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.     

Direct and indirect effects of predatory young‐of‐year fishes in a dryland river food web

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From autochthonous to allochthonous resources: seasonal shifts in food use by stream‐dwelling YOY Arctic charr Salvelinus alpinus through the ice‐free season

Substantial seasonal changes in resource use associated with enhanced water‐column use were revealed in stream‐living YOY Arctic charr Salvelinus alpinus during the ice‐free season. In July, YOY individuals showed a diet dominated by aquatic invertebrates (mainly Chironomidae larvae), but despite the small size of the fish, the abundance of terrestrial insects in their diet increased markedly from July to September (from 1·9 to 62·8%). Similarly, the frequency of surface drifting foragers, i.e. individuals feeding on allochthonous resources, increased from July to September (from 20·6 to 80%); allochthonous resources thus constituting an important energy subsidy for YOY S. alpinus during the late sub‐Arctic summer.     

The trophic interactions of young Arctic grayling (Thymallus arcticus) in an Arctic tundra stream

1. Young (0+) Arctic grayling (Thymallus arcticus) have the potential to control the trophic structure of Arctic tundra streams through consumption, nutrient excretion and the modification of prey behaviour. The effect of young grayling on three trophic levels (algae, invertebrates and fish) was investigated by manipulating fish density and by fertilizing the river with phosphorus (P). 2. Nutrients, epilithic chlorophyll a, benthic invertebrates and fish biomass were measured within each fish density treatment (0, 4, and 40 fish m^–2) within the P-limited reference zone and the P-enriched fertilized zone of the Kuparuk River, Alaska. 3. Epilithic chlorophyll a increased with increased fish density in both reference and fertilized zones, while mayfly density decreased with increased fish density in the fertilized zone only. Final mean mass of young grayling in the 40 fish m^–2 cages was lower than mean mass in the 4 fish m^–2 cages. 4. Young grayling may produce a top-down cascading trophic effect in areas where nutrients are not limited. 5. River nutrient status and river discharge may modify the strength of top-down control by young grayling.     

Prey size,prey abundance,and temperature as correlates of growth in stream populations of cutthroat trout

Growth and maximum size of stream fishes can be highly variable across populations. For salmonid fishes in streams, individuals from populations confined to headwater streams often exhibit small size at maturity in comparison to populations with access to main-stem rivers. Differences in prey size, prey availability, and metabolic constraints based on temperature may explain patterns of maximum size and growth. In this study, cutthroat trout from headwater stream populations that were isolated above a waterfall were compared to individuals from populations in similar sized streams without a movement barrier and from large main-stem rivers. Cutthroat trout from smaller streams with or without a movement barrier were significantly smaller at a given age than fish from main-stem rivers, where individuals were able to achieve a much larger maximum size. Comparisons of invertebrate drift abundance and size in the three types of streams revealed that drift size did not differ between stream categories, but was highest per volume of water in large main-stem rivers. Across all stream types, prey abundance declined from summer to fall. Temperature declined over the course of the season in a similar manner across all stream types, but remained relatively high later in the season in main-stem river habitats. Prey availability and temperature conditions in main-stem rivers may provide more optimal growing conditions for fish as individuals increase in size and become constrained by prey availability and temperature conditions in small streams. Maintaining connectivity between small spawning and rearing tributary streams and main-stem river habitats may be critical in maintaining large-bodied populations of stream salmonids.     

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.     

Occupancy of young-of-year Arctic grayling (Thymallus arcticus) in Barrenland streams

Hydrobiologia - Arctic grayling (Thymallus arcticus) is an iconic fish species that is present across the remote subarctic Barrenlands, yet our lack of understanding of their distributional...     

Cost of predation avoidance in young-of-year lake trout (Salvelinus namaycush): growth differential in sub-optimal habitats

Selection of habitat to avoid predation may affect the diet of young-of-year (YOY) lake trout (Salvelinus namaycush). YOY lake trout may use inshore habitat to avoid predation; this habitat may be sub-optimal for growth. To test this, YOY lake trout were penned in nearshore and offshore pelagic areas of two arctic lakes. Toolik Lake had a lake trout population, the other lake, S6, did not. YOY lake trout in Toolik Lake lost weight, but those offshore lost less weight. The YOY lake trout in Lake S6 gained weight and those offshore gained more weight. The primary diet item of the YOY lake trout in both lakes during this experiment was the zooplankter Diaptomis probilofensis; it was also one of the most abundant species. However, its density inshore in Lake S6 was similar to inshore and offshore densities in Toolik Lake. The increased availability of alternative zooplankton prey in Lake S6 may account for the growth differential of YOY lake trout in Lake S6 relative to Toolik Lake. Bioenergetic modeling of YOY lake trout suggests that growth similar to that in the offshore of Lake S6 would be necessary for successful recruitment. If the reduced zooplankton availability in Toolik Lake leads to the reduced growth of YOY in the inshore and offshore pelagic areas, then these fish will be more susceptable to winter predation/starvation. For YOY lake trout to survive in Toolik Lake they most likely shift to feeding on benthic prey before the end of their first summer. Dept. of Chemical Engineering     

Ontogenetic diet shifts and resource partitioning among piscivorous fishes in the Venezuelan ilanos

Synopsis Resource utilization by nine abundant piscivores from a diverse tropical fish assemblage was examined over the course of a year. All nine species exhibited peak reproduction during the early wet season and a similar sequence of size-dependent shifts from a diet composed primarily of microcrustacea, to aquatic insects, and finally fishes. Three piranha species specialized on fish fins, particularly at subadult size classes (SL 30–80 mm). Gradual dessication of the floodplain during the transition season was associated with fish growth, increased fish density, and decreased aquatic primary productivity and availability of invertebrate prey. Based on 118 resource categories, average pairwise diet overlap was low during all three seasons: wet, transition, and dry. Of 72 species pairings, only one pair of fin-nipping piranhas exhibited high overlap simultaneously on three niche dimensions: food type, food size, and habitat. Adults of two species, a gymnotid knifefish and pimelodid catfish, were largely nocturnal. Patterns of habitat utilization indicate that piranhas may restrict diurnal use of the open-water region by other piscivores. Collective diet overlap of individual piscivore species with the other eight feeding guild members and collective overlaps with the entire fish community each revealed two basic seasonal trends. Four species that showed an early switch to piscivory also showed a high degree of diet separation with both the guild and community at large on a year-round basis. The five remaining species showed lowest collective diet overlaps during the transition season when availability of invertebrates was reduced and fish densities were maximal. Whereas predation may play a role in habitat separation, diffuse competition for food resources during the approximately four-month transition season probably is the principal factor yielding patterns of diet specialization.     

Cascading life-history interactions: alternative density-dependent pathways drive recruitment dynamics in a freshwater fish

Although density-dependent mechanisms in early life-history are important regulators of recruitment in many taxa, consequences of such mechanisms on other life-history stages are poorly understood. To examine interacting and cascading effects of mechanisms acting on different life-history stages, we stocked experimental ponds with fathead minnow (Pimephales promelas) at two different densities. We quantified growth and survival of the stocked fish, the eggs they produced, and the resulting offspring during their first season of life. Per-capita production and survival of eggs were inversely related to density of stocked fish; significant egg cannibalism by stocked minnows resulted in initial young-of-the-year (YOY) densities that were inversely related to adult densities. Subsequent growth and survival of YOY were then inversely related to these initial YOY densities, and survival of YOY was selective for larger fish. Because of these compensatory processes in the egg and YOY stages, treatments did not differ in YOY abundance and mean size at the end of the growing season. Because of differences in the intensity of size-selective mortality, however, variation in end-of season sizes of YOY was strongly (and inversely) related to densities of stocked fish. When mortality was severe in the egg stage (high densities of stocked fish), final YOY size distributions were more variable than when the dominant mortality was size-selective in the YOY stage (low stocked fish densities). These differences in size variation could have subsequent recruitment consequences, as overwinter survival is typically selective for YOY fish larger than a critical threshold size. Density-dependent effects on a given life stage are not independent, but will be influenced by earlier stages; alternative recruitment pathways can result when processes at earlier stages differ in magnitude or selectivity. Appreciation of these cascading effects should enhance our overall understanding of the dynamics of stage-structured populations.     

Diel feeding periodicity, daily ration and prey selection of a riverine population of juvenile chinook salmon, Oncorhynchus tshawytscha (Walbaum)

The diel feeding periodicity, daily ration and prey selection of juvenile chinook salmon, Oncorhynchus tshawytscha , were studied in relation to the available prey. Maximum dry weight of food intake occurred about dawn, when mayflies were the major prey, but the greatest number of freshly eaten prey occurred during the afternoon, when chironomids and terrestrial dipterans predominated. Feeding activity at night was low, with smaller mayflies comprising up to 50% of the prey. During the day the young salmon fed selectively on chironomids and the larger mayflies, while trichopterans and terrestrial taxa were under-represented in the diet. Food consumption over the 24-h period averaged 8.3% of the fish dry body weight. Prey abundance in the drift explained about 50% of the composition of the diet. Although the fish selected larger mayflies, size apparently was not a main criterion for selection because chironomids, although smaller than mayflies, were also frequently eaten. Previous dietary experience of the fish and the diel pattern of prey abundance appear to best explain the selective feeding of juvenile chinook salmon.     

Differences in composition of small,motile invertebrate assemblages from seagrass and unvegetated habitats in a southern Australian estuary

Assemblages of small, motile invertebrates (epifauna) from eelgrass(Zostera muelleri) and unvegetated habitats in a shallow, marine-dominatedestuary were compared at five sampling periods over one year. Assemblagesbased on abundance and biomass of 21 taxa from the two habitats groupedseparately in multivariate analyses (MDS ordination), and these groupingswere shown to be significant using an analysis of similarities (ANOSIM)randomisation routine. Secondarily to habitat differences, weak influencesof water temperature and distance to open water, but not of salinity, weredetected at some periods. Abundance and biomass of key taxa and all speciescombined were higha in eelgrass than in unvegetated habitat. Cumaceans wereexceptional in being collected predominantly from unvegetated habitat. Totalepifaunal production and crustacean production estimated using twovariables, (1) the biomass of individuals of each size class, and (2) watertemperature, were also higher in eelgrass than in unvegetated habitat. Thehigher abundance in eelgrass of taxa such as amphipods, harpacticoidcopepods and polychaetes that are major components of the diets of smallfish is consistent with a model explaining higher fish numbers in eelgrassin terms of prey availability.     

Feeding of tropical freshwater fishes: seasonality in resource availability and resource use

Summary Food resources in the environment and in the diets of small fish inhabiting two water bodies in a tropical savanna were studied during both wet and dry seasons. During the wet season (high water, abundant food) most fish species in both habitats fed predominantly on vegetation-dwelling invertebrates. Most fish species switched to alternative foods (algae and detritus) following the drastic decline in invertebrate food available towards the end of the dry season. In one habitat, this change in diet was accompanied by an increase in the volume of food intake. In the second habitat, only two larger species foraged intensively, while smaller species showed low food intake or almost ceased feeding. These differences may be explained by the high risk of predation for small fish in the second habitat. Dietary overlaps among fish species were high at the end of the dry season and moderate in the wet season. However, critical analysis of such factors as food abundance, the size and number of shared prey, and diet breadth showed that all significant overlaps were ecologically unimportant i.e. there was only weak competition for food.     

Effects of headwater impoundment and channelization on invertebrate drift

The construction of a flood control impoundment on Twitty's Creek added large numbers of organisms of limnetic origin to the stream ecosystem. However, the number of limnetic organisms per unit volume of water decreased rapidly as the distance downstream from the reservoir increased and, during most sampling periods, made up an insignificant portion of the total drift biomass at 7.2 km downstream. Factors favoring the extended downstream drift of limnetic organisms were high stream discharge and low water temperature.Several taxa of benthic organisms had much lower drift rates in the station immediately below the dam than at other stations and several taxa commonly taken at other stations were not captured immediately below the reservoir outfall. One possible explanation is that these organisms may have longer drift recruitment distances than the distance from the reservoir outfall to the sample location.A comparison of drift densities of organisms of benthic origin and benthic standing crop densities in channeled and unchanneled streams revealed that drift densities were higher in channeled streams than in unchanneled streams for most taxa of invertebrates. In addition, channeled streams appeared to have lower benthic standing crops than unchanneled streams for most taxa of invertebrates.In stream sections impacted by either channelization or the Twitty Lake outfall, the energy dynamics of the stream ecosystems were altered by increased density of drifting invertebrates. From the standpoint of increasing food availability to the fish fauna of the stream, these changes would appear to benefit drift feeding species and negatively impact bottom feeding species.     

Habitat effects on invertebrate drift in a small trout stream: implications for prey availability to drift-feeding fish

In this study, we focused on the drivers of micro- and mesohabitat variation of drift in a small trout stream with the goal of understanding the factors that influence the abundance of prey for drift-feeding fish. We hypothesized that there would be a positive relationship between velocity and drift abundance (biomass concentration, mg/m³) across multiple spatial scales, and compared seasonal variation in abundance of drifting terrestrial and aquatic invertebrates in habitats that represent the fundamental constituents of stream channels (pools, glides, runs, and riffles). We also examined how drift abundance varied spatially within the water column. We found no relationship between drift concentration and velocity at the microhabitat scale within individual pools or riffles, suggesting that turbulence and short distances between high- and low-velocity microhabitats minimize changes in drift concentration through settlement in slower velocity microhabitats. There were also minimal differences in summer low-flow drift abundance at the mesohabitat scale, although drift concentration was highest in riffle habitats. Similarly, there was no differentiation of drifting invertebrate community structure among summer samples collected from pools, glides, runs, and riffles. Drift concentration was significantly higher in winter than in summer, and variation in drift within individual mesohabitat types (e.g., pools or riffles) was lower during winter high flows. As expected, summer surface samples also had a significantly higher proportion of terrestrial invertebrates and higher overall biomass than samples collected from within the water column. Our results suggest that turbulence and the short length of different habitat types in small streams tend to homogenize drift concentration, and that spatial variation in drift concentrations may be affected as much by fish predation as by entrainment rates from the benthos. Handling editor: Robert Bailey     

A trait‐based approach reveals the feeding selectivity of a small endangered Mediterranean fish

Functional traits are growing in popularity in modern ecology, but feeding studies remain primarily rooted in a taxonomic‐based perspective. However, consumers do not have any reason to select their prey using a taxonomic criterion, and prey assemblages are variable in space and time, which makes taxon‐based studies assemblage‐specific. To illustrate the benefits of the trait‐based approach to assessing food choice, we studied the feeding ecology of the endangered freshwater fish Barbus meridionalis. We hypothesized that B. meridionalis is a selective predator which food choice depends on several prey morphological and behavioral traits, and thus, its top‐down pressure may lead to changes in the functional composition of in‐stream macroinvertebrate communities. Feeding selectivity was inferred by comparing taxonomic and functional composition (13 traits) between ingested and free‐living potential prey using the Jacob's electivity index. Our results showed that the fish diet was influenced by 10 of the 13 traits tested. Barbus meridionalis preferred prey with a potential size of 5–10 mm, with a medium–high drift tendency, and that drift during daylight. Potential prey with no body flexibility, conical shape, concealment traits (presence of nets and/or cases, or patterned coloration), and high aggregation tendency had a low predation risk. Similarly, surface swimmers and interstitial taxa were low vulnerable to predation. Feeding selectivity altered the functional composition of the macroinvertebrate communities. Fish absence favored taxa with weak aggregation tendency, weak flexibility, and a relatively large size (10–20 mm of potential size). Besides, predatory invertebrates may increase in fish absence. In conclusion, our study shows that the incorporation of the trait‐based approach in diet studies is a promising avenue to improve our mechanistic understanding of predator–prey interactions and to help predict the ecological outcomes of predator invasions and extinctions.     

Fish predation affects the structure of a benthic community

1. We conducted an experimental study of predation by benthivorous fish on a natural community of stream invertebrates using a reach‐scale approach. Over a 2‐year period (experimental phase), the benthic invertebrate community of a stretch containing two species of benthivorous fish was compared with a fishless stretch. Thereafter, all fish were removed and benthic community structure was analysed again to account for natural differences between the two stretches (reference phase). 2. Benthivorous fish at the moderate densities investigated did not affect total benthic biomass or density, but did alter species composition. In addition, the fish effect differed between pool and riffle habitats, with larger effects in the pools indicating a habitat‐specific predation effect. In the reference phase, when all fish were removed from the stream, the difference between the two stretches was reduced. 3. The benthivorous fish reduced the densities of four taxa (Pisidium sp., Dugesia gonocephala, Gammarus pulex, Limoniidae), representing 29% of total biomass. It is possible that density reductions of other species were masked by prey migration despite the relatively large spatial scale. Indeed, higher drift activity in the upstream fishless stretch could have increased the density of Baetis rhodani in the fish stretch, as indicated by the results of a drift model. 4. Our results provide insights into stream food web ecology because fish predation showed effects even in a natural system where habitat complexity was high, environmental factors were highly variable and many predator and prey species interacted and because benthivorous fish were the focus, whereas the majority of previous predation experiments in streams have used drift‐feeding trout.     

Invertebrate responses to short-term water abstraction in small New Zealand streams

1. Small permanent streams are coming under increasing pressure for water abstraction. Although these abstractions might only be required on a short‐term basis (e.g. summer time irrigation), the highest demand for water often coincides with seasonal low flows. 2. We constructed weirs and diversions that reduced discharge in three small streams (<4 m width) to test the hypotheses that short‐term water abstractions would decrease habitat availability and suitability for invertebrates, resulting in increased invertebrate drift, reduced taxonomic richness and decreased benthic invertebrate densities. 3. We sampled benthic invertebrates, invertebrate drift and periphyton at control (upstream) and impact (downstream) sites on each stream before and during 1 month of discharge reduction. 4. Discharge decreased by an average of 89–98% at impact sites and wetted width decreased by 24–30%. Water depth decreased by 28–64% while velocity decreased by 50–62%. Water conductivity, temperature and dissolved oxygen showed varying responses to flow reduction among the three streams, whereas algal biomass and pH were unaffected in all streams. 5. The densities of invertebrate taxa tended to increase in the impact reaches of these streams, even though invertebrate drift increased at impact sites in the first few days following discharge reduction. There were a higher proportion of mayflies, stoneflies and caddisflies at the impact site on one stream after flow reduction. There were no changes to the number of taxa or species evenness at impact sites. 6. Our results suggest that for these small streams, the response of invertebrates to short‐term discharge reduction was to accumulate in the decreased available area, increasing local invertebrate density.     

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.     

Summer diet of beluga whales inferred by fatty acid analysis of the eastern Beaufort Sea food web

Beluga whales (Delphinapterus leucas) are the most abundant odontocetes in Arctic waters and are thus thought to influence food web structure and function. The diet of the Beaufort Sea beluga population is not well known, partly due to the inherent difficulty of observing feeding behaviour in Arctic marine cetaceans. To determine which prey items are critical to the Beaufort Sea beluga diet we first examine and describe the Mackenzie Delta and Beaufort Sea food web using fatty acid analyses. Fatty acid profiles effectively partitioned prey items into groups associated with their habitat and feeding ecology. Next, the relative contribution of various prey items to beluga diet was investigated using fatty acids. Finally, beluga diet variability was examined as a function of body size, a known correlate of habitat use. Beluga appeared to feed predominantly on Arctic cod (Boreogadus saida) collected from near shore and offshore regions. Size related dietary differences suggested larger sized beluga preferred offshore Arctic cod given the shared high levels of long chain monounsaturates, whereas smaller sized beluga appeared to feed on prey in their near shore habitats that included near shore Arctic cod. The presence of Arctic cod groups in shallow near shore and deep offshore habitats may facilitate the behavioural segregation of beluga habitat use as it relates to their size and resource requirements. Given Arctic cod are a sea ice associated fish combined with the accelerated sea ice loss in this region, beluga whales may need to adapt to new dietary regimes.