Predator effects on prey population dynamics in open systems

Animal population dynamics in open systems are affected not only by agents of mortality and the influence of species interactions on behavior and life histories, but also by dispersal and recruitment. We used an extensive data set to compare natural loss rates of two mayfly species that co-occur in high-elevation streams varying in predation risk, and experience different abiotic conditions during larval development. Our goals were to generate hypotheses relating predation to variation in prey population dynamics and to evaluate alternative mechanisms to explain such variation. While neither loss rates nor abundance of the species that develops during snowmelt (Baetis bicaudatus) varied systematically with fish, loss rates of the species that develops during baseflow (Baetis B) were higher in streams containing brook trout than streams without fish; and surprisingly, larvae of this species were most abundant in trout streams. This counter-intuitive pattern could not be explained by a trophic cascade, because densities of intermediate predators (stoneflies) did not differ between fish and fishless streams and predation by trout on stoneflies was negligible. A statistical model estimated that higher recruitment and accelerated development enables Baetis B to maintain larger populations in trout streams despite higher mortality from predation. Experimental estimates suggested that predation by trout potentially accounts for natural losses of Baetis B, but not Baetis bicaudatus. Predation by stoneflies on Baetis is negligible in fish streams, but could make an important contribution to observed losses of both species in fishless streams. Non-predatory sources of loss were higher for B. bicaudatus in trout streams, and for Baetis B in fishless streams. We conclude that predation alone cannot explain variation in population dynamics of either species; and the relative importance of predation is species- and environment-specific compared to non-predatory losses, such as other agents of mortality and non-consumptive effects of predators. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Balancing risks? Responses and non-responses of mayfly larvae to fish and stonefly predators

Summary In a series of laboratory experiments we examined the hypothesis that larvae of stream mayflies would respond to the presence of two different types of predators in such a way as to minimize their risk of being consumed by each. Positioning of larvae (whether they frequent the top, sides, or bottom of stones) of Baetis tricaudatus and Ephemerella subvaria was altered by the presence of predaceous stoneflies (Agnetina capitata) with a larger proportion of the population occurring on the upper surfaces, where the probability of encountering the predator was lowest. The presence of a benthivorous fish (Cottus bairdi) had no significant effects on positioning of the mayfly larvae. Lack of fish effects may reflect an inability of the mayflies to detect or respond to sculpins, or alternately may indicate that sculpins do not normally present a important predation risk for these mayflies. Failure of mayfly prey to account for fish predators when responding to the presence of stoneflies appcars to explain facilitation previously observed between stoneflies and sculpins.     

The influence of predatory fish on mayfly drift: extrapolating from experiments to nature

1. A knowledge of how individual behaviour affects populations in nature is needed to understand many ecologically important processes, such as the dispersal of larval insects in streams. The influence of chemical cues from drift‐feeding fish on the drift dispersal of mayflies has been documented in small experimental channels (i.e. < 3 m), but their influence on dispersal in natural systems (e.g. 30 m stream reaches) is unclear. 2. Using surveys in 10 Rocky Mountain streams in Western Colorado we examined whether the effects of predatory brook trout (Salvelinus fontinalis) on mayfly drift, that were apparent in stream‐side channels, could also be detected in natural streams. 3. In channel experiments, the drift of Baetis bicaudatus (Baetidae) was more responsive to variation in the concentration of chemical cues from brook trout than that of another mayfly, Epeorus deceptivus (Heptageniidae). The rate of brook trout predation on drifting mayflies of both species in a 2‐m long observation tank was higher during the day (60–75%) but still measurable at night (5–10%). Epeorus individuals released into the water column were more vulnerable to trout predation by both day and night than were Baetis larvae treated similarly. 4. Drift of all mayfly taxa in five fishless streams was aperiodic, whereas their drift was nocturnal in five trout streams. The propensity of mayflies to drift was decreased during the day and increased during the night in trout streams compared with fishless streams. In contrast to the channel experiments, fish biomass and density did not alter the nocturnal nature nor magnitude of mayfly drift in natural streams. 5. In combination, these results indicate that mayflies respond to subtle differences in concentration of fish cues in experimental channels. However, temporal and spatial variation in fish cues available to mayflies in natural streams may have obscured our ability to detect responses at larger scales.     

How do grazers affect periphyton heterogeneity in streams?

The effects of grazing by stream invertebrates on algal biomass and spatial heterogeneity were tested experimentally in flow-through microcosms with natural substrates (rocks). One experiment tested the effects of fixed densities of three species of grazers (the caddisfly Allomyia sp. and two mayflies, Epeorus deceptivus and Baetis bicaudatus) on periphyton. Baetis was tested with and without chemical cues from fish predators, which reduced grazer foraging activity to levels similar to the less mobile mayfly (Epeorus). Mean algal biomass (chlorophyll a; chl a) was reduced in grazer treatments compared to ungrazed controls, but there were no differences among grazer treatments. Algal heterogeneity (Morisita index) increased with grazer mobility, with the highest heterogeneity occurring in the Baetis-no fish treatment (most mobile grazer) and the lowest in the caddisfly treatment (most sedentary grazer). A second experiment used a three factorial design, and tested whether initial resource distribution (homogeneous vs. heterogeneous), Baetis density (high vs. low) and fish odor (present vs. absent) affected grazer impact on algal resources. Abundances of Baetis and chl a on individual rocks were recorded to explore the mechanisms responsible for the observed distributions of algae. Initial resource heterogeneity was maintained despite being subjected to grazing. Mean chl a was highest in controls, as in experiment I, and effects of Baetis on algal biomass increased with grazer density. There were no fish effects on algal biomass and no effects of grazer density or fish on algal heterogeneity. At the scale of individual rocks Baetis was unselective when food was homogeneously distributed, but chose high-food rocks when it was heterogeneously distributed. Results of these mechanistic experiments showed that Baetis can track resources at the scale of single rocks; and at moderate densities mobile grazers could potentially maintain periphyton distributions observed in natural streams.     

Differential behavioural responses of mayflies from streams with and without fish to trout odour

1. In streams, mayflies (Order Ephemeroptera) are at risk from fish feeding visually in the water column. The effect of fish odour on the behaviour of Baetis bicaudatus from a fishless stream and a trout stream was investigated in four large oval tanks supplied with water from the fishless stream.
2. For each mayfly population, mayfly positioning on the substratum and movement in the water column (drift) were measured during the day and night, over 3 days. Brook trout ( Salvelinus fontinalis ) odour was added to two tanks to test the effect of a threat from fish.
3. Throughout the experiment more mayflies from the trout stream were observed on the substratum surface and in the water column during the night than the day, but the magnitude of night drift was less in tanks with fish odour.
4. Baetis from the fishless stream also displayed a nocturnal periodicity in drift and positioning, but their night-time drift was not affected by the presence of fish odour. On the first day of the experiment, however, more mayflies were observed on the substratum surface and drifting in tanks without fish odour during the day.
5. Sensitivity to fish odour may enable mayflies to alter their behaviour according to the risk of predation from fish.     

Differential behavioural responses of mayflies from streams with and without fish to trout odour

1. In streams, mayflies (Order Ephemeroptera) are at risk from fish feeding visually in the water column. The effect of fish odour on the behaviour of Baetis bicaudatus from a fishless stream and a trout stream was investigated in four large oval tanks supplied with water from the fishless stream.
2. For each mayfly population, mayfly positioning on the substratum and movement in the water column (drift) were measured during the day and night, over 3 days. Brook trout ( Salvelinus fontinalis ) odour was added to two tanks to test the effect of a threat from fish.
3. Throughout the experiment more mayflies from the trout stream were observed on the substratum surface and in the water column during the night than the day, but the magnitude of night drift was less in tanks with fish odour.
4. Baetis from the fishless stream also displayed a nocturnal periodicity in drift and positioning, but their night-time drift was not affected by the presence of fish odour. On the first day of the experiment, however, more mayflies were observed on the substratum surface and drifting in tanks without fish odour during the day.
5. Sensitivity to fish odour may enable mayflies to alter their behaviour according to the risk of predation from fish.     

Can current velocity mediate trophic cascades in a mountain stream?

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Stonefly nymphs use hydrodynamic cues to discriminate between prey

Summary Playback experiments conducted in a Rocky Mountain, USA, stream determined whether predatory stonefly nymphs (Kogotus modestus; Plecoptera: PerlodiMae) used hydrodynamic cues to discriminate prey species from nonprey species. In the laboratory we recorded pressure wave patterns associated with swimming escape behavior of Baetis bicaudatus (Baetidae), the favored mayfly prey species, and those of a nonprey mayfly, Ephemerella infrequens (Ephemerellidae). We video taped the responses of 24-h starved Kogotus to Baetis playbacks, Ephemerella playbacks or no playbacks made by oscillating (or not) live mayflies (Ephemerella) or clear plastic models placed within in situ flow-through observation boxes. The probability of attacks per encounter with Baetis playbacks was highest and independent of the model type used, but Kogotus also showed an unexpected high probability of attacks per encounter when Ephemerella playbacks were made through live Ephemerella. Thus, Kogotus discriminated between Baetis and Ephemerella swimming patterns but only when playbacks were made through the plastic model. Kogotus never attacked motionless mayflies or motionless plastic models. We allowed some Kogotus to successfully capture one small Baetis immediately before playbacks, which resulted in a much higher probability of attacks per encounter with Baetis playbacks on either model and a heightened discrimination of prey versus nonprey playbacks. The probability of attacks per encounter by Kogotus with live Baetis swimming under similar experimental conditions was strikingly similar to its response to Baetis playbacks made by oscillating the plastic model after a successful capture. Order of playback presentation (Baetis first or Ephemerella first) did not influence predatory responses to mayfly swimming patterns. This study is the first to document the use of hydrodynamic cues by stream-dwelling predators for discrimination of prey from nonprey and provides a mechanism to explain selective predation by stoneflies on Baetis in nature.     

A laboratory study of predator-prey interactions of stoneflies and mayflies

SUMMARY. 1. Reactions of individual predatory stoneflies to contact by individual mayfly prey, and vice versa, were studied in the laboratory using two sets of species, one from a stream in southern Ontario. Canada, and the other from a stream in North Wales. 2. Based largely on information received via their antennae, the stoneflies were capable of distinguishing between conspecifics and mayfly prey and between prey species. A clear preference for certain prey species was evident but less preferred species were made more attractive when damaged or when the stoneflies were very hungry. Detection of conspecifics by stoneflies was confused in the presence of prey or the local smell of prey body fluids and this led to interference during hunting and feeding at high predator densities. 3. Some mayfly species were capable of distinguishing between conspecifics. other mayflies and predators. The cerci were important in intra- and interspecific contact encounters between mayflies and were used in a fencing fashion to establish a minimum distance between nymphs. However, the cerci did not appear to play an anti-predation role as stoneflies captured and ate mayflies with and without cerci in equal numbers. 4. Reactions to conspecifics by the Palaearetie mayflies Rhithrogena semicolorata (Curt.) and Baetis rhodani (Pictet) were similar to those of the ecologically-equivalent Nearctic species Stenonema vicarium (Walker) and Baetis vagans McDunnough. 5. In encounters between mayflies, intra- and interspecific responses were similar. However, the responses of some mayfly species to conspeeifics were modified in the physical presence of a predator or the water from its holding tank. Different mayfly species made specific responses to predaeeous stoneflies. Responses to a herbivorous stonefly were either similar to conspeeific encounters or were intermediate between them and encounters with predators. 6. Experiments to reveal the exact nature of the stonefly stimulus detected by the mayfly nymphs strongly suggested that there were two components to the stimulus. One was physical contact by the predator's antennae, the other was chemical in nature and could be detected only over short distances (up to a few mm). Independently each could elicit a response, but combined their effect appeared to be synergistic. 7. Different mayfly species responded differently to the stonefly Phasganophora capitata (Pictet) and this sequence of sensitivity was mirrored by the predator's choice of prey species—the most sensitive to the predator being the favoured prey and the less sensitive the least favoured. The significance of these findings to optimal foraging theory, as applied to tactile predators, is discussed.     

Ecosystem engineering by beavers affects mayfly life histories

1. The North American beaver has been studied as a model ecosystem engineer for many decades. Previous studies have documented physical, chemical and biological impacts attributed to beaver engineering in both aquatic and terrestrial environments. This study focused on the effects of ecosystem engineering by beavers on life histories of a common mayfly and on the potential consequences for mayfly populations. 2. We studied 18 montane beaver ponds of varying size and shape in western Colorado near the Rocky Mountain Biological Laboratory. Our goal was to test whether variation in beaver pond morphology (pond size and shape) explains downstream changes in stream temperature, mayfly size and timing of emergence. 3. Downstream water temperatures varied predictably with pond morphology, being colder downstream of high‐head dams and warmer downstream of low‐head dams. Pond morphology was also a significant predictor of variation in the size of mature female Baetis bicaudatus (the most abundant mayfly), with larger females emerging downstream of high‐head dams and smaller females downstream of low‐head dams. The size of male B. bicaudatus was not significantly related to pond morphology or stream temperature. There was no relationship between pond morphology and variation in the timing of emergence of Baetis (males or females) between upstream and downstream reaches. 4. Our results have implications for the effects of beaver ponds on Baetis individual fitness because large Baetis females are more fecund. Therefore, predictable female size variation associated with beaver pond morphology makes it possible to model the effects of beaver activity on local contributions of Baetis to the regional pool of reproductive adults at the catchment scale. Additionally, predictable changes in the size of emerging mayflies may have important consequences for the magnitude of aquatic to terrestrial resource subsidies in beaver‐modified systems.     

Nested species subsets, gaps, and discrepancy

Chemical cues from fish can alter the behaviour of stream invertebrates in experimental tanks but their effect in natural streams has received little attention. By adding brook trout (Salvelinus fontinalis) odour to a trout stream in the Rocky Mountains of Colorado, USA, we tested whether changes in the concentration of chemical cues from visually feeding predatory fish would alter the drift of mayfly nymphs (Ephemeroptera). Stream water was piped from stream-side tanks with (odour) and without (control) three brook trout to two locations in the stream 3.5 m upstream of drift nets at six replicate sites. Five-minute drift samples were collected downstream from odour and control pipes before, during and after the release of water from the tanks into the stream during both the day and night. Almost all drift occurred at night and consisted predominantly of Baetis bicaudatus nymphs. The odour manipulation had no measurable effect on Baetis drift during the day but statistical power was low. During the night, however, the drift of large (>0.65 mm head capsule width, HCW) Baetis nymphs decreased significantly during the odour addition compared to control drift. In contrast, the drift of small nymphs (≤0.65 mm HCW) increased both during and after the odour addition in comparison to control drift. Since the stream contains brook trout (0.04–0.18 m⁻²), and water from the stream (presumably containing fish odour) altered the behaviour of fishless-stream Baetis nymphs in another experiment, we conclude that the changes in Baetis drift density were a response to an increase in the concentration of fish odour in the stream. Furthermore, we were able to detect the effect within 5 min. of odour addition, indicating that mayfly behavioural response to trout odour was rapid. These results suggest that mayflies can distinguish different concentrations of trout odour in natural streams and that the response is size-specific, according to the relative risk of predation of large and small Baetis. Received: 12 May 1998 / Accepted: 23 October 1998     

Does light intensity modify the effect mayfly grazers have on periphyton?

1. A factorial experiment was conducted in artificial outdoor streams to quantify the effects of irradiance (two levels) and two mayfly grazers (four densities of each) on periphytic community structure. The mayflies were Ecdyonurus venosus (Heptageniidae), a grazer using brushing mouthparts, and Baetis spp. (Baetidae) a grazer which uses mandibles and maxilla to scrape and gather periphyton. The experiment ran for 16 days. 2. Grazer densities in channels approximated those existing in a shoreline habitat in the River Sihl, Switzerland. Light treatments were natural (daily mean = 810 μmol m^–2 s^–1) and shaded (daily mean = 286 μmol m^–2 s^–1). 3. Higher irradiance increased total algal abundance by a factor of 4. Algae most affected were prostrate/motile and erect diatoms, filamentous chlorophytes and Hydrurus foetidus. 4. Both species of mayfly reduced periphytic and algal biomass. Mayfly–mayfly interactions, however, were associated with statistical increases in algal biovolume and chlorophyll-a content, indicating that the two grazers may have interfered with one another as their densities increased. The mayfly–mayfly interaction did not influence periphytic ash-free dry mass (AFDM). Light modified the influence of Ecdyonurus such that this mayfly produced greater reductions in algal biovolume under high irradiance. 5. Despite efforts to exclude other grazers, chironomids colonized experimental channels. Chironomid biomass was approximately eight times less than mayflies across treatments and was positively correlated with all measures of periphytic abundance, suggesting that these grazers were responding to periphyton rather than controlling it. Chironomids were also associated with an increase in the abundance of diatoms having a prostrate/motile physiognomy. The only physiognomy to show a negative relationship with chironomid biomass was the thallus type, a form which comprised less than 1% of the algal biovolume across channels. 6. Ecdyonurus and Baetis had distinct influences on algal physiognomy. Ecdyonurus, for example, reduced adnate, stalked and Achnanthes-type physiognomies, but was associated with a significant increase in the abundance of filamentous chlorophytes (primarily Ulothrix sp.). Baetis reduced erect, Achnanthes-type and thallus physiognomies. Neither mayfly influenced the abundance of prostrate/motile diatoms; a physiognomy that comprised 21% of the algae in channels. 7. Light and mayfly interactions affected algal community structure. The interaction of Ecdyonurus with light had a negative effect on erect diatoms, filamentous chlorophytes and the thallus physiognomy, but a positive effect on stalked and Achnanthes-type physiognomies. Baetis interacting with light had a positive effect on adnate diatoms. 8. Although both mayfly taxa influenced periphytic community structure, physiognomy was not a good predictor of algal susceptibility to grazing. The type of substratum to which an alga is attached (detritus or algal filaments vs hard surfaces) and location within the periphytic matrix may be better indicators of vulnerability to grazing than physiognomy.     

The influence of moss on grazers in high‐altitude streams: food,refuge or both?

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Predator detection and avoidance by lotic mayfly nymphs of different size

We studied antipredatory responses of lotic mayfly (Baetis) nymphs in a factorial experiment with four levels of fish presence: (1) a freely foraging fish (the European minnow,Phoxinus phoxinus), (2) a constrained fish, (3) water from a fish stream, (4) water from a fishless stream. LargeBaetis nymphs drifted mainly during night-time in treatments involving either the chemical or actual presence of fish, whereas no diel periodicity was observed when the water was not conditioned with fish odour. The response was strongest when the fish was uncaged, which suggests that visual or hydrodynamic cues are needed in addition to chemical ones for an accurate assessment of predation risk. Fish presence had no effect on the drift rates of small nymphs. Instead, they increased their refuge use in the presence of a live fish. Chemical cues alone did not have any effect on the refuge use of any of theBaetis size classes. Our results indicate active drift entry by mayfly nymphs. Because predation pressure is spatially and temporally variable, nymphs must sample the environment in order to locate predator-free areas or areas with low predation risk. Drifting should be the most energy-saving way to do this. To avoid the risk from visually feeding fish, large individuals can sample safely (i.e. enter drift) only at night-time, while the small ones can also do this safely during the day. We suggest that, contrary to some earlier assumptions, mayfly drift is not a fixed prey response. Instead,Baetis nymphs are able to assess the prevailing predation pressure, and they adjust their foraging behaviour accordingly.     

Mechanisms of prey selection by predaceous stoneflies: roles of prey morphology,behavior and predator hunger

Summary Laboratory feeding experiments using Hesperoperla pacifica (Banks), Perlidae, and Megarcys signata (Hagen), Perlodidae, as predators and Baetis tricaudatus Dodds and Ephemerella altana Allen as prey indicate a strong effect of prey morphology and mobility and predator hunger on prey selection by stoneflies. Knowledge of both dietary composition and feeding behavior was necessary to fully understand prey selection by these stoneflies.Fasted stoneflies presented with live prey ate more E. altana while satiated stoneflies ate approximately equal numbers of the two mayfly species. This pattern of dietary composition was the result of a reduction of attack frequency on the slower swimming E. altana with predator satiation and a continued high attack rate on B. tricaudatus regardless of recent feeding history. In contrast, fasted H. pacifica fed fresh frozen mayflies ate more B. tricaudatus indicating the importance of differences in prey mobility in controlling dietary composition.The high degree of similarity in patterns of feeding and mechanisms underlying those patterns for H. pacifica and M. signata suggest that they may be using similar rules for choosing mayfly prey and we suggest that mayfly prey are ranked by stoneflies on the basis of handling times. A general mechanistic model for stoneflies feeding on mayflies is presented and discussed.     

Cascading effects of predatory fish on the composition of benthic algae in high‐altitude streams

Cascading effects of predators can affect ecosystem properties by changing plant biomass, distribution and assemblage composition. Using data from field surveys and whole‐stream experiments we tested the hypothesis that predatory trout change assemblage composition of benthic algae in high‐elevation streams mediated by grazer behavior. Field surveys revealed that the taxonomic composition of algal assemblages differed significantly between streams that contained trout and those that were fishless; but comparisons of palatable versus unpalatable algal taxa between fish and fishless streams were equivocal because of high natural variability. Therefore, we tested for a behavioral (non‐consumptive) trophic cascade experimentally by adding brook trout chemical cues to six naturally fishless streams for 25 days and compared responses of grazers and algae to six reference streams without fish cues added. Algal response variables included rates of change in the abundance of three physiognomic categories, from most palatable (attached erect and prostrate diatoms) to least palatable (non‐diatoms), as determined from food selectivity analyses of the most common grazers (mayflies and caddisflies). Fish cues did not affect the mean densities or changes in densities of total grazers or any individual grazer species. However, in streams where fish cues were added, rates of accrual of attached erect diatoms, which was the preferred algal type for the grazer most vulnerable to trout predation (Baetis), were higher and their densities increased significantly faster with increasing densities of this grazer species than in reference streams. Results of his experiment support the hypothesis that predator induced suppression of grazer foraging behavior, rather than cascading effects of top predators on grazer density, may contribute to variation in the composition of algal assemblages among streams by allowing proliferation of most palatable algal species.     

Are populations of mayflies living in adjacent fish and fishless streams genetically differentiated?

1. Conspecific populations living in habitats with different risks of predation often show phenotypic variation in defensive traits. Traits of two species of mayflies (Baetidae: Baetis bicaudatus and Baetis sp. nov.) differ between populations living in fish and fishless streams in a high altitude drainage basin in western Colorado, U.S.A. We tested for genetic differentiation between mayfly populations in these two habitat types, assuming that lack of genetic differentiation would be consistent with the hypothesis that those traits are phenotypically plastic. 2. Previous work has shown that larvae of both species behave differently and undergo different developmental pathways in adjacent fish and fishless streams. These phenotypic differences in behaviour and development have been induced experimentally, suggesting that populations from fishless streams have the genetic capability to respond to fish. 3. During summer 2001 we collected Baetis larvae from several fish and fishless streams, and from fish and fishless sections of the same streams. We used allozymes and a fragment of the cytochrome oxidase subunit 1 mitochondrial gene to examine genetic variation of Baetis individuals within and among streams. 4. Results showed that genetic variation exists among populations of the same species of Baetis from different streams, but none of that variation was associated with the presence or absence of fish. These data confirm that populations of Baetis living in fish and fishless streams are not genetically distinct, and are consistent with the hypothesis that traits associated with environments of different risk are phenotypically plastic.     

Trout reverse the effect of water temperature on the foraging of a mayfly

Climate change is likely to increase the metabolisms of ectothermic animals living below their thermal optimum. While ectothermic top predators may compensate by increasing foraging, ectothermic prey may be unable to increase foraging because of increased predation risk from ectothermic predators. We examined how the diurnal drift behavior (i.e., the downstream movement associated with foraging) of the mayfly Baetis, an ectothermic herbivore, responds to changing temperature in the implied presence and absence of trout, an ectothermic predator. In an experiment replicated at the catchment scale, water temperature and trout presence strongly interacted to affect the diurnal drift of Baetis from artificial channels lacking periphyton over a water temperature range of 4.2–14.8 °C. In fishless streams, daytime drift increased with increasing water temperature, likely because of increased metabolic demand for food. However, in trout-bearing streams, daytime drift decreased with increasing water temperature. Our interpretation is that the perceived threat of trout rose with increasing water temperature, causing mayflies to reduce foraging despite heightened metabolic demand. These results suggest that anticipated increases in stream temperature due to climate change may further escalate divergence in structure and process between fishless and trout-bearing streams. Similar dynamics may occur in other ecosystems with ectothermic predators and prey living below their thermal optima.     

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.     

Unidirectional prey-predator facilitation: apparent prey enhance predators' foraging success on cryptic prey

Food availability can strongly affect predator-prey dynamics. When change in habitat condition reduces the availability of one prey type, predators often search for other prey, perhaps in a different habitat. Interactions between behavioural and morphological traits of different prey may influence foraging success of visual predators through trait-mediated indirect interactions (TMIIs), such as prey activity and body coloration. We tested the hypothesis that foraging success of stream-dwelling cutthroat trout (Onchorhyncus clarki) on cryptically coloured, less-active benthic prey (larval mayfly; Paraleptophebia sp.) can be enhanced by the presence of distinctly coloured, active prey (larval stonefly shredder; Despaxia augusta). Cutthroat trout preyed on benthic insects when drifting invertebrates were unavailable. When stonefly larvae were present, the trout ate most of the stoneflies and also consumed a higher proportion of mayflies than under mayfly only treatment. The putative mechanism is that active stonefly larvae supplied visual cues to the predator that alerted trout to the mayfly larvae. Foraging success of visual predators on cryptic prey can be enhanced by distinctly coloured, active benthic taxa through unidirectional facilitation to the predators, which is a functional change of interspecific interaction caused by a third species. This study suggests that prey-predator facilitation through TMIIs can modify species interactions, affecting community dynamics.