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

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

Impact of whitefish on an enclosure ecosystem in a shallow eutrophic lake: selective feeding of fish and predation effects on the zooplankton communities

Bag-type enclosures (75 m³) with bottom sheets and tube-type enclosures (105 m³) open to the bottom sediment were stocked with exotic whitefish (Coregonus lavaretus maraena) to study their predation effects on the plankton community. The fish fed mainly on adult chironomids during the period of their emergence (earlier part of the experimental period). Thereafter, the food preference was shifted to larvae of chironomids and crustacean zooplankters. The predation effects on the plankton community were not evident in the bag-type enclosures where zooplankton densities were consistently low. The fish reduced the crustacean populations composed ofBosmina fatalis, B. longirostris andCyclops vicinus in the tube-type enclosures where the prey density was high (above ca. 50 individuals 1⁻¹). The results suggested that the intensity of predation depended on the prey density. Rotifers increased in the fish enclosure, probably becauseCoregonus reduced the predation pressure byCyclops vicinus on rotifers and allowed the latter to increase. In the fish enclosures, no marked changes in species composition were observed. Zooplankton predated by the fish seemed to be distributed near the walls of the enclosures. Problems of enclosure experiments for examining the effects of fish predation on pelagic zooplankton communities are discussed.     

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.     

Influence of the predatory backswimmer, Notonecta maculata, on invertebrate community structure

1. The impact of the predator, Notonecta maculata , on invertebrate community structure was investigated by a combination of a natural pool survey, an outdoor artificial pool experiment, and a laboratory prey preference experiment.
2. In a survey of forty natural pools in the Negev Desert, Israel, pelagic/neustonic species richness and overall species richness, but not benthic species richness, were negatively associated with N. maculata density.
3. In the artificial pool experiment, in which the presence or absence of Notonecta was manipulated, Notonecta caused a reduction in the richness of species colonizing the pools. Notonecta strongly reduced densities of the pelagic dipterans, Culiseta longiareolata (often 100% reduction of later-instar larvae and pupae) and ephydrids. Fewer Culiseta egg rafts were found in Notonecta pools, indicating that part of the observed reduction of Culiseta is not due to consumption of Culiseta larvae by Notonecta, but to habitat selection by ovipositing Culiseta females in response to risk of predation by Notonecta . Notonecta did not affect densities of (benthic) chironomids or the small cladoceran, Ceriodaphnia cf. quadrangula . Laboratory assessments of selectivity by Notonecta for the various prey matched patterns found in the field experiment.
4. This work provides strong evidence that N. maculata is an important organizer of community structure: it strongly reduces or even eliminates larger pelagic or neustonic species, but does not affect densities of small or benthic species.     

Predatory fish impact on competition between stream insect grazers: a consideration of behaviorally- and density-mediated effects on an apparent coexistence pattern

A manipulative field experiment and theoretical analyses of a simple competition model were used to show how exploitative competition between a caddisfly (Glossosoma sp.) and three mayfly grazers (Ameletus sp., Baetisthermicus and Cinygmula sp.) was mediated by a predatory fish, freshwater sculpin (Cottus nozawae). The field experiment followed a two‐factorial design, with Glossosoma densities (natural vs reduced) and sculpin presence (present vs absent) as treatments. Diet analysis revealed that all four prey species were eaten under the natural condition and the sculpin preferred mayfly grazers to Glossosoma. Our experiment showed that although mayfly densities in the presence of either sculpin or Glossosoma were lower than in the no‐sculpin plus reduced‐Glossosoma treatment, no difference in mayfly densities was found between the following three treatments: sculpin plus natural‐Glossosoma, no‐sculpin plus natural‐Glossosoma, and sculpin plus reduced‐Glossosoma. These results indicated that fish predator produced no effects additive to the competitive operation on the mayfly density, and also that competitive operation of Glossosoma on the mayfly densities produced no additional effects to fish predators. In addition, although the competitive effect of the mayflies on Glossosoma could not be manipulated in isolation, the density of Glossosoma in the presence of sculpin was greater than that in the absence of sculpin in the natural‐Glossosoma treatment. Thus, the densities of these competing grazers either stayed the same or increased in the presence of the fish predator relative to predator‐free treatments. A theoretical model, developed to explain the experimental results here, predicted that the densities of the two competing prey under predation pressure could be simultaneously greater than those under predator absent conditions when the behaviorally‐mediated effects of the predator were strongly operative. Although we were unable to distinguish experimentally the two different effects of predator on the prey competition, the behaviorally‐ and density‐mediated effects, the reality of the behaviorally‐mediated effects in the experiment was discussed.     

Diel interactions between prey behaviour and feeding in an invasive fish, the round goby, in a North American river

1. We studied the diet of the invasive round goby (Neogobius melanostomus) on a diel basis in the Flint River, a warmwater stream in Michigan, U.S.A. Diet and available prey samples were collected seven times over a 24 h period in four consecutive months. The section of river studied lacked zebra mussels (Dreissena polymorpha), the primary prey of adult round gobies elsewhere in the Great Lakes region. 2. Diet changed on a diel basis with hydropsychid caddisfly and chironomid larvae predominating during the day, chironomid pupae dominating in the evening and heptageniid mayflies dominating at night. Simultaneous study of macroinvertebrate drift suggested that caddisfly and chironomid larvae were most likely picked from submerged rocks, chironomid pupae were most likely taken during their emergent ascent and mayflies were either captured from the drift or picked from rocks. 3. The Flint River lacks a diverse darter (Family: Percidae) and sculpin (Family: Cottidae) fauna and it appears that the round goby has occupied a generalised darter/sculpin niche. Our results indicate that round gobies have the potential to invade successfully riverine systems, particularly those lacking a diverse benthic fish assemblage.     

Effect of habitat complexity on the predation of Buenoa fuscipennis (Heteroptera: Notonectidae) on mosquito immature stages and alternative prey

Notonectids are well‐known predators in aquatic habitats, where mosquito larvae, chironomids, and cladocerans constitute their main diet. Our purpose was to assess the effect of structural complexity on the predatory ability of Buenoa fuscipennis, a common predator in aquatic habitats of Buenos Aires city (Argentina). Buenoa fuscipennis showed type 2 functional responses in both the presence and absence of prey refuge and no differences in attack rate or handling time between refuge treatments. Regarding mosquito size classes, B. fuscipennis exhibited a significantly higher preference for 2^nd instar larvae and no predation on pupae. In the presence of mosquito larvae and alternative prey, B. fuscipennis preferred mosquitoes over chironomid larvae and adult cladocerans over mosquito larvae. No switching behavior was detected in our experiments. Habitat structure only slightly affected the predator´s consumption rates on mosquito larvae. Overall, preference for prey did not vary with the presence of refuge, except for the preference for mosquitoes over chironomid larvae, which was significantly decreased in the presence of refuge as a consequence of reduced predation on mosquito larvae. The results suggest that B. fuscipennis could efficiently control mosquitoes in structurally simple habitats where chironomids are the most abundant alternative prey but not in temporary pools where cladocerans are abundant.     

Learning to Avoid Dangerous Habitat Types by Aquatic Salamanders,Eurycea tynerensis

There should be intense selection for predation avoidance mechanisms when prey live in close proximity to their predators. Prey individuals that can learn to associate habitat features with high levels of predation risk should experience increased survival if they subsequently avoid those habitats. We tested whether or not habitat learning occurred in a benthic stream community consisting of adult Oklahoma salamander (Eurycea tynerensis) prey and a syntopic predatory fish, the banded sculpin (Cottus carolinae). We exposed individual salamanders to chemical stimuli from sculpin, non‐predatory tadpoles, or a blank control in training tanks containing either rocks or grass. Two days later, the salamanders were tested in tanks that offered a choice of rocks or grass. Salamanders showed significant avoidance of the habitat where they had previously encountered chemical cues from sculpin in comparison to the non‐predatory controls. Learning to avoid dangerous habitats may be particularly important for prey whose predators are visually cryptic ambush foragers, such as sculpin.     

Experimental analysis of the predation impact of the larvae of pond smelt (<Emphasis Type="Italic">Hypomesus transpacificus nipponensis</Emphasis>) on zooplankton populations established in mesocosms

The predation impact of the larvae of pond smelt Hypomesus transpacificus nipponensis on a zooplankton community was studied using mesocosms. The fish significantly depressed the abundances of copepod nauplii and rotifers, especially Hexarthra mira. The vulnerabilities of these prey might be determined by their swimming behavior and population density, suggesting that larval fish selectively prey on zooplankton that have a high encounter rate with the predator. The larvae did not have a negative effect on the densities of cladocerans, but fish predation altered the cladoceran community structure from the dominance of B. longirostris to that of B. fatalis. This result suggests that larval fish predation is an important factor that shifts the species composition of Bosmina in some lakes, the shift occurring in the season when fish larvae are abundant. Our results have shown that predation by the larval fish would control not only the abundance, but also the community structure of the small-sized zooplankton prey.     

The effects of spate-induced disturbance, predation and environmental complexity on macroinvertebrates in a tropical stream

• 1 The combined effects of fish predation, substrate complexity and flow on benthic macroinvertebrates inhabiting riffles was investigated in a Hong Kong stream. Predation was manipulated using fish inclusion/exclusion cages containing complex (= many refuges) or simple (= few refuges) substrates. Experiments were undertaken during the winter dry season, when disturbance due to flow events was minimal, and repeated during the summer monsoon, when the stream experienced spates of varying intensity and duration.
• 2 Predation by the fish, Oreonectes platycephalus, significantly reduced the abundance of macroinvertebrates, especially chironomids and mayflies. Because chironomids (mostly Chironominae) were the dominant cage colonists, there was also a reduction in total macroinvertebrate density.
• 3 Predator impacts were significantly lessened during the wet season, when macroinvertebrate densities increased considerably, but significant reductions in the densities of vulnerable taxa and total macroinvertebrate abundance were nevertheless apparent.
• 4 Substrate complexity (the presence of prey refuges) had no significant effect on the ability of predators to reduce prey abundance.
• 5 Detritus accumulated in cages during the latter part of the study, and densities of most taxa were correlated with detrital standing stocks.
• 6 The results of this experiment indicate that biotic interactions such as predation may be suppressed during periods of spate-induced disturbance, although they can still influence benthic communities significantly. However, the effects of predation are highly taxon specific and may vary among streams in response to changes in predator and prey species composition, or the severity and duration of spates.

     

Predation on meiobenthic assemblages: resource use of a tanypod guild (Chironomidae, Diptera) in a gravel stream

1. Three predatory chironomid species constituted numerically 8.8% (± 95% CL 2.2) of the macro- and meiobenthic community at the sediment surface and in the hyporheic zone of Oberer Seebach, a gravel stream in Lower Austria. Larvae of Thienemannimyia geijskesi (Goetghebuer) and Nilotanypus dubius (Meigen) occurred in higher densities in sediment depths between 10 and 40 cm, whereas Conchapelopia pallidula (Meigen) achieved higher densities at the sediment surface. The three species completed one generation in a year. 2. A total of ninety-seven prey species and instars were identified by gut analyses, of which forty-one benthic rotifer species constituted 69.5% of individuals and twenty-three chironomid species and their instars, 22.9%. The three tanypod species showed shifts from mainly rotifer species in early instars to chironomids and diverse other meio- and macrofaunal taxa in later instars. Rather than shifting towards larger prey sizes, growing predators expanded their upper size thresholds and continued to include smaller prey species in their diet. The extent to which tanypod instars fed on similar prey size classes declined with increasing larval size. Predation by tanypods amounted to 2.2% (± 95% CL 0.1) of the combined prey densities and prey consumption averaged 1.32 (bootstrap 95% CL 1.26–1.39) individuals per predator individual. 3. Preferences for microhabitat flow differed between predator species and in the prey assemblage. Prey densities and densities of T. geijskesi and C. pallidula were highest in pool areas, whereas N. dubius achieved high densities in riffle sites. 4. Tanypod larvae fed non-selectively among prey types. To test the significance of observed size(instar)-specific spatial and dietary overlap values amongst tanypod species, simulations were generated from random models for pairs of intra- and interspecific associations of individuals and groups of prey and predator species. Groups and individuals of tanypod instars fed near randomly on groups of prey types and a high proportion (P > 0.60) of prey individuals are quasi-randomly chosen by tanypods in those patches. Tanypod instar-pairs did not show a sustained trophic resource partitioning in time, thus reducing the degree of competitive interactions for food in this predator guild. Spatially segregated and non-segregated tanypod instars formed random aggregations independent of each other at different flow microhabitats. 5. Species-rich prey assemblages such as benthic rotifers and larval chironomids increased the probability of non-selective feeding upon a wide spectrum of prey species by tanypods. Prey choice was governed by prey availability and tanypod individuals fed on many species at rather even proportions independent of each other.     

Effects of vertical prey distribution on the food-searching behaviour of Danube bleak Chalcalburnus chalcoides mento

Planktivorous fishes frequently have to cope with spatial dynamics of their prey and should be flexible in their vertical food search position. In two experiments Danube bleak Chalcalburnus chalcoides mento, a planktivorous cyprinid, was conditioned to forage on chironomid larvae within the bottom zone and on the cladoceran Leptodora kindtii in the open water of experimental tanks. The fish responded to the presence of chironomids by orienting their snapping behaviour towards the bottom. After most of the prey were depleted, the fish shifted their search position to the open water. Fish were offered L. kindtii at higher densities than the chironomids in the second experiment. With one exception all fish snapped at a high rate into the open water with no observed vertical spatial shifts. One highly active individual showed a clear shift from the open water towards the bottom after depleting most of the L. kindtii. After previous conditioning with chironomids, Danube bleak responded to the presentation of a conditioned stimulus by searching at a deeper position than during control recordings of spontaneous snappings and during a conditioned response test after training with L. kindtii. These results provide preliminary evidence that Danube bleak can detect variation in vertical prey distribution and can build up internal spatial representations.     

Patch based predation in a southern Appalachian stream

Streams are characterized by high degrees of patchiness that could influence the role of predators in these systems. Here we assess the impact of predatory benthic fishes on benthic macroinvertebrate density, biomass, and community structure at the patch scale in a fourth order stream in the southern Appalachians. We tested the role of predation in two different patch types: patches inhabited by adult mottled sculpin ( Cottus bairdi ) and random patches. We placed 30 basket pairs (one open to fish predation, and one from which fish predators were excluded) in the streambed at each patch type. We also tested for potential basket effects by setting up a basket control area. Although there was some evidence of basket artifacts on macroinvertebrate density in sculpin patches, these artifacts were not consistent and we do not feel that they affected our results because predators did not affect macroinvertebrate density. In random patches, predation did not significantly affect macroinvertebrate density or biomass. Predators significantly reduced macroinvertebrate biomass in sculpin patches but did not affect prey density. When the data-set was size-limited to exclude macroinvertebrates too large for consumption by sculpin, macroinvertebrate biomass did not differ significantly between exclusion and open baskets. This suggests that sculpin can reduce macroinvertebrate biomass through a combination of consumption and by predator-induced emigration of large macroinvertebrates into areas that are protected from sculpin. In addition, invertebrate predator biomass was higher in predator exclusion baskets in sculpin patches indicating that predation pressure remained high in the exclusion baskets despite fish exclusion. These results illustrate the heterogeneity of streams and the effect of small-scale differences (e.g. location of predators' territories) on local processes. Experiments that utilize these differences can provide insights into these stream processes.     

Interactions ofMonoporeia affinis (Lindström) (Amphipoda) with sedentary Chironomidae

Interannual population oscillations ofMonoporeia affinis and sedentary chironomids are negatively correlated in Lake Mälaren. Sedentary chironomid abundance regressed againstM. affinis density was highly significant at a lag response of one year (adj. R²=0.54, P=0.0001). The inverse correlation between Tanytarsini (Micropsectra sp. andTanytarsus sp.) andM. affinis densities supports a recruitment limitation hypothesis,viz. that the growth or survival of early instar chironomids is low at high amphipod densities. Microcosm studies showed density-dependent effects on the growth and survival of second instarCh. riparius larvae across an amphipod density gradient. No significant effects were found on growth or survival of third instars. These findings support the chironomid recruitment limitation hypothesis. Amphipod predation on early instar chironomid larvae is suggested as a population controlling mechanism.     

A trophic pathway from biogenic methane supports fish biomass in a temperate lake ecosystem

Although some primary consumers such as chironomid larvae are known to exploit methane‐derived carbon via microbial consortia within aquatic food webs, few studies have traced the onward transfer of such carbon to their predators. The ruffe Gymnocephalus cernuus is a widespread benthivorous fish which feeds predominantly on chironomid larvae and is well adapted for foraging at lower depths than other percids. Therefore, any transfer of methanogenic carbon to higher trophic levels might be particularly evident in ruffe. We sampled ruffe and chironomid larvae from the littoral, sub‐littoral and profundal areas of Jyväsjärvi, Finland, a lake which has previously been shown to contain chironomid larvae exhibiting the very low stable carbon isotope ratios indicative of methane exploitation. A combination of fish gut content examination and stable isotope analysis was used to determine trophic linkages between fish and their putative prey. Irrespective of the depth from which the ruffe were caught, their diet was dominated by chironomids and pupae although the proportions of taxa changed. Zooplankton made a negligible contribution to ruffe diet. A progressive decrease in δ¹³C and δ¹⁵N values with increasing water column depth was observed for both chironomid larvae and ruffe, but not for other species of benthivorous fish. Furthermore, ruffe feeding at greater depths were significantly larger than those feeding in the littoral, suggesting an ontogenetic shift in habitat use, rather than diet, as chironomids remained the predominant prey item. The outputs from isotope mixing models suggested that the incorporation of methane‐derived carbon to larval chironomid biomass through feeding on methanotrophic bacteria increased at greater depth, varying from 0% in the littoral to 28% in the profundal. Using these outputs and the proportions of littoral, sub‐littoral or profundal chironomids contributing to ruffe biomass, we estimated that 17% of ruffe biomass in this lake was ultimately derived from chemoautotrophic sources. Methanogenic carbon thus supports considerable production of higher trophic levels in lakes.     

An experimental study of the effects of predatory fish on macroinvertebrates in a Hong Kong stream

SUMMARY. 1. Predation upon macroinvertebrates by the loach Oreonectes platycephalus Günther (Cobitidae) was studied using predator inclusion/exclusion cages in a series of pools along a Hong Kong stream. Treatments employed were predator exclusion, medium (approximately natural) predator densities (1 fish cage⁻¹) and high predator densities (2 fish cage⁻¹). Macroinvertebrate abundance in cages was monitored after 2 and 4-weeks exposure to predators.
2. The presence of fish was associated with significant declines in the total numbers of macroinvertebrates colonizing cages. However, taxa were influenced differently, with mayflies decreasing by a factor of two while the more mobile shrimps (Atyidae) were unaffected. Chironomid abundance (largely Chironominae) was unaffected by predator density and increased in week 4. Detritus acted as a confounding variable at this time because chironomid abundance was significantly correlated with the weight of accumulated detritus in cages.
3. While invertebrates were more abundant in cages lacking fish, there were no fewer invertebrates in cages with 2 fish than with 1 fish. This may indicate the presence of secure refuges among substrates in the cages, preventing the additional fish from depleting prey further, or a lack of precision of methods due to natural variations in prey densities and spatial patchiness.
4. No significant effects of predators on relative prey abundance or species richness were detected.
5. The impact of predation on prey abundance weakened on week 4, perhaps due to extra refuges among the accumulated detritus. However, drying of the stream increased fish densities in pools so that cages may have become zones of relative safety that were colonized readily by macroinvertebrates. This result highlights the need for year-round investigations to quantify predation effects in Hong Kong's seasonal tropical climate.     

Is coarse woody debris in lakes a refuge or a trap for benthic invertebrates exposed to fish predation?

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Opportunistic foraging by heteropteran mosquito predators

Tropical aquatic environments host a large number of predatory insects including heteropteran water bugs Anisops bouvieri Kirkaldy, 1704 (Heteroptera: Notonectidae), Diplonychus (=Sphaerodema) rusticus Fabricius, 1781 (Heteroptera: Belostomatidae), and Diplonychus (=Sphaerodema) annulatus Fabricius, 1781 (Heteroptera: Belostomatidae) feeding on a range of organisms. In tropical and subtropical wetlands, ponds, and temporary pools these predators play a role in regulation of dipteran populations, particularly mosquitoes and chironomids. Their relative abilities to control mosquitoes depend in part on predator preference for mosquitoes in relation to other natural prey, and the predators’ propensities to switch to mosquitoes as mosquito density increases. The prey electivity and switching dynamics of these predatory water bugs were evaluated in the laboratory under various prey densities, using two instars (II and IV) of chironomid and mosquito larvae as prey. Studies of electivity at relatively high densities (20 prey L⁻¹) in small (5 L) vessels demonstrated that all predators showed opportunistic foraging as the mosquito:chironomid ratio changed, with some evidence that mosquito larvae were positively selected over chironomids. In particular, Anisops showed strong electivity for mosquitoes when presented with any ratio of large mosquito and chironomid prey in the high density experiment, although the preference was not expressed in lower density (2.5 prey L⁻¹) treatments executed in 40 L vessels. In these lower density treatments, D. rusticus demonstrated higher electivity for mosquitoes when the mosquito:chironomid ratio was high, consistent with non-significant trends observed in the higher density experiment. The positive electivity of D. rusticus for mosquitoes was reinforced in an experiment executed over 16 days at varying prey ratios, in which D. rusticus mosquito electivity was high and consistent while D. annulatus showed slight avoidance of mosquito larvae, and Anisops remained largely opportunistic in foraging on prey in proportion with availability. Anisops and D. rusticus are potentially good biocontrol agents for mosquito larvae, in that they preferentially consume mosquitoes under many circumstances but can readily forage on other prey when mosquito density is low.     

Predation of perch on vendace larvae: diet composition in an oligotrophic lake and digestion time of the larvae

The diet of perch Perca fluviatilis was studied to reveal possible predation on vendace Coregonus albula larvae in an oligotrophic lake. Perch diet changed with the size of the fish: small perch ate mainly zooplankton and the diet shifted more to benthic invertebrates and fishes in larger perch. There were also annual and spatial differences in the diet, probably reflecting differences in the availability of prey animals. Perch predation on vendace larvae was only observed in the area with high availability of the larvae. The result suggested strengthened predation when the density of the larvae increases. According to bioenergetics modelling, the perch population increased natural mortality of vendace larvae only marginally. Food intake of spawning female perch was slightly reduced, whereas spawning males fed similarly to non-spawning males. Hence, the spawning period of perch was only a minor refuge for vendace larvae. Laboratory experiments of perch digestion rate demonstrated that, due to rapid digestion of the small fish larvae, diet sampling interval should not be >2 h in the field.     

Effects of Predation Risk on Habitat Selection by Water Column Fish, Benthic Fish and Crayfish in Stream Pools

Predation risk can affect habitat selection by water column stream fish and crayfish, but little is known regarding effects of predation risk on habitat selection by benthic fish or assemblages of fish and crayfish. I used comparative studies and manipulative field experiments to determine whether, (1) habitat selection by stream fish and crayfish is affected by predation risk, and (2) benthic fish, water column fish, and crayfish differ in their habitat selection and response to predation risk. Snorkeling was used to observe fish and crayfish in, (1) unmanipulated stream pools with and without large smallmouth bass predators (Micropterus dolomieui >200 mm total length, TL) and (2) manipulated stream pools before and after addition of a single large smallmouth bass, to determine if prey size and presence of large fish predators affected habitat selection. Observations of microhabitat use were compared with microhabitat availability to determine microhabitat selection. Small fish (60–100 mm TL, except darters that were 30–100 mm TL) and crayfish (40–100 mm rostrum to telson length; TL) had significantly reduced densities in pools with large bass, whereas densities of large fish and crayfish (> 100 mm TL) did not differ significantly between pools with and without large bass. Small orangethroat darters (Etheostoma spectabile), northern crayfish (Orconectes virilis), and creek chubs (Semotilus atromaculatus) showed significantly greater densities in pools without large bass. The presence of large smallmouth bass did not significantly affect depths selected by fish and crayfish, except minnows, which were found significantly more often at medium depths when bass were present. Small minnows and large and small crayfish showed the greatest response to additions of bass to stream pools by moving away from bass locations and into shallow water. Small darters and sunfish showed an intermediate response, whereas large minnows showed no significant response to bass additions. Response to predation risk was dependent on prey size and species, with preferred prey, crayfish and small minnows, showing the greatest response. Small benthic fish, such as darters, are intermediate between small water column fish and crayfish and large water column fish in their risk of predation from large smallmouth bass.