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     

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.     

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.     

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.     

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.     

Effects of temperature and a piscivorous fish on diel winter behaviour of juvenile brown trout (Salmo trutta)

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Diel,size-differential drift patterns of three macroinvertebrate species in the lower Mississippi River,Louisiana (USA)

We explored macroinvertebrate size-differential drift in the lower Mississippi River (a 9th order system). Because this river system is highly turbid, we hypothesized that visually-dependent vertebrate predators feeding on drifting organisms would be at a disadvantage. Thus, size-differential drift should not occur. For one 24-hour period in both January and April, six drift nets were used to sample surface drift. Nets were emptied once every four hours. Individual intra-ocular distances of three macroinvertebrate species (Hydropsyche orris: Trichoptera, Hexagenia limbata: Ephemeroptera, Macrobrachium ohione: Crustacea) were measured. Percentages of size classes in the drift were determined. In both months, large individuals of H. orris and H. limbata were prevalent in the nocturnal but scarce in the diurnal drift. In January, large M. ohione drifted regardless of time. In April, large M. ohione predominated the nocturnal drift. Our results could not be attributed solely to vertebrate predator avoidance. Other mechanisms such as diel microhabitat migration and current velocity may have accounted for the results.     

Invertebrate drift in the regulated River Tees, and an unregulated tributary Maize Beck, below Cow Green dam

SUMMARY. The Cow Green dam was completed in the summer of 1970 and invertebrate drift was sampled below the dam and in an adjacent tributary, Maize Beck, on thirty-one occasions between July 1970 and September 1973. Drift was sampled by pumping river water through a filter. The intake was placed in Maize Beck for the first sample and in the Tees for the second, and so on alternately for the rest of the sampling period. Nets were used on ten occasions, nine of these in winter months and once when the pump broke down. A total of ninety-five taxa were recognized, of which eighty-six occurred in Maize Beck and seventy-one in the Tees. The Tees fauna was dominated numerically and in terms of biomass by a large population of micro-crustaceans originating in the reservoir. Hydra and Naididae also formed a large proportion of the Tees drift but contributed little to the biomass. Ephemeroptera were most abundant in Maize Beck samples. Diptera were abundant in drift catches in both streams with simuliid larvae most numerous in Maize Beck and chironomid larvae most numerous in the Tees. The greatest drift densities of the benthic fauna were observed between April and October; the mean number of organisms per 10 m³ were seventy-three in Maize Beck and 144 in the Tees. The mean densities in winter were very low, respectively two and seventeen per 10 m³ in the two rivers. There was no significant difference between the mean levels of the total bottom fauna (numbers and biomass) in the drift in the two rivers during the period April-October, but vrtnter biomass was significantly greater in the Tees. In July 1970 micro-crustaceans represented 29% (14 per 10 m³) of total drift numbers and 3% (0.7 mg wet-weight per 10 m³) of the biomass, whereas in 1973 they represented 99% of both the numbers (37 670 per 10 m³) and weight (2.2 g wet-weight per 10 m³). The relation between benthos and drift was examined. In the drift Plecoptera and Baetidae were more abundant in Maize Beck than in the Tees. Only Chironomidae and Nais spp. were more abundant in the Tees, In the benthos the density of Plecoptera and Baetidae was not significantly different in the two rivers, but all other groups with the exception of Simuliidae occurred at greater densities in the Tees. The proportion of baetids present in the drift was greatest in Maize Beck. No such difference was demonstrated for total fauna. Diel rhythms were observed in baetids and simuhids with densities greater in night catches. Nocturnal peaks of these organisms were less pronounced in the Tees. Chironomid larvae showed no diel changes in abundance. Significant diel changes in the mean weights of individual animals were not detected in baetid nymphs or chironomids. Micro-crustaceans showed no nocturnal peaks of abundance. Preliminary observations on the quality and quantity of seston caught in drift samples between April and October showed great differences between the rivers. In the Tees the bulk ofeach sample consisted of algal filaments derived from the river and micro-crustaceans from the reservoir. In Maize Beck algae were un-common and the sample was composed of peat and mineral particles. Data are presented on seston output at different discharges.     

Independent and combined effects of multiple predators across ontogeny of a dominant grazer

Ecosystems host multiple coexisting predator species whose interactions may strengthen or weaken top–down control of grazers. Grazer populations often exhibit size‐structure, but the nature of multiple predator effects on suppression of size‐structured prey has seldom been explicitly considered. In a southeastern US salt‐marsh, we used both field (additive design) and mesocosm (additive‐substitutive design) experiments to test the independent and combined effects of two species of predatory crab on the survival and predator‐avoidance behavior (i.e. a non‐consumptive effect) of both juveniles and adults of a dominant grazing snail. Results showed: 1) juvenile snails were more vulnerable to predation; 2) consumptive impacts of predators were hierarchically nested, i.e. the larger predator consumed both juvenile and adult snails, while the smaller‐bodied predator consumed only juvenile snails; 3) there were no emergent multiple predator effects on snail consumption; and 4) non‐consumptive effects differed from consumptive effects, with only the large predator inducing predator‐avoidance behavior of individuals within either snail ontogenetic class. The smaller predator therefore played a functionally redundant trophic role across the prey classes considered, augmenting and potentially stabilizing trophic regulation of juvenile snails. Meanwhile, the larger predator played a complementary and functionally unique role by both expanding the size‐spectrum of prey trophic regulation and non‐consumptively altering prey behavior. While our study suggests that nestedness of consumptive interactions determined by predator and prey body sizes may allow prediction of the functional redundancy of particular predator species, it also shows that traits beyond predator body size (e.g. habitat domain) may be required to predict potentially cascading non‐consumptive effects. Future studies of multiple predators (and predator biodiversity) should continue to strive towards greater realism by incorporating not only size‐structured prey, but also other aspects of resource and environmental heterogeneity typical of natural ecosystems.     

Piscivore addition causes a trophic cascade within and across ecosystem boundaries

The addition of predators can play a key role in structuring ecological communities through both consumptive and non‐consumptive effects. Stocking of piscivorous fish in lakes and similar experimental introductions have provided fundamental evidence in support of trophic cascade theory. Yet, the impact of piscivore addition on cross ecosystem subsidies and meso‐predator resource use has not been well studied. Here, we use a replicated pond experiment to document the trophic impacts of a piscivore, cutthroat trout Onchorhynchus clarkii, on aquatic communities already containing a meso‐predatory fish (threespine stickleback Gasterosteus aculeatus) and neighbouring terrestrial ecosystems. We find that piscivore addition led to a trophic cascade that extended across an ecosystem boundary: trout addition increased the biomass and average size of insects emerging into the terrestrial system. Piscivores caused a diet shift in stickleback, a non‐consumptive effect that was likely mainly responsible for the increase in emerging insect biomass. We additionally show that heterogeneity in the strength of the pelagic trophic cascade was more closely correlated with the magnitude of diet shift (reflecting a non‐consumptive effect) than decreases in stickleback abundance (a consumptive effect). Taken together, our experiment demonstrates that the addition of a piscivore causes a trophic cascade that can extend beyond the aquatic system and suggests that non‐consumptive effects may more strongly influence the strength of a trophic cascade than has been previously recognized.     

Plant resistance attenuates the consumptive and non‐consumptive impacts of predators on prey

Plant resistance and predation have strong independent and interacting effects on herbivore survival, behavior, and patterns of herbivory. Historically, research has emphasized variation in the consumption of herbivores by enemies. Recent work, however, demonstrates that predators also elicit important changes in the traits of their prey, but we do not know how this is influenced by plant quality. In this study, we quantify how the consumptive and non‐consumptive effects of predators vary along a gradient of plant resistance using tomato plants (Solanum lycopersicum), tobacco hornworms (Manduca sexta), and predaceous stinkbugs (Podisus maculiventris). We manipulated resource quality using three tomato lines that vary in the expression of the jasmonate pathway, a phytohormonal pathway that is central in mediating resistance to insects. Resistant plants had higher levels of defensive proteins and glandular trichomes than low resistance plants. The consumptive and non‐consumptive effects of predators were quantified on the three tomato lines by comparing the impact of ‘lethal’ predators that both kill and scare prey with ‘risk’ predators whose mouthparts were surgically impaired to prevent killing. Across several field experiments, the total cascading effect of predators on plant damage was 80.4% lower on jasmonate‐overex‐pressing (highly resistant) plants compared to that on wild‐type or jasmonate‐insensitive (low resistance) plants. This dramatic attenuation of predator effects was due to a 66% reduction in consumption on high resistance plants, and also because of a 65% decline in non‐consumptive effects. Numerous studies in natural and agricultural habitats have documented that predator effects tend to be weaker on well‐defended plants; our results provide novel mechanistic insight into this pattern by demonstrating that plant resistance substantially weakens both the consumptive and non‐consumptive impacts of predators.     

Relationship between the drift of macroinvertebrates and the activity of brown trout in a small stream

Brown trout Salmo trutta were most active in a small stream at night, dusk and dawn when drift rate was highest, but correlations between hourly drift rates and the trout's activity varied substantially between individuals, between different dates for a single individual, and between different periods of the daily cycle. On some occasions, the trout were responsive to the total drift rate, either at night or during the day, and on others to the largest drifting organisms only (terrestrial organisms, adults of Ephemeroptera, Diptera and Trichoptera). The study supports the idea that trout adapt their activity pattern to the abundance of drifting prey, either as generalists towards any organism, or as specialists towards the largest ones.     

The influence of habitat structure and flow permanence on macroinvertebrate assemblages in temporary rivers in northwestern Zimbabwe

Temporary rivers within the Nyaodza-Gachegache subcatchment in northwestern Zimbabwe were investigated to examine the role of flow permanence and habitat structure on macroinvertebrate community composition. Macroinvertebrate communities of intermittent and ephemeral rivers displayed significant differences in the number of taxa, macroinvertebrate abundance, Shannon and Simpson diversity indices and in size class structure. Intermittent sites were characterised by higher numbers of taxa, diversity and Ephemeroptera and Trichoptera richness compared to ephemeral sites. The fauna of ephemeral sites was dominated by a single taxon (Afrobaetodes) (Ephemeroptera, Baetidae) whilst larger sized taxa (e.g. Elassoneuria (Ephemeroptera, Oligoneuriidae), Dicentroptilum (Ephemeroptera, Baetidae), Aethaloptera (Trichoptera, Hydropsychidae), Pseudagrion (Odonata, Coenagrionidae) and Tholymis (Odonata, Libellulidae) were exclusively restricted to intermittent sites. Clear differences were observed between sand, gravel, cobble and vegetation habitats. Vegetation and cobbles supported distinct communities, with some taxa exclusively restricted either to vegetation (e.g. Pseudagrion, Leptocerina (Trichoptera, Leptoceridae), Cloeon (Ephemeroptera, Baetidae), Afronurus (Ephemeroptera, Heptageniidae) and Povilla (Ephemeroptera, Polymitarcidae) or cobble (e.g. Aethaloptera and Dicentroptilum) habitats. In terms of ensuring optimum diversity within the subcatchment, we consider conservation of critical habitats (cobbles and vegetation) and maintenance of natural flows as the appropriate management actions. Handling editor: D. Dudgeon     

Interspecific differences in antipredator strategies determine the strength of non‐consumptive predator effects on stream detritivores

Animal species differ considerably in their response to predation risks. Interspecific variability in prey behaviour and morphology can alter cascading effects of predators on ecosystem structure and functioning. We tested whether species‐specific morphological defenses may affect responses of leaf litter consuming invertebrate prey to sit‐and‐wait predators, the odonate Cordulegaster boltonii larvae, in aquatic food webs. Partly or completely blocking the predator mouthparts (mandibles and/or extensible labium), thus eliminating consumptive (i.e. lethal) predator effects, we created a gradient of predator‐prey interaction intensities (no predator < predator – no attack < predator – non‐lethal attacks < lethal predator). A field experiment was first used to assess both consumptive and non‐consumptive predator effects on leaf litter decomposition and prey abundances. Laboratory microcosms were then used to examine behavioural responses of armored and non‐armored prey to predation risk and their consequences on litter decomposition. Results show that armored and non‐armored prey responded to both acute (predator – non‐lethal attacks) and chronic (predator – no attack) predation risks. Acute predation risk had stronger effects on litter decomposition, prey feeding rate and prey habitat use than predator presence alone (chronic predation risk). Predator presence induced a reduction in feeding activity (i.e. resource consumption) of both prey types but a shift to predator‐free habitat patches in non‐armored detritivores only. Non‐consumptive predator effects on prey subsequently decreased litter decomposition rate. Species‐specific prey morphological defenses and behaviour should thus be considered when studying non‐consumptive predator effects on prey community structure and ecosystem functioning.     

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.     

Artificial light as a disturbance to light‐naïve streams

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Predator‐Naïve Brown Trout (Salmo trutta) Show Antipredator Behaviours to Scent from an Introduced Piscivorous Mammalian Predator Fed Conspecifics

Introduced mammalian predators may pose a high risk for native and naïve prey populations, but little is known about how native fish species may recognize and respond to scents from introduced mammalian predators. We investigated the role of diet‐released chemical cues in facilitating predator recognition, hypothesizing that native brown trout (Salmo trutta) would exhibit antipredator behaviours to faeces scents from the introduced American mink (Neovision vison) fed conspecifics, but not to non‐trout diets. In treatments‐control and replicate stream tank experiments, brown trout showed significant antipredator responses to faeces scent from mink fed conspecifics, but not to faeces scent from mink fed a non‐trout diet (chicken), or the non‐predator food control, Eurasian beaver (Castor fiber). We conclude that native and naïve brown trout show relevant antipredator behaviours to an introduced mammalian predator, presumably based on diet‐released conspecific alarm cues and thereby estimate the predation risk.     

The Diet of the Brown Trout Salmo trutta (L.) during the Reproductive Period: Size‐Related and Sexual Effects

We investigated the autumnal diet of the brown trout Salmo trutta, in a Prepyrenean stream (NW Iberian Peninsula) focusing on intraspecific dietary differences related to size and sex. The diet of trout included 18 types of prey, with Plecoptera and Ephemeroptera nymphs and Diptera larvae as the most consumed taxa. Large trout ate larger prey, than did small trout, and also increased the consumption of terrestrial‐surface prey with respect to aquatic‐benthic prey. As terrestrial‐surface preys were larger than aquatic‐benthic prey, the size‐related differences in the diet of trout were related to gape‐limitations. Although male and female trout did not differ in size, we found that males foraged on a more diverse type of prey than females, probably owing to male territoriality during the reproductive period. This study provides new evidence of dietary plasticity in the brown trout and confirms the importance of local dietary studies to better understand factors which drive trophic ecology of predators. (© 2006 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Night — day drift patterns and the size of larvae of two aquatic insects

A test of Allan's (1978) hypothesis about differential drift abundance of mayflies of the genus Baetis between night and day, and the size of larvae was performed at a mountain stream in Idaho. Palisades Creek, Idaho, contains a different species of mayfly, B. tricaudatus, and vertebrate predator, Salmo clarki, than Cement Creek, Colorado (B. bicaudatus, and brook trout, Salvelinus fontinalis). Consequently it was not known if B. tricaudatus would exhibit a similar pattern as its congener in Cement Creek, with large instars tending to avoid daylight drift, as found by Allan (1978). However, similar results were observed in the present study. It appears that the earlier hypothesis may have generality for geographically distinct streams with a different vertebrate predator and mayfly prey. The existence of a similar pattern for chironomid larvae was also tested, however, no such pattern existed. This discrepency between taxa may be due to differential predation, or to inherent differences in drift abilities.