Top-down and bottom-up impacts of juvenile fish in a littoral reed stand

1. Research has often focused on pelagic food chains and processes of lakes; less is known about the contribution of benthic energy flows to whole‐lake ecosystem energetics. This stems from the fact that the shoreline and littoral habitats, which provide a key linkage between sediment and water column, have only recently become a significant focus for study. 2. This study aimed to quantify the feeding and phosphorus allocation of a juvenile fish community in a littoral zone of a shallow lake in response to the biomass succession of the invertebrate prey community. Habitats comprising reed and adjacent open water were sampled over two consecutive years during day and night. 3. Although there were substantial year‐to‐year differences in the biomass of invertebrates, the fish community composition, diet consumption rates and phosphorus allocations were very similar in both study years. Biomasses and predation impacts by juvenile fish on prey groups were substantially higher within the reeds than in the adjacent open water habitat. This may be explained by the refuge‐seeking behaviour of the fish. 4. In general, invertebrates were negligibly influenced by fish feeding, with the exception for a strong top‐down control of large cladocerans. In response to the resulting low Daphnia biomass, fish were forced to switch to a higher degree of benthivory. Consequently, juvenile fish in littoral reed stands may shift benthic‐derived energy and phosphorus via the excretion of soluble reactive phosphorus into the open water.     

Response of littoral invertebrates to reduction of fish density: simultaneous experiments in ponds with different fish assemblages

1. We experimentally reduced densities of predatory fish in replicated 2 m² areas of the littoral zone in two ponds to test whether density and biomass of invertebrates would respond to release from fish predation. The ponds are of similar size and in close proximity, but support different fish assemblages: bluegills ( Lepomis macrochirus Rafinesque) and largemouth bass ( Micropterus salmoides (Lacepede)) in one pond, and bluespotted sunfish ( Enneacanthus gloriosus (Holbrook)) and chain pickerel ( Esox niger Lesueur) in the other. Fish densities were reduced to less than 15% of ambient levels in both experiments.
2. In the bluegill–bass pond, density and biomass of most invertebrate taxa and size classes were unaffected by the fish manipulation. Total invertebrate densities did not differ significantly between fish treatments, but total invertebrate biomass was significantly greater where fish density was reduced, averaging 30% higher over the course of the study. Likewise, manipulation of fish in the bluespotted sunfish–pickerel pond had few significant effects on individual taxa and size classes. There were no significant effects on total invertebrate abundance in the bluespotted sunfish–pickerel pond.
3. Our results provide direct experimental evidence consistent with the collective evidence from previous work, suggesting that the impact of fish predation on density and biomass of invertebrate prey in littoral habitats is variable, but generally weak. Invertebrates that coexist successfully with fish in littoral systems probably are adept at taking advantage of refugia offered by the structurally complex physical environment.     

Response of littoral invertebrates to reduction of fish density: simultaneous experiments in ponds with different fish assemblages

1. We experimentally reduced densities of predatory fish in replicated 2 m² areas of the littoral zone in two ponds to test whether density and biomass of invertebrates would respond to release from fish predation. The ponds are of similar size and in close proximity, but support different fish assemblages: bluegills ( Lepomis macrochirus Rafinesque) and largemouth bass ( Micropterus salmoides (Lacepede)) in one pond, and bluespotted sunfish ( Enneacanthus gloriosus (Holbrook)) and chain pickerel ( Esox niger Lesueur) in the other. Fish densities were reduced to less than 15% of ambient levels in both experiments.
2. In the bluegill–bass pond, density and biomass of most invertebrate taxa and size classes were unaffected by the fish manipulation. Total invertebrate densities did not differ significantly between fish treatments, but total invertebrate biomass was significantly greater where fish density was reduced, averaging 30% higher over the course of the study. Likewise, manipulation of fish in the bluespotted sunfish–pickerel pond had few significant effects on individual taxa and size classes. There were no significant effects on total invertebrate abundance in the bluespotted sunfish–pickerel pond.
3. Our results provide direct experimental evidence consistent with the collective evidence from previous work, suggesting that the impact of fish predation on density and biomass of invertebrate prey in littoral habitats is variable, but generally weak. Invertebrates that coexist successfully with fish in littoral systems probably are adept at taking advantage of refugia offered by the structurally complex physical environment.     

The effects of predation by juvenile fish on the meiobenthic community structure in a natural pond

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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.     

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

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Impact of Holoshestes heterodon Eigenmann (Pisces: Characidae) on the plankton community of a subtropical reservoir: the importance of predation by Chaoborus larvae

We analyzed the effects of planktivorous Holeshestes heterodon Eigenmann (Characidae) predation on the plankton community of a small subtropical reservoir, using four enclosures (volume about 17.5 m³), open to the sediment, established in the littoral zone. Two enclosures were stocked with fish (mean TL 5.7 cm), at a density of about 4–5 fish m^–3 (approx. 8 g m^–3), whereas two remained fishless. The experiment lasted a little longer than one month. In the fish enclosures, most Crustacea and Chaoborus larvae remained scarce, probably as a result of visually selective fish predation. In both fishless enclosures, Chaoborus larvae became abundant. However, in only one of these did large individuals become relatively numerous; this discrepancy in the demographic structure of the Chaoborus populations between the two fishless enclosures is unexplained. Only in the fishless enclosure without appreciable numbers of large Chaoborus did densities of Crustacea increase greatly. It is suggested that in the enclosure containing large Chaoborus individuals, crustacean populations were prevented from developing due to predation pressure, while the small Chaoborus larvae of the other enclosure could not readily consume these prey. Rotifers were low in abundance in the absence of fish, probably as a consequence of Chaoborus predation. Phytoplankton density increased in all four enclosures, due probably to the lack of water flow. Only in the fishless enclosure with high densities of crustaceans did phytoplankton abundance decrease markedly at the end of the experiment, perhaps because of grazing losses.     

Indirect effects of fish on macrophytes in Bays Mountain Lake: evidence for a littoral trophic cascade

Summary We began this experiment to test specific hypotheses regarding direct and indirect effects of fish predation on the littoral macroinvertebrate community of Bays Mountain Lake, Tennessee. We used 24 m² enclosures in which we manipulated the presence and absence of large redear sunfish (Lepomis microlophus>150 mm SL), and small sunfish (L. macrochirus and L. microlophus <50 mm SL) over a 16-mo period. Here we report on effects of fish predation on gastropod grazers that appear to cascade to periphyton and macrophytes.Both large redear sunfish and small sunfish maintained low snail biomass, but snails in fish-free controls increased significantly during the first 2-mo of the experiment. By late summer of the first year of the experiment, the difference in biomass between enclosures with and without fish had increased dramatically (>10×). Midway through the second summer of the experiment, we noted apparent differences in the abundance of periphyton between enclosures containing fish and those that did not. We also noted differences in the macrophyte distribution among enclosures. To document these responses, we estimated periphyton cover, biovolume and cell size frequencies as well as macrophyte distributions among enclosures at the end of the experiment. When fish were absent, periphyton percent cover was significantly reduced compared to when fish were present. Periphyton cell-size distributions in enclosures without fish were skewed toward small cells (only 12% were greater than 200 m³), which is consistent with intense snail grazing. The macrophyte Najas flexilis had more than 60 x higher biomass in the fish-free enclosures than in enclosures containing fish; Potamogeton diversifolius was found only in fish-free enclosures. These results suggest a chain of strong interactions (i.e. from fish to snails to periphyton to macrophytes) that may be important in lake littoral systems. This contrasts sharply with earlier predictions based on cascading trophic interactions that propose that fish predation on snails would enhance macrophyte biomass.     

Food selection of Coregonus lavaretus in a brackish water ecosystem

This study examined how variability in the abundance and biomass structure of benthic invertebrates affected the feeding choice of the whitefish Coregonus lavaretus on a hard bottom habitat of the brackish Baltic Sea. In general, crustaceans such as Idotea balthica and Gammarus spp. were preferred over molluscs. Although being the most numerous taxon in the invertebrate samples, Mytilus trossulus was the lowest ranking in C. lavaretus food preference. The availability of benthic invertebrate prey set the dietary range of fish but the selectivity largely described fish feeding within this range. There was no clear link between fish predation and the dominance structure of benthic invertebrate communities, suggesting that species composition, abundance and biomass of invertebrate species had no impact on the feeding selectivity of the fish. Thus, while fish predation may not affect the dominant species within a benthic community, due to strong selectivity fish may impose strong pressure on some rarer but highly preferred invertebrate prey species.     

Top-down control of a meiobenthic community by two juvenile freshwater fish species

Top-down control of prey assemblages by fish predation has been clearly demonstrated for zooplankton and macroinvertebrates. However, in the benthic communities of freshwater ecosystems, the impact of fish predation on meiofaunal assemblages is nearly unknown. In this study, the predation effects of juvenile carp (Cyprinus carpio) and gudgeon (Gobio gobio) on meiofaunal abundance, biomass, community structure, and the diversity of nematodes were examined using microcosms that were sampled repeatedly over 64 days. Significant differences in abundance and biomass were found between the two fish treatments (carp and gudgeon) and their respective controls for nematodes, oligochaetes, and crustaceans (copepods, harpacticoids, ostracods, and cladocerans), but not for rotifers. These changes were consistent with top-down control of the freshwater meiofaunal assemblages in the microcosms over time. By contrast, small-bodied meiofauna was more abundant, suggesting indirect facilitation. Neither the species richness nor the diversity of the nematode community was affected by fish predation. The results indicate that predation by juvenile freshwater fish depresses the overall abundance and biomass of meiofaunal assemblages, except for rotifers, and alters the size structure of the meiofaunal community. Therefore, the meiofaunal assemblages of freshwater ecosystems may be influenced by bottom-feeding juvenile fish, e.g., carp and gudgeon, through top-down control of meiofaunal populations.     

Body size and substrate association of littoral insects in relation to vegetation structure

Variation in substrate association types and maximum size of aquatic insects were studied in a vegetated littoral zone of three lake basins. The basins differed from each other in trophic status, biomass of benthivorous fish, and abundance of macrophytes. Four types of substrate association – swimmers, crawlers, semisessiles and burrowers, respectively – were assumed to represent decreasing vulnerability to fish predators. Large-sized species were also hypothesised to be more vulnerable to fish predators. The distributions of species traits were examined in relation to vegetation density. Inferring from ``predation hypothesis' opposite selection pressures on the species traits were expected along the vegetation density. Dense macrophyte beds were thought to be dominated by invertebrate predators and open water by fish predators, since the predation efficiency of fish decreases in complex environments. In the case of invertebrate predator domination, large size and higher activity should be favoured traits among the prey species. Distribution patterns of modes of the two studied traits were explored separately for predatory and non-predatory insects. As expected, swimmers and large-sized crawlers were characteristic of the insect assemblages of dense macrophyte beds. The densities of Odonata, Corixidae, Dytiscidae, Ephemeroptera and Sialidae were higher among macrophytes than in open water, where these insect taxa were possibly depleted by fish. On the other hand, the small-sized and fairly immobile Chironomidae were the most abundant group in open water. These results support the existence of a predator transition zone among littoral vegetation, ranging from domination of invertebrate predation among the dense beds to that of fish predation in open water.     

Influences of slimy sculpin (Cottus cognatus) predation on the rocky littoral invertebrate community in an arctic lake

We tested the role of the slimy sculpin (Cottus cognatus), a benthic fish, in structuring the rocky littoral invertebrate community in Toolik Lake, Alaska. Comparisons of sculpin gut contents and prey community structure indicated that these fish forage selectively, eating proportionally more large and motile prey, and proportionally fewer small and sessile forms. Field experiments compared the effects of natural, reduced and elevated sculpin densities on benthic community structure. At natural levels of sculpin density, biomass of trichopteran larvae were reduced by more than 50%, and predatory chironomid larvae by 27%, in comparison to areas where sculpin were excluded. Tube-dwelling and small free living chironomid larvae were unaffected at normal sculpin densities. Under artificially high sculpin densities, there was some reduction of tube-dwelling chironomids, but the small free living ones remained unaffected. There appears to be a threshold length of about 3.5 mm, below which chironomid larvae are free form sculpin predation. Tube-dwelling chironomids may be longer than this threshold, but still avoid predation by having most of their body hidden in their tubes.     

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

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Seasonal influence of brook trout and mottled sculpin on lower trophic levels in an Appalachian stream

1. In some situations fish have strong top‐down effects in stream communities while in others they seem to be relatively unimportant. Differences in the impact of fish may depend on a variety of factors including the foraging mode of the fish, interactions among fish species and temporal variation in environmental conditions and species interactions. 2. We investigated the effect of brook trout (Salvelinus fontinalis) and mottled sculpin (Cottus bairdi) on lower trophic levels in Appalachian streams and whether or not interactions between these fish changed their influence. Mesocosms were placed in a headwater stream in a randomized complete block design. Within blocks, mesocosms were randomly assigned to one of the following treatments: (i) no fish; (ii) sculpin only; (iii) trout only and (iv) both sculpin and trout. Fish biomass was the same in all three fish treatments. Invertebrate density and algal biomass in mesocosms were determined after 3 weeks. We repeated the experiment in the autumn, spring and summer to test for seasonality of fish effects. 3. The effect of fish on invertebrate assemblages was seasonal and depended on prey identity. Sculpin strongly suppressed grazer abundance in spring while trout had little effect on grazers in any season. The influence of both fish on insect predators was similar and relatively constant across seasons. We found little evidence of an interaction between sculpin and trout that strongly influenced their effect on prey across seasons. 4. None of the fish treatments influenced algal biomass during any of the seasons. Algal growth was also seasonal, with a two‐ to four‐fold increase in algal biomass in spring compared to autumn and summer. 5. Our results indicate that benthic and drift feeding fish differ in their effects on some, but not all prey. Furthermore, fish effects on prey were strongly seasonal for some, but not all prey types. While the temporal context is not commonly considered, our results indicate seasonality can be an important component of predator–prey interactions in streams.     

Long-term changes in littoral fish community structure and resilience of total catch to re-oligotrophication in a large,peri-alpine European lake

1. The littoral zone of lakes is used as spawning, shelter, or feeding habitat for many fish species and hence is of key importance for overall lake functioning. Despite this, hardly any studies exist examining the long-term dynamics and response of the littoral fish community, composed mostly of juvenile fish, to environmental change. Here, we study the response of total catch per unit effort (CPUE) and individual species CPUE of such a community to 17 years of oligotrophication and examine whether the species responses can be characterised as synchronous or asynchronous.
2. We analyse a data set of beach seine catches carried out during morning and twilight, late spring and late summer at three sites in large and deep Lake Constance from 1997 to 2014. Generalised additive mixed models were used to explore changes in CPUE of the overall community and of the most frequently occurring species, and Kendall's W test was applied to examine whether the dynamics of fish species were synchronous or asynchronous.
3. Species-specific and total CPUE strongly differed between morning and twilight and between spring and summer indicating an important role of behavioural and life cycle adaptations of species for CPUE. In addition, also the CPUE of some species seeking shelter behind larger stones was lower at sites without these.
4. Total CPUE did not decline suggesting the overall abundance of littoral fish was resilient to declining nutrients. In contrast, fish community composition changed strongly during the study period due to increases in some species (dace, loach, perch) and decreases in others (bream, burbot, chub, ruffe), indicating response diversity of fish to oligotrophication. The type of community dynamics was scale-dependent, whereby significantly synchronous dynamics according to Kendall's W were observed when taking seasonal variability into account. In contrast, significantly asynchronous species dynamics were observed when only the low-frequency variability of species dynamics was considered separately for spring and summer time series.
5. Resilience of littoral fish total CPUE to oligotrophication might have important consequences for ecosystem dynamics and ecosystem services beyond the littoral zone. As small fish often impose strong predation pressure on zooplankton, their resilience might sustain a high top-down control on zooplankton resulting in a further reduction of zooplankton biomass. This could contribute to delayed food web responses and reduced growth of fish with oligotrophication.

     

Predator evasion in <Emphasis Type="Italic">Daphnia</Emphasis>: the adaptive value of aggregation associated with attack abatement

To evaluate the adaptive value of daphnian aggregation associated with attack abatement, a combined laboratory and field experiment was performed. As the lake investigated does not contain planktivorous fish, only invertebrate predation is important. In the laboratory, newly collected Chaoborus flavicans, among the most important predators to exploit the Daphnia population in the lake, were individually placed in a spherical flask containing different densities of juvenile Daphnia prey. The number of successful attacks of Chaoborus was independent of prey abundance. Thus, in a given density regime, each individual prey gained protection from a dilution effect. In the field, dense aggregates of Daphnia were created by continuously adding food at a localized point in the littoral. Even though the food input produced aggregates of Daphnia which were maintained for 3 h, the number of invertebrate predators did not increase in the input area. It is concluded that each Daphnia in a group reduces the probability of predation through an effect of attack abatement which is approximately proportional to the number of prey in the group. However, when considering the high median density of Daphnia, together with their strong response to food input, it seems more plausible that natural aggregates are driven by a patchy food distribution, whereas attack abatement is only a secondary effect of aggregation.     

In situ experiments on the effects of increased sediment loads on littoral rocky shore communities in Lake Tanganyika, East Africa

1. Two in situ experiments investigated the responses to artificially increased sediment loads of the gastropod and fish communities, and of the whole benthos, of the rocky shores of Lake Tanganyika. Sediments were emptied once onto randomly selected quadrats using SCUBA, with control quadrats receiving no addition of sediment. Quadrats were monitored up to 6 months after sediment addition. 2. While the abundance of Lavigeria grandis (Mollusca; Gastropoda) was reduced greatly by the addition of sediment, low abundance and biomass of the whole gastropod community on sediment‐impacted quadrats 6 months after sediment addition suggests more general impacts of inundation in the longer term. 3. Benthic invertebrate communities inundated by sediment had significantly fewer individuals and numbers of taxa than control communities. No significant community recovery was recorded on sediment‐impacted quadrats within the timescale of the experiment. A short‐term period of high turbidity brought about by the emptying of sediments onto sediment‐impacted quadrats also caused severe decreases in benthic invertebrate abundance on controls 10 days after sediment addition. This suggests that even short‐term pulses of sediment may have significant detrimental effects on littoral communities. 4. Although significant impacts of sediment on fish community dynamics were found, and a significant reduction in the number of benthic algivorous species was recorded immediately after sediment addition, littoral fish communities showed strong resistance overall to the addition of sediments. This apparent resistance, however, may be a consequence of the small scale at which the experiment took place. Littoral fish communities were also shown to be more dynamic than indicated previously.     

Crayfish in lakes and streams: individual and population responses to predation, productivity and substratum availability

1. In a correlative study, we investigated the relative importance of fish predation, refuge availability and resource supply in determining the abundance and size distributions of the introduced and omnivorous signal crayfish (Pacifastacus leniusculus) in lakes and streams. Moreover, the biomass and food selection of predatory fish was estimated in each habitat type and stable isotopes of carbon and nitrogen were measured in perch (Perca fluviatilis), the dominant predator in the lakes, and in its potential food sources (crayfish, juvenile roach and isopods). 2. In lakes, crayfish were the most frequent prey in large perch (46%), followed by other macroinvertebrates (26%, including the isopod Asellus aquaticus) and small fish (25%). Crayfish and fish dominated the gut contents of large perch with respect to biomass. Nitrogen signatures showed that perch were one trophic level above crayfish (approx. 3.4‰) and a two‐source mixing model using nitrogen isotope values indicated that crayfish (81%) contributed significantly more to perch isotope values than did juvenile roach (19%). A positive correlation was found between the abundance of crayfish and the biomass of large perch. Crayfish abundance in lakes was also positively correlated with the proportion of cobbles in the littoral zone. Lake productivity (chlorophyll a) was positively correlated with crayfish size, but not with crayfish abundance. 3. In streams, brown trout (Salmo trutta) were the most abundant predatory fish. Gut contents of large trout in a forested stream showed that terrestrial insects were the most frequently found prey (60%), followed by small crayfish (27%) and isopods (27%). In contrast to lakes, the relative abundance of crayfish was negatively correlated with the total biomass of predatory fish and with total biomass of trout. However, abundance of crayfish at sites with a low biomass of predatory fish varied considerably and was related to substratum grain size, with fewer crayfish being caught when the substratum was sandy or dominated by large boulders. The mean size of crayfish was greater at stream sites with a high standing stock of periphyton, but neither predator biomass nor substratum grain size was correlated with crayfish size. 4. Our results suggest that bottom‐up processes influence crayfish size in lakes and streams independent of predator biomass and substratum availability. However, bottom‐up processes do not influence crayfish abundance. Instead, substratum availability (lakes) and interactions between predation and substratum grain size (streams) need to be considered in order to predict crayfish abundance.     

Interspecific differences in drift behaviour between the native <Emphasis Type="Italic">Gammarus pulex</Emphasis> and the exotic <Emphasis Type="Italic">Gammarus roeseli</Emphasis> and possible implications for the invader’s success

“Drifting” is known to subject aquatic invertebrates to intense predation by drift feeding fish. Consequently, interspecific variations in drifting behaviour could lead to differences in predation pressure between coexisting prey species. Predation being an important factor determining the success of invaders, differences in drift patterns could advantage either native or exotic invertebrates through differential predation by native fish predators. The exotic freshwater amphipod (Gammarus roeseli) has now largely colonized Western Europe where it is often found in sympatry with a native species (Gammarus pulex). Here we documented interspecific differences in drifting behaviour that might have favored the invader’s success through differential predation. Benthic and drifting amphipods were sampled three times at the same site to compare the proportion of each species within and between sample types (benthos or drift) across time. Compared with the benthos, where the invader (G. roeseli) was significantly less abundant than the native (G. pulex), G. roeseli was proportionally overrepresented in the drift but displayed a very different drifting pattern. While G. pulex drift rates remained roughly constant over a 24 h period, G. roeseli showed a marked diel periodicity with low diurnal and high nocturnal drift rates. Such drifting behaviour could procure this species with a competitive advantage regarding predation as most drift feeding fish are diurnal. As a result, the native appears more disadvantaged with respect to drift. This may partly explain the ability of G. roeseli to coexist with G. pulex in a habitat more suitable to the native.     

Influence of fishes on macroinvertebrate communities and insect emergence production in intermittent stream refuges

1. Drying intermittent stream networks often have permanent water refuges that are important for recolonisation. These habitats may be hotspots for interactions between fishes and invertebrates as they become isolated, but densities and diversity of fishes in these refuges can be highly variable across time and space.
2. Insect emergence from streams provides energy and nutrient subsidies to riparian habitats. The magnitude of such subsidies may be influenced by in-stream predators such as fishes.
3. We examined whether benthic macroinvertebrate communities, emerging adult insects, and algal biomass in permanent grassland stream pools differed among sites with naturally varying densities of fishes. We also manipulated fish densities in a mesocosm experiment to address how fishes might affect colonisation during recovery from hydrologic disturbance.
4. Fish biomass had a negative impact on invertebrate abundance, but not biomass or taxa richness, in natural pools. Total fish biomass was not correlated with total insect emergence in natural pools, but orangethroat darter (Etheostoma spectabile) biomass was inversely correlated with emerging Chironomidae biomass and individual midge body size. The interaction in our models between predatory fish biomass and date suggested that fishes may also delay insect emergence from natural pools, altering the timing of aquatic–terrestrial subsidies.
5. There was an increase over time in algal biomass (chlorophyll-a) in mesocosms, but this did not differ among fish density treatments. Regardless, fish presence in mesocosms reduced the abundance of colonising insects and total invertebrate biomass. Mesocosm invertebrate communities in treatments without fishes were characterised by more Chironomidae, Culicidae, and Corduliidae.
6. Results suggest that fishes influence invertebrates in habitats that represent important refuges during hydrologic disturbance, hot spots for subsidy exports to riparian food webs, and source areas for colonists during recovery from hydrologic disturbance. Fish effects in these systems include decreasing invertebrate abundance, shifting community structure, and altering patterns of invertebrate emergence and colonisation.