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 (Salmotrutta, a drift‐feeding fish), one with alpine bullhead (Cottuspoecilopus, 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.
JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact support@jstor.org.. British Ecological Society is collaborating with JSTOR to digitize, preserve and extend access to Journal of Animal Ecology. Summary 1. Prey preference is determined by active predator choice and by the relative vulnerability of prey taxa. In this study, we addressed the mechanisms of prey preference in the perlodid stonefly Diura bicaudata. 2. Components of the predator-prey interaction between the stonefly and its prey were quantified in laboratory observations. These data were compared to prey selection in preference trials and to gut contents of field-collected stoneflies. Experiments were conducted in spring (May) and in autumn (September), using prey taxa commonly available in each season. 3. In the September trials, Diura exhibited positive selection for black fly larvae, whereas Heptagenia, Ephemerella and large Baetis mayflies were avoided. Encounter rates did not affect preference: these were highest for heptageniids and lowest for black flies. Once contacted, black flies were practically always attacked with a high capture probability. Attack propensity and capture success were very low for all other prey types, including Baetis mayflies. 4. In May, female Diura avoided Ephemerella mayflies and Asellus isopods, but showed a positive, albeit non-significant, preference for Nemoura stoneflies. Males did not select any of the prey types. Again, encounter rate was the least important determinant of preference: nemourid stoneflies were encountered less frequently than other prey, especially by female Diura. Females attacked Asellus more frequently than other prey types. Baetis was not a preferred prey for either of the sexes. 5. Our results show that D. bicaudata prefers sedentary or slowly moving prey types. Preference was determined both by active predator choice and differential prey vulnerability. We suggest that although mobile prey such as Baetis are encountered frequently, they are difficult to capture, and are thus relatively safe from stonefly predation when sedentary prey are also available. 6. Microhabitat overlap between predator and prey may determine encounter rates in the field, but this may not translate into prey preferences. Prey with efficient antipredatory behaviours can risk predator encounters, whereas prey with less efficient escape mechanisms may have to select microhabitats avoided by the predator. It is thus essential that laboratory systems incorporate at least some of the structural complexity of natural streams. However, even relatively simple laboratory systems may provide the complexity needed, as long as they contain prey refuges.
Summary1. We examined direct and indirect behavioural responses of grazing may¯y nymphs (Baetis rhodani) to predation risk from a drift-feeding ®sh (European minnow; odour manipulated), and two types of invertebrate predators, Diura bicaudata (stone¯y) and Rhyacophila nubila (caddis larva). We also assessed the direct responses of the invertebrate predators to ®sh chemicals. 2. Based on diel gut content periodicity, D. bicaudata nymphs were strongly nocturnal foragers. R. nubila was also nocturnal, but only in a ®sh stream; in a stream lacking ®sh, their gut contents did not vary signi®cantly on a diel basis. In the laboratory, Diura was nocturnal even in the absence of ®sh and almost ceased moving when ®sh was present. Rhyacophila shifted from aperiodic to nocturnal foraging in the presence of ®sh. 3. The contrasting behaviours of the two predators may be explained by their respective ®eld distributions: D. bicaudata always co-occurs with ®sh, whereas R. nubila is also found in streams without ®sh. Therefore, a capacity for plastic anti®sh responses is bene®cial for R. nubila, but not for D. bicaudata. 4. Drift of large Baetis nymphs was aperiodic when Rhyacophila was present, but nocturnal in all other treatments. Drift rate was highest when both Diura and ®sh were present and lowest in treatments with Rhyacophila. Predatory stone¯ies, but not the caddis larva, induced a night-time peak in the drift of both Baetis sizeclasses. In the absence of predators, small Baetis drifted aperiodically. Interaction terms in three-way ANOVAs testing for the indirect eects of ®sh and invertebrate predators on may¯y drift periodicity and drift rate were all non-signi®cant, indicating that the response of Baetis to one type of predator was not modi®ed by the other predator. 5. Our results indicate that the drift periodicity of lotic may¯ies may be ®ne-tuned to variations in the multi-predator environment and that prey responses are sizespeci®c. We conclude that the eects of invertebrate predators on prey behaviour vary in relation to predator's foraging strategy and generalizations based on studies with only one type of predator should be avoided.
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.
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