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.     

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     

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.     

Fitness and community consequences of avoiding multiple predators

We investigated the fitness and community consequences of behavioural interactions with multiple predators in a four-trophic-level system. We conducted an experiment in oval flow-through artificial-stream tanks to examine the single and interactive sublethal effects of brook trout and stoneflies on the size at emergence of Baetis bicaudatus (Ephemeroptera: Baetidae), and the cascading trophic effects on algal biomass, the food resource of the mayflies. No predation was allowed in the experiment, so that all effects were mediated through predator modifications of prey behaviour. We reared trout stream Baetis larvae from just before egg development until emergence in tanks with four treatments: (1) water from a holding tank with two brook trout (trout odour), (2) no trout odour + eight stoneflies with glued mouthparts, (3) trout odour + stoneflies and (4) no trout odour or stoneflies. We ended the experiment after 3 weeks when ten male and ten female subimagos had emerged from each tank, measured the size of ten male and ten female mature nymphs (with black wing pads), and collected algal samples from rocks at six locations in each tank. To determine the mechanism responsible for sublethal and cascading effects on lower trophic levels we made day and night observations of mayfly behaviour for the first 6 days by counting mayflies drifting in the water column and visible on natural substrata in the artificial streams. Trout odour and stoneflies similarly reduced the size of male and female Baetis emerging from artificial streams, with non-additive effects of both predators. While smaller females are less fecund, a fitness cost of small male size has not been determined. The mechanism causing sublethal effects on Baetis differed between predators. While trout stream Baetis retained their nocturnal periodicity in all treatments, stoneflies increased drift dispersal of mayflies at night, and trout suppressed night-time feeding and drift of mayflies. Stoneflies had less effect on Baetis behaviour when fish odour was present. Thus, we attribute the non-additivity of effects of fish and stoneflies on mayfly growth to an interaction modification whereby trout odour reduced the impact of stoneflies on Baetis behaviour. Since stonefly activity was also reduced in the presence of fish odour, this modification may be attributed to the effect of fish odour on stonefly behaviour. Only stoneflies delayed Baetis emergence, suggesting that stoneflies had a greater sublethal effect on Baetis fitness than did trout. Delayed emergence may reduce Baetis fitness by increasing risks of predation and parasitism on larvae, and increasing competition for mates or oviposition sites among adults. Finally, algal biomass was higher in tanks with both predators than in the other three treatments. These data implicate a behavioural trophic cascade because predators were not allowed to consume prey. Therefore, differences in algal biomass were attributed to predator-induced changes in mayfly behaviour. Our study demonstrates the importance of considering multiple predators when measuring direct sublethal effects of predators on prey fitness and indirect effects on lower trophic levels. Identification of an interaction modification illustrates the value of obtaining detailed information on behavioural mechanisms as an aid to understanding the complex interactions occurring among components of ecological communities. Received: 20 March 1997 / Accepted: 29 September 1997     

Nocturnal drift of mayfly nymphs as a post-contact antipredator mechanism

1. The predominantly nocturnal constrained drift of stream invertebrates is commonly regarded as a behaviour that avoids encounters with visually foraging fish in the water column. The alternative explanation, that drift peaks are caused by bottom-feeding, nocturnal predators, has rarely been tested.
2. We examined these hypotheses by collecting invertebrate drift in five streams in northern Finland: one with brown trout ( Salmo trutta , a drift-feeding fish), one with alpine bullhead ( Cottus poecilopus , a benthic fish), one with both species, and two fishless streams.
3. Drift by Baetis mayflies was aperiodic or slightly diurnal in both fishless streams on all sampling occasions. In contrast, drift was nocturnal in streams with trout and, to a lesser extent, in the stream with bullhead. Non-dipteran prey drifted mainly nocturnally in all streams with fish, whereas Diptera larvae were less responsive to the presence of fish.
4. In laboratory experiments, bullheads were night-active, causing a much higher frequency of drift by touching Baetis at night than during the day. Thus, increased nocturnal drift may serve to avoid both visual predators (a pre-contact response) and benthic fish (a post-contact response). In streams with bottom-feeding fish, nocturnal drift should be caused by increased drift by night rather than by reduced drift by day.     

Effects of trout on the diel periodicity of drifting in baetid mayflies

Some benthic invertebrates in streams make frequent, short journeys downstream in the water column (=drifting). In most streams there are larger numbers of invertebrates in the drift at night than during the day. We tested the hypothesis that nocturnal drifting is a response to avoid predation from fish that feed in the water column during the day. We surveyed diel patterns of drifting by nymphs of the mayfly Baetis coelestis in several streams containing (n=5) and lacking (n=7) populations of rainbow trout, Oncorhynchus mykiss. Drifting was more nocturnal in the presence of trout (85% of daily drift occurred at night) than in their absence (50% of daily drift occurred at night). This shift in periodicity is due to reduced daytime drifting in streams with trout, because at a given nighttime drift density, the daytime drift density of B. coelestis was lower in streams occupied by trout than in troutless streams. Large size classes of B. coelestis were underrepresented in the daytime drift in trout streams compared to nighttime drift in trout streams, and to both day and night drift in troutless streams. Differences in daytime drift density between streams with and without trout were the result of differences in mayfly drift behaviour among streams because predation rates by trout were too low to significantly reduce densities of drifting B. coelestis. We tested for rapid (over 3 days) phenotypic responses to trout presence by adding trout in cages to three of the troutless streams. Nighttime drifting was unaffected by the addition of trout, but daytime drift densities were reduced by 28% below cages containing trout relative to control cages (lacking trout) placed upstream. Drift responses were measured 15 m downstream of the cages suggesting that mayflies detected trout using chemical cues. Overall, these data support the hypothesis that infrequent daytime drifting is an avoidance response to fish that feed in the water column during the day. Avoidance is more pronounced in large individuals and is, at least partially, a phenotypic response mediated by chemical cues.     

Interactions between fish,grazing invertebrates and algae in a New Zealand stream: a trophic cascade mediated by fish-induced changes to grazer behaviour?

Experiments in laboratory stream channels compared the behaviour of Deleatidium mayfly nymphs in the absence of fish with that in the presence of either native common river galaxias (Galaxias vulgaris Stokell) or introduced brown trout (Salmo trutta L.). Galaxias present similar predation risks to prey during day and night but are more active at night. Whereas, trout present a higher predation risk during the day. Deleatidium maintained a fixed nocturnal drift periodicity that is characteristic of streams containing visually feeding fish regardless of the nature of the predation regime presented in the laboratory. However, the number on the substratum surface, and therefore able to graze algae, was lower when fish were present than when they were absent. The number was lower during the day in the presence of trout, when they present the highest predation risk, and lower during the night compared to the day in trials with galaxias when galaxias activity disturbs Deleatidium from the substratum. Increases in the probability of Deleatidium leaving a patch, reductions in the proportion of mayflies on high quality patches and reductions in the distance travelled from refuge also reflected variations in the predation regime. Similar differences in positioning were observed under the same predation regimes in in situ channels in the Shag River and these were associated with differences in algal biomass. Algal ash-free dry mass (AFDM) and chlorophyll a (chl a) were higher on the tops of cobbles when fish were present. Fish also affected the biomass and the distribution of algae on cobbles as AFDM and chl a were higher on the sides of cobbles from channels with trout compared to those with galaxias. Changes in grazing behaviour, caused by predator avoidance, are likely to have been responsible for differences in algal biomass because no significant differences were detected between treatments in the biomass of Deleatidium or of total invertebrates.     

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.     

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 influence of upstream predation on the expression of fish effects in downstream patches

1. Enclosures were installed in a fishless stream and divided transversely into upstream and downstream sections. Downstream sections were further divided longitudinally, and one of the downstream sections in each enclosure was stocked with juvenile coho salmon ( Oncorhynchus kisutch ), and the other side was left as a fishless control. Densities of juvenile coho were then manipulated in upper sections of enclosures to determine the effect of upstream predation intensity on fish predation effects in downstream sections.
2. Significantly fewer mayflies (primarily Ameletus sp.) were observed grazing on unglazed ceramic tiles in lower enclosure sections with fish present. There was no detectable effect of fish density in upstream enclosure sections on the number of mayflies observed grazing on ceramic tiles in lower sections of the enclosures.
3. There was a significant positive effect of both fish presence and upstream density on chlorophyll a concentrations on ceramic tiles in lower enclosure sections, but not on chlorophyll a on natural gravel substratum.
4. Behavioural experiments with mayflies and coho in streamside troughs suggest that Ameletus sp. responds primarily to mechanical rather than chemical cues from coho parr.     

Assessing the influence of water and substratum quality on benthic macroinvertebrate communities in a metal-polluted stream: an experimental approach

SUMMARY 1. Field and laboratory experiments were conducted to assess the relative influence of water quality and substratum quality on benthic macroinvertebrate communities in the Animas River, a metal-polluted stream in south-western Colorado (U.S.A.).
2. A community-level in situ toxicity test measured direct effects of Animas River water on benthic invertebrates collected from a reference stream (Elk Creek). The effects of metal-contaminated biofilm were examined by comparing macroinvertebrate colonisation of clean and contaminated substrata placed in Elk Creek. A feeding experiment with the mayfly Baetis tricaudatus Dodds (Ephemeroptera: Baetidae) examined metal bioaccumulation and effects of metal-contaminated biofilm on growth and survival.
3. Animas River water was acutely toxic to most taxa, with greatest effects observed on mayflies (Heptageniidae, Ephemerellidae) and stoneflies (Taeniopterygidae and Capniidae).
4. Although Animas River biofilm was characterised by high concentrations of metals and low algal biomass, most taxa colonised substratum from the reference stream and the Animas River equally. The exceptions were Ephemerellidae, Taeniopterygidae and Simuliidae, which were less abundant on Animas River substratum. Mayflies grazing Animas River biofilm accumulated significantly more metals and showed reduced growth compared with organisms feeding on Elk Creek biofilm.
5. Results of our experiments demonstrated that effects of heavy metals on benthic community structure in the Animas River were complex, and that responses to metals in water and contaminated substratum were species-specific. Predicting recovery of benthic communities following remediation requires an understanding of these species-specific responses.     

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.     

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.     

Facilitation and interference among three predators affect their consumption of a stream-dwelling mayfly

1. We experimentally tested if a multiplicative risk model accurately predicted the consumption of a common mayfly at risk of predation from three predator species in New Zealand streams. Deviations between model predictions and experimental observations were interpreted as indicators of ecologically important interactions between predators. 2. The predators included a drift‐feeding fish [brown trout (T), Salmo trutta], a benthivorous fish [galaxiid (G), koaro, Galaxias brevipennis] and a benthic predatory stonefly (S; Stenoperla sp.) with Deleatidium sp. mayflies as prey. Eight treatments with all predator species combinations and a predator‐free control were used. Experiments were performed in aquaria with cobbles as predator refuges for mayflies and we measured the proportion of prey consumed after 6 h for both day and night trials. 3. Trout consumed a higher proportion of prey than other predators. For the two predator treatments we found less than expected prey consumption in the galaxiid + trout treatment (G + T) for both day and night trials, whereas a higher than expected proportion of prey was consumed during night time in the stonefly + trout (S + T) treatment. 4. The results indicate interference (G + T) and facilitation (S + T) between predators depending on predator identity and time of day. Thus, to make accurate predictions of interspecific interactions, it is necessary to consider the ecology of individual species and how differences influence the direction and magnitude of interactions.     

Drift dispersal of mayfly nymphs in the presence of chemical and visual cues from diurnal drift- and nocturnal benthic-foraging fishes

1. The drift of Baetis thermicus nymphs in the presence of chemical, visual and hydrodynamic cues, considered individually and in combination, produced by different predatory fishes was examined experimentally in laboratory streams. Masu salmon ( Oncorhynchus masou ) and freshwater sculpin ( Cottus nozawae ) are typical drift- and benthic-foraging fishes, respectively.
2. Observations of fish swimming in the streams revealed differing diel periodicity between the species; sculpin were nocturnal foragers and salmon diurnal.
3. The drift rate of Baetis by night increased in the presence of chemical cues from sculpin, with other cues having no interactive effects. In contrast, the drift rate increased primarily in the presence of both chemical and, particularly, visual cues from salmon, although no additional effects were found for any non-visual cues. Visual cues could enable Baetis to assess precisely the predation risk from foraging salmon by day, whereas Baetis could not use visual cues to detect sculpin either at night, because of the low light intensity, or during the day, because of the low activity of sculpin at that time.
4. In natural streams, which are often inhabited by several predatory fish employing different modes of foraging, invertebrates may be able to precisely assess the risk and effectively to avoid predators by using cues unique to each.     

Reduction of predation risk under the cover of darkness: Avoidance responses of mayfly larvae to a benthic fish

Summary Mayfly larvae of Paraleptophlebia heteronea (McDunnough) had two antipredator responses to a nocturnal fish predator (Rhinichthys cataractae (Valenciennes)): flight into the drift and retreat into interstitial crevices. Drift rates of Paraleptophlebia abruptly increased by 30 fold when fish were actively foraging in the laboratory streams but, even before fish were removed, drift began returning to control levels because larvae settled to the substrate and moved to areas of low risk beneath stones. This drifting response was used as an immediate escape behavior which likely decreases risk of capture from predators which forage actively at night. Surprisingly, drift most often occurred before contact between predator and prey, and we suggest that in darkness this mayfly may use hydrodynamic pressure waves for predator detection, rather than chemical cues, since fish forage in an upstream direction. Although drifting may represent a cost to mayfly larvae in terms of relocation to a new foraging area with unknown food resources, the immediate mortality risk probably out-weighs the importance of staying within a profitable food patch because larvae can survive starvation for at least 2 d. In addition to drifting, mayflies retreated from upper, exposed substrate surfaces to concealed interstitial crevices immediately after a predator encounter, or subsequent to resettlement on the substrate after predator-induced drift. A latency period was associated with this response and mayflies remained in these concealed locations for at least 3 h after dace foraging ceased. Because this mayfly feeds at night and food levels are significantly lower in field refugia under stones, relative to exposed stone surfaces, predator avoidance activity may limit foraging time and, ultimately, reduce the food intake of this stream mayfly.     

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.     

Incorporating invertebrate predators into theory regarding the timing of invertebrate drift

Theory concerning the timing of lotic invertebrate drift suggests that daytime-feeding fish cause invertebrates to restrict their drift behavior to the nighttime. However, there is growing evidence that the nighttime foraging of invertebrate predators also contributes to the nocturnal timing of drift, though it is unclear whether the nocturnal behavior of invertebrate predators is innate or proximately caused by fish. In two experiments, one conducted in a fish-bearing stream and a second in a fishless stream, we compared the drift patterns of Baetidae (Ephemeroptera) from channels with and without benthic invertebrate predators. We tested whether invertebrate predators affect the timing of drift, either as a proximate cause of nocturnal drift in the fishless stream (diel periodicity) or as a proximate cause of a pre-dawn peak in drift in the fish-bearing stream (nocturnal periodicity). In the fish-bearing stream experiment, a pre-dawn increase of baetid drift occurred independently of invertebrate predators, indicating that invertebrate predators were not the proximate cause of nocturnal periodicity in the stream. In the fishless stream experiment, invertebrate predators caused more baetid drift at night than during the day, indicating that invertebrate predators caused the nocturnal drift pattern we observed in the stream, and that invertebrate predators can influence drift timing independently of fish. Therefore, we suggest that both visually feeding fish and nocturnally foraging benthic predators, when present, affect the timing of invertebrate drift; visually feeding fish by reducing daytime drift, and benthic predators by increasing nighttime drift.     

Benthivorous fish reduce stream invertebrate drift in a large-scale field experiment

Drift as a low-energy cost means of migration may enable stream invertebrates to leave risky habitats or to escape after encountering a predator. While the control of the diurnal patterns of invertebrate drift activity by fish predators has received considerable interest, it remains unclear whether benthivorous fish reduce or increase drift activity. We performed a large-scale field experiment in a second-order stream to test if invertebrate drift was controlled by two benthivorous fish species (gudgeon Gobio gobio and stone loach Barbatula barbatula). An almost fishless reference reach was compared with a reach stocked with gudgeon and loach, and density and structure of the invertebrate communities in the benthos and in the drift were quantified in both reaches. The presence of gudgeon and stone loach reduced the nocturnal drift of larvae of the mayfly Baetis rhodani significantly, in contrast to the findings of most previous studies that fish predators induced higher night-time drift. Both drift density and relative drift activity of B. rhodani were lower at the fish reach during the study period that spanned 3 years. Total invertebrate drift was not reduced, by contrast, possibly due to differences in vulnerability to predation or mobility between the common invertebrate taxa. For instance, Chironomidae only showed a slight reduction in drift activity at the fish reach, and Oligochaeta showed no reduction at all. Although benthic community composition was similar at both reaches, drift composition differed significantly between reaches, implying that these differences were caused by behavioural changes of the invertebrates rather than by preferential fish consumption. The direction and intensity of changes in the drift activity of stream invertebrates in response to the presence of benthivorous fish may depend on the extent to which invertebrate taxa can control their drifting behaviour (i.e. active versus passive drift). We conclude that invertebrate drift is not always a mechanism of active escape from fish predators in natural streams, especially when benthos-feeding fish are present.     

Diel vertical movements by lotic mayfly nymphs under variable predation risk

1. The diel foraging periodicities of two grazing mayfly (Ephemeroptera) nymphs, Heptagenia dalecarlica and Baetis rhodani, under variable fish (European minnow) predation risk were examined in a series of laboratory experiments. 2. Heptagenia dalecarlica were almost exclusively nocturnal in their use of feeding areas on stone tops. There was a sharp increase in the proportion of nymphs out of refuge at nightfall, both in the control and fish-odour treatments. In a treatment with freely moving fish, H. dalecarlica decreased their activity during both the day and night. In feeding trials with three freely foraging minnows, nymphs were completely safe when they had access to refuges beneath stones, whereas almost half the nymphs were consumed when no refuges were available. 3. Baetis rhodani nymphs reduced their use of stone tops when exposed to four caged minnows or a freely moving fish, but this occurred both day and night. In feeding trials, B. rhodani were captured only while in the water column, and their mortality risk was independent of refuge availability. 4. It is suggested that because H. dalecarlica lack efficient post-encounter defences, they must rely on pre-encounter mechanisms to reduce the threat of fish predation. It appears that in order to dwell sympatrically with fish, H. dalecarlica have evolved a coexistence by hiding strategy. In contrast, Baetis are vulnerable to fish attacks only if they enter drift in daylight, and are thus able to remain on stone tops both during the day and at night.