause salamanders to move farther from streams. 5. Our results provide novel insight on the extent of terrestrial habitat use by G. porphyriticus. More broadly, these results indicate that major habitat gradients, such as forest succession, can affect the strength and scale of terrestrial‐aquatic linkages. Application of this insight to the design of vegetation buffers along headwater streams would have widespread benefits to freshwater ecosystems.     

Asymmetric competition between distant taxa: poecilid fishes and water striders

Summary Competition for water surface prey between fish (Priapichtus annectens: Poeciliidae) and water striders (Potamobates unidentatus: Gerridae) was studied in the laboratory and in pools in a small tropical stream. Laboratory experiments showed that fish depressed activity and foraging success of water striders. Large fish (4–5 cm) had a greater effect than small fish (2–3 cm). The field experiment showed that competition was highly asymmetric. Presence of fish decreased water strider foraging success while the reverse interaction was insignificant. It is suggested that the higher individual foraging success of the fish, harassment of water striders by fish and the use of an exclusive resource, benthic invertebrates, by the fish, contribute to this pattern. Habitat use differed between the two species. Fish used the deeper parts of stream pools and water striders used the shallower parts of the pools. Asymmetric interference and exploitation competition may force water striders to use shallow edge habitats.     

Trophic interception: how a boundary-foraging organism influences cross-ecosystem fluxes

The rate at which subsidies move between habitats is a function not only of those factors that facilitate such transfers, but also of factors that limit or restrict the movement of subsidies. The interruption or redirection of subsidies by organisms foraging at the boundary between habitats, or trophic interception, has the potential to substantially restructure the food webs of recipient habitats. In this study we describe how a waterstrider, Aquarius remigis , limits the transfer of a subsidy across the land-water boundary. Prey interception varied with the type of stream habitat; on average, waterstriders in isolated pools intercepted 71.8% of experimental prey inputs of all sizes compared with 21.5% in connected pools and 0.8% in riffles. Across all stream habitat types, waterstriders consumed 43.2% of the experimental inputs of the smallest prey used, the prey size class most similar to natural inputs in our study area. We estimate that foraging waterstriders may intercept 0.3–1.2 kg of terrestrial prey subsidy for every 100 m of stream channel during three summer months, resources that could otherwise support 13–58 young-of-the-year salmonids. In controlled trials, waterstriders significantly altered the amount of terrestrial prey directly consumed by fish, while fish also altered waterstrider interception of those prey. Interestingly, when waterstriders and fish were present together, more terrestrial prey were lost to the streambed than when either or both of these consumers were absent, making this resource available to benthic detritivores, and facilitating the direct incorporation of terrestrial nutrients into stream detrital webs. Overall, we demonstrate that organisms that forage at habitat boundaries can control the quantity and quality of subsidies arriving in recipient habitats, potentially altering food web structures within those habitats.     

The piscivore feeding guild of fishes in small freshwater ecosystems

Synopsis Analysis of the piscivore guild in fish species-rich lake and stream systems in eastern Ontario showed the co-occurrence of three types: (1) specialists that became piscivorous at the age of a few weeks (Esox, Micropterus); (2) ‘secondary’ piscivores that are fish-eaters only later in life (Perca, Ambloplites); and (3) species in which fish consumption is limited to taking some larvae (Lepomis macrochirus). In the first group the basic series of dietary shifts that characterize many long-lived fish (i.e. zooplankton followed by small invertebrates then large invertebrates and finally fish); is greatly accelerated. Prey size increases with growth. Overall prey selection was on the basis of body size and abundance. Most piscivores took a range of fish prey. There was little evidence of specialization at the species level. Esox and Micropterus spawn some weeks ahead of their major prey species. This is seen as adaptive. Their young harvest the larvae of the latter. The ensuing predator/prey association with growth is highly advantageous to the piscivore as prey of optimum body size are thus continually available.     

Individual specialization and trophic adaptability of northern pike (Esox lucius): an isotope and dietary analysis

Northern pike (Esox lucius) are often considered to be specialist piscivores, but under some circumstances will continue to eat invertebrates as adults. To examine effects of fish assemblage composition on the trophic ecology of pike, we combined stable isotope analysis (SIA) of carbon and nitrogen and stomach content analysis (SCA) on pike from five lakes in northern Alberta, three of which contain only pike (“pike-only”) and two that also contain yellow perch (Perca flavescens) or white sucker (Catostomus commersoni) (“pike-other”). Fish were more important as prey and empty stomachs, which often characterize piscivores, were significantly more frequent in pike-other than in pike-only lakes. However, even though invertebrates were more important for pike in pike-only lakes, SIA and SCA indicated that invertebrates were also an important component of pike diets in pike-other lakes. SIA and SCA also revealed considerable intrapopulation variation in trophic ecology, with individuals in some populations differing by as much as two trophic levels. Comparisons of stomach contents and isotope signatures of the same fish suggested that within these variable populations, specialization on invertebrates or fish was a long-term trait of some individuals. SIA indicated that trophic position increased and diets shifted to a greater importance of littoral prey as pike grew in pike-only lakes, but not in lakes with other fish present. Trophic adaptability in northern pike is expressed at both the population level, where the trophic ecology is sensitive to differences in prey regimes, and at the organismal level, in the form of intrapopulation variation and individual specialization. Received: 1 July 1998 / Accepted: 3 February 1999