Ontogenetic dietary changes of whitefish larvae: insights from field and experimental observations

Ontogenetic changes in resource use are widespread in many fish species. This study investigated the feeding habits of whitefish (C. lavaretus L.) larvae in Lake Annecy (France) coupled with experimental behavioral studies in order to identify the underlying mechanisms of the ontogenetic shifts in the diet. The predatory behavior of wild larvae, and the escape responses of their zooplankton prey were both videorecorded in experimental tanks under controlled laboratory conditions. Ontogenetic diet patterns showed that young whitefish larvae have a preference for small cyclops, while older larvae selectively predate cladocerans. Our experimental observations showed that the capture success rate also varied in relation to ontogenetic development in fish. Young larvae were more successful in capturing small copepods, whereas old larvae were more successful in capturing Daphnia. In addition, the larvae were able to adjust their predatory behavior (speed, pursuit) according to the swimming pattern of the prey. These observations suggest that the selective predation on cladocerans observed in old larvae is the outcome of both active and passive choices depending on the escape swimming behavior of the prey, and handling time of the predator.     

Differential susceptibility of marine invertebrate larvae: Laboratory predation of sand dollar, Dendrasterexcentricus (Eschscholtz), embryos and larvae by zoeae of the red crab, Cancer productus Randall

Predation on eggs, embryos, and larvae of the sand dollar, Dendraster excentricus (Eschscholtz) was investigated in a series of laboratory feeding experiments. Dendraster susceptibility to predation by zoea larvae of the red crab, Cancer productus Randall was strongly dependent on developmental stage and ontogenetic differences in motility. Clearance rates by C. productus were highest for eggs and averaged 0.551·zoea⁻¹·day⁻¹. Embryos and prism larvae of Dendraster were consumed at an intermediate rate, while pluteus larvae were captured at a relatively low rate. Clearance rates decreased from 0.18 to 0.031·zoea⁻¹·day⁻¹ during the transition of prism larvae into echinoplutei. Differences in Dendraster susceptibility to predation cannot be attributed to increasing prey body size because dwarf plutei were captured at the same rate as normal plutei. Reduced capture rates by Cancer productus zoeae are dependent on the development of Dendraster swimming behavior. Periodic reversals in the direction of ciliary beating and backwards swimming effectively remove Dendraster plutei from the immediate capture sphere of Cancer productus. Reversed swimming appears to function as a post-contact encounter response that reduces the mortality rates of Dendraster plutei.     

Feeding by marine fish larvae: developmental and functional responses

Synopsis The relationship between prey consumption rate and prey concentration (functional response), and its change with growth (developmental response) were examined in the laboratory for three species of marine fish larvae: bay anchovy Anchoa mitchilli (Engraulidae), sea bream Archosargus rhomboidalis (Sparidae) and lined sole Achirus lineatus (Soleidae). The major objective was to determine relative predatory abilities of the larvae by fitting feeding rate data to developmental and functional response models. Feeding success, prey capture success, attack rates, handling times and search rates were estimated. Prey consumption rates and attack rates of bay anchovy usually were highest, but at the lowest prey level (50 per liter) first-feeding sea bream larvae had the highest consumption rate. Sea bream could consume prey at near-maximum rates at prey levels lower than those required by the other species. As larvae grew, time searching per attack decreased rapidly for all species, especially at low prey levels. Handling time also decreased, but most rapidly for bay anchovy. Search rates were highest for bay anchovy and lowest for lined sole. Bay anchovy had the best apparent predation ability, but when previous results on larval growth rates, survival rates and growth efficiencies were considered, sea bream larvae were the most efficient predators and the least likely of the three species to be limited by low prey levels.     

Impact of predation on early stages of the armoured catfish Hoplosternum thoracatum (Siluriformes-Callichthyidae) and implications for the syntopic occurrence with other related catfishes in a neotropical multi-predator swamp

This study investigated the role of predators in preventing competitive exclusion among three closely related armoured catfishes (Callichthys callichthys, Hoplosternum littorale and H. thoracatum) that occur synthopically in multi-predator freshwater swamps of Suriname, South America. The potential impact of predation on armoured catfish was determined by combining laboratory measurements of predation rates on five early developmental stages of the armoured catfish H. thoracatum for 24 aquatic predators with field studies of the density of the predators in the swamps. The contribution of a particular predator to the total predation pressure on its prey was determined to a large extent by the density of the predator in the swamp. Seemingly innocuous predators with low or moderate predation rates in the laboratory may be extremely important in the swamps due to their high abundance. Small-sized omnivorous fishes and aquatic invertebrates were major predators of early developmental stages of armoured catfish. Both qualitative and quantitative ontogenetic changes in the predation pressure on armoured catfish were observed. Major predation on eggs, larvae and juveniles of H. thoracatum resulted from a different set of predators in each developmental stage of the prey. In all developmental stages of H. thoracatum the predation pressure involved several predator species and not a single, dominant predator. The potential predation pressure of the 24 predators taken together and the number of predators that were able to prey on H. thoracatum decreased sharply with increasing age (size) of the prey. Even if egg (nest) predation is prevented by the guarding male, the potential impact of the 24 predators on the populations of armoured catfish is large. Predation may account for the high mortality of H. thoracatum observed in the swamps. The high predation pressure on callichthyid catfishes may help to explain the coexistence of three closely related and morphologically quite similar armoured catfishes in Surinamese swamps.     

Intraspecific variation in Gape–prey size Relationships and Feeding Success During Early Ontogeny in Red Drum, Sciaenops Ocellatus

The relationship between predator gape and prey consumption in laboratory-reared larva and field-caught early juvenile red drum, Sciaenops ocellatus, was investigated in light of the hypothesis that feeding success varies throughout the early life history intervals of marine fishes. We expected the feeding ability of red drum to be more strongly constrained by mouth gape in smaller fish and expected this ability to improve with gape size. To test this hypothesis, field-caught, early juvenile red drum were examined to determine the relationship between gape size and prey size consumed. In field-caught early juveniles, gape (height and width) and prey size consumed (length and width) increased linearly with standard length (SL); however, mean width of prey consumed was only 20–47% of gape width. Furthermore, when regressed on SL, gape width yielded a higher slope than prey width. To further test this hypothesis on less developed, pre-metamorphic fish, age-specific differences in gape, number of prey and size of prey consumed prior to metamorphosis were determined from laboratory-reared red drum larvae. Similar patterns were observed for gape height– and gape width–SL relationships in laboratory-reared red drum larvae. Size of consumed prey increased from three days from hatching (dfh) to 18dfh. The percentage of feeding larvae also increased from 3% at 3dfh to 97% at 18dfh. In both field-caught, early juvenile red drum and laboratory-reared larvae, there was little evidence that the size of prey consumed was constrained by mouth gape. It is hypothesized that besides gape size, the development of other features of the feeding mechanism (e.g., hyoid and opercular series) influences prey-capture performance prior to settlement in marine fishes.     

Effects of prey density, prey instar, and patch size on the development of the predatory mosquito, Toxorhynchites towadensis

Effects of prey density, prey instar, and patch size on the development of the predatory mosquito larva, Toxorhynchites towadensis, were investigated in the laboratory. Survivors of T. towadensis showed different developmental patterns in relation to prey age structure. All predatory larvae in containers with only second instar prey developed into the third instar. However, in several containers with fourth instar prey, mortality of predators was observed. During the third instar, no predatory larva died, but both prey density and prey instar significantly affected the survival of predators during their fourth instar. Large prey size promoted large predator adults, and predatory larvae which grew up in small surface containers responded by developing to large sizes than those in large containers. Larval developmental time of the predators differed in each treatment. During first and second instars, faster predator development was observed in containers with small surface areas and containing young prey individuals. However, when development was enhanced by the presence of old prey individuals, no surface effect was observed. The fastest predator development was observed with prey of mixed instars and high density. This study suggests that a small surface container containing prey of mixed instars and high density is suitable for development of predators.     

Habitat-related predation on juvenile wild-caught and hatchery-reared red drum Sciaenops ocellatus (Linnaeus)

We examined the patterns of habitat-specific mortality for newly settled red drum (Sciaenops ocellatus) using an experimental mesocosm approach. Experiments were designed to analyze prey vulnerability and fish rearing-type (wild-caught or hatchery-reared) in estuarine habitats of varying structural complexity including marsh (Spartina alterniflora Loisel), oyster reef (Crassostrea virginica Gmelin), seagrass (Halodule wrightii Aschers), and nonvegetated sand bottom. We used two different predators, pinfish (Lagodon rhomboides Linnaeus) and spotted seatrout (Cynoscion nebulosus Cuvier). For both predators, vulnerability of wild-caught red drum was significantly lower in structurally complex habitats such as seagrass and oyster reef; the highest vulnerability was associated with the nonvegetated bottom. This habitat effect was not apparent for hatchery-reared prey. In trials using a combination of both rearing-types, there was no significant habitat effect on prey selection, but hatchery-reared red drum suffered higher overall mortality than wild-caught fish from pinfish predators. In these trials, spotted seatrout did not select for either prey type. Differences we observed in prey vulnerability were likely caused by behavioral differences between wild-caught and hatchery-reared red drum. Our results reinforce the conclusion that structural complexity in estuarine habitats increases survival of newly settled fishes. Our data also suggest that hatchery-reared red drum may be more vulnerable to predation than natural fishes, and that survival of stocked fish may be enhanced through habitat-related behavior modification.     

The food and feeding relationships of larval and 0+ year juvenile fishes in lowland rivers and connected waterbodies. II. Prey selection and the influence of gape

The relative importance of taxa- and size-specific prey selection, and the influence of gape on the prey consumed by the larvae and 0+ year juveniles of four fish species were investigated in 'main river', 'marina' and 'pond' macrohabitats in the lower River Trent, England. A general sequence of ontogenetic shifts in food consumption was reflected in the electivity indices of particular prey taxa, partly due to the restrictions imposed by the gape of 0+ year fishes. Certain taxa, however, were consistently selected over others, irrespective of size, suggesting that taxa-specific, as well as size-specific, prey characteristics may be important in the selection process. There were significant, positive relationships between maximum prey (zooplankton) length and maximum gape height for larvae, but not for 0+ year juveniles. The majority of fishes, however, consumed prey substantially smaller than the maximum theoretically possible inferred from their gape. The greater size ranges of zooplankton in connected waterbodies compared with main river channels provide suitable prey for a range of developmental steps and fish species, and may, thus, enhance recruitment success.     

Emergent impacts of cannibalism and size refuges in prey on intraguild predation systems

Many organisms undergo ontogenetic niche shifts due to considerable changes in size during their development. These ontogenetic shifts can alter the trophic position of individuals, the type and strength of ecological interactions across species, and allow for cannibalism within species. In this study we ask if and how the interaction of a size refuge and cannibalism in the prey alters the dynamics of intraguild predation (IGP) systems. By manipulating the composition of large cannibalistic (Aeshna umbrosa) and predatory (Anax junius) dragonfly larvae in mesocosms we show that the interaction of cannibals and predators was non-linear and increased the survival of prey. The structure of the final resource community shared by prey and predator differed between small and large dragonfly treatments but not within size classes across species. In general, the small prey stage showed similar shifts in microhabitat use and refuge use when exposed to either conspecific cannibals or predators, while large cannibals showed no clear anti-predator response. However, further behavioral experiments revealed that specific behavioral components, such as distances between individuals or number of movements, differed when individuals were exposed to either cannibals or predators. This indicates that individuals discriminated between conspecific or heterospecific predators. Furthermore, in similar experiments large cannibals and predators showed different behaviors when exposed to conspecifics rather than to each other. These changes in behavior are consistent with the observed increase in prey survival. In general, the results indicate that cannibalism and ontogenetic niche shifts can result in behavior-mediated indirect interactions that reduce the impact of the predator on the mortality of its prey and alter the interactions of IGP systems. However, they also indicate that size is not the sole determinant and that we also need to account for the species identity when predicting the dynamics of communities.     

Vulnerability of marine forage fishes to piscivory: effects of prey behavior on susceptibility to attack and capture

We conducted a series of size-structured laboratory experiments to quantify and compare the susceptibility of several estuarine and marine forage fishes to attack and capture by piscivorous predators. Size-dependent estimates of capture success, handling time, and prey profitability were generated from single-species experiments offering bay anchovy, Atlantic menhaden, Atlantic silverside, and age-0 striped bass to piscivores. Bay anchovy and Atlantic menhaden were most susceptible to capture and yielded high profitability compared to Atlantic silverside and age-0 striped bass prey. Variation in capture success among forage species was particularly influential in generating disparate profitability functions. Although morphological differences among forage species contributed to variation in susceptibility to predation, behavioral analyses indicated that variable reaction distances to approaching predators and activity levels of prey may explain a large fraction of the observed differences in susceptibility. When several forage species were offered to predators simultaneously in larger enclosures, mortality was highest and occurred earlier for bay anchovy and Atlantic menhaden compared to other prey, which points to the strong influence of predator capture success on overall forage fish vulnerability. Our results demonstrate species-specific differences among forage fishes in the ability to avoid attack and capture by piscivores, and we conclude that the expression of antipredator behaviors contributes significantly to variation in forage species vulnerability.     

Susceptibility of Atlantic herring and plaice larvae to predation by juvenile cod and herring at two constant temperatures

Predation encounters were staged in the laboratory to compare prey responsiveness, predator error rate, and predator capture success for juvenile cod Gadus morhua (a suction feeder) and herring Clupea harengus (a biting predator) preying on herring and plaice Pleuronectes platessa larvae. Trials were conducted at near natural temperature extremes for the larvae (8 and 13°C) to assess the importance of water temperature to the interaction. Herring larvae were significantly more responsive to attacks by herring than were plaice larvae (5·7 vs 0'0%). The two prey species were equally responsive to attacks by cod (2–6 vs 10%). Cod caught 91% of herring larvae attacked and juvenile herring caught 87%. Cod were successful in 96% of attacks on plaice, but juvenile herring caught significantly fewer (83%) plaice larvae. For each predator species, capture success did not vary significantly with prey species. Overall capture success for herring was significantly lower than that for cod. Responsiveness of herring larvae to attacks by juvenile herring increased with temperature, but predator error rate and capture success were not altered by water temperature.     

The influence of oral anatomy on prey selection during the ontogeny of two percoid fishes,Lagodon rhomboides and Centropomus undecimalis

Synopsis Ontogenetic increases in mouth size and changes in dentition of percoid fishes may affect the size and species of prey selected, thus influencing the fundamental trophic niche. To examine the influence of oral anatomy on prey selectivity by pinfish, Lagodon rhomboides, and snook, Centropomus undecimalis, two co-occurring percoid fishes with contrasting mouth morphologies, the mouth size, dentition, stomach contents, and available prey during ontogeny were quantified. Based on the presence of prey fragments in stomach contents and direct behavioral observation, prey were categorized by the feeding mode used during capture (suction/ramfeeding or biting). Centropomus has a larger size-specific gape than Lagodon during all ontogenetic stages. Although both feeding modes were used by Lagodon during ontogeny, the amount of prey captured using suction/ram-feeding declined and the amount of prey captured by biting increased with standard length. This change in feeding mode was associated with a change in incisor shape and width: Lagodon < 39 mm SL possessed narrow, pointed incisors and strongly selected amphipods, which are captured using suction/ram-feeding; Lagodon> 40 mm SL possessed wide, flat-topped incisors and significantly increased their selectivity for polychaetes, which are captured by biting. Centropomus used ram-feeding to capture prey at all ontogenetic stages. Size-selective feeding by Centropomus was apparent but could not be due to gape-limitation alone, because average prey body depth was only 45% of gape and was not proportional to absolute mouth size increase during ontogeny. Dietary diversity was greatest during the transition from suction/ram-feeding to biting in Lagodon. Lagodon had a higher dietary diversity at all ontogenetic stages than Centropomus, due in part to Lagodon's use of multiple feeding modes.     

Predicting predation through prey ontogeny using size-dependent functional response models

The functional response is a critical link between consumer and resource dynamics, describing how a consumer's feeding rate varies with prey density. Functional response models often assume homogenous prey size and size-independent feeding rates. However, variation in prey size due to ontogeny and competition is ubiquitous, and predation rates are often size dependent. Thus, functional responses that ignore prey size may not effectively predict predation rates through ontogeny or in heterogeneous populations. Here, we use short-term response-surface experiments and statistical modeling to develop and test prey size-dependent functional responses for water bugs and dragonfly larvae feeding on red-eyed treefrog tadpoles. We then extend these models through simulations to predict mortality through time for growing prey. Both conventional and size-dependent functional response models predicted average overall mortality in short-term mixed-cohort experiments, but only the size-dependent models accurately captured how mortality was spread across sizes. As a result, simulations that extrapolated these results through prey ontogeny showed that differences in size-specific mortality are compounded as prey grow, causing predictions from conventional and size-dependent functional response models to diverge dramatically through time. Our results highlight the importance of incorporating prey size when modeling consumer-prey dynamics in size-structured, growing prey populations.     

The foraging ecology of larval and juvenile fishes

Knowledge of the foraging ecology of fishes is fundamental both to understanding the processes that function at the individual, population and community levels, and for the management and conservation of their populations and habitats. Furthermore, the factors that influence the acquisition and assimilation of food can have significant consequences for the condition, growth, survival and recruitment of fishes. The majority of marine and freshwater fish species are planktivorous at the onset of exogenous nutrition and have a limited ability to detect, capture, ingest and digest prey. Improvements in vision, development of fins and associated improvements in swimming performance, increases in gape size and development of the alimentary tract during ontogeny often lead to shifts in diet composition. Prey size, morphology, behaviour and abundance can all influence the prey selection of larval and juvenile fishes. Differences in feeding behaviour between fish species, individuals or during ontogeny can also be important, as can inter- and intraspecific interactions (competition, predation risk). Temporal (diel, seasonal, annual) and spatial (microhabitat, mesohabitat, macrohabitat, regional) variations in prey availability can have important implications for the prey selection, diet composition, growth, survival, condition and, ultimately, recruitment success of fishes. For fish populations to persist, habitat must be available in sufficient quality and quantity for the range of activities undertaken during all periods of development. Habitats that enhance the diversity, size ranges and abundance of zooplankton should ensure that sufficient food resources are available to larval and juvenile fishes.     

Group predatory behavior by the assassin bugAgriosphodrus dohrni Signoret (Hemiptera: Reduviidae)

Nymphs of Agriosphodrus dohrniSignoret (Reduviidae) have a strong gregariousness and show group predatory behavior. This study was conducted to clarify adaptive significance of group predation of this species, including laboratory observations and 6-year field surveys. In the laboratory, observations on both solitary and group attacking against armyworms were made at varying prey size classes to compare the capture success rate by solitary predators with that by groups. The efficiency in capturing the prey was significantly higher in group attacking at any prey size class compared. Data obtained from the field surveys indicated the tnedency for searching nymphs to feed in group and to increase the number of predators feeding per prey item with increasing prey size. Average sizes of prey captured were also larger in group feeding throughout the nymphal stage. In particular, it was remarkable that, when prey were “creeping” types, the upper size limit of prey eaten was dramatically increased.     

The effectiveness of eyespots and masquerade in protecting artificial prey across ontogenetic and seasonal shifts

When constraints on antipredator coloration shift over the course of development, it can be advantageous for animals to adopt different color strategies for each life stage. Many caterpillars in the genus Papilio exhibit unique ontogenetic color sequences: for example, early instars that masquerade as bird feces, with later instars possessing eyespots. I hypothesize that larvae abandon feces masquerade in favor of eyespots due to ontogenetic changes in signaler size. This ontogenetic pattern also occurs within broader seasonal shifts in background color and predator activity. I conducted predation experiments with artificial prey to determine how potential signaling constraints (specifically size and season) shape predation risk, and consequently the expression of ontogenetic color change in Papilio larvae. Seasonally, both predation and background greenness declined significantly from July to September, though there was little evidence that these patterns impacted the effectiveness of either color strategy. Caterpillar size and color strongly affected the attack rate of avian predators: attacks increased with prey size regardless of color, and eyespotted prey were attacked more than masquerading prey overall. These results may reflect a key size-mediated tradeoff between conspicuousness and intimidation in eyespotted prey, and raise questions about how interwoven aspects of behavior and signal environment might maintain the prevalence of large, eyespotted larvae in nature.     

Do Experience and Body Size Play a Role in Responses of Larval Ringed Salamanders,Ambystoma annulatum,to Predator Kairomones? Laboratory and Field Assays

Prey may experience ontogenetic changes in vulnerability to some predators, either because of changes in morphology or experience. If prey match their level of antipredator behavior to the level of predatory threat, prey responses to predators should reflect the appropriate level of threat for their stage of development. For larval salamanders, responses to predators may change with body size because larger larvae are less vulnerable to predation by gape‐limited predators or because fleeing responses by large salamanders may be more effective than for smaller salamanders. In a field experiment, small larval ringed salamanders, Ambystoma annulatum, responded to chemical stimuli (‘kairomones’) from predatory newts, Notophthalmus viridescens, with an antipredator response (decreased activity). Laboratory‐reared larvae decreased their activity following exposure to newt kairomones, indicating that larval ringed salamanders do not require experience with newts to recognize them as predators. In both experiments, larvae distinguished between chemical stimuli from newts and stimuli from tadpoles (non‐predators) and a blank control. In a third experiment, field‐caught (experienced) larvae showed a graded response to newt kairomones based on their body size: small larvae tended to decrease their activity while larger larvae showed no change or an increase in activity. This graded response was not observed for neutral stimuli, indicating that it is predator‐specific. Therefore, ringed salamander larvae exhibit threat‐sensitive ontogenetic changes in their response to chemical stimuli from predatory newts.     

A parasite's modification of host behavior reduces predation on its host

Parasite modification of host behavior is common, and the literature is dominated by demonstrations of enhanced predation on parasitized prey resulting in transmission of parasites to their next host. We present a case in which predation on parasitized prey is reduced. Despite theoretical modeling suggesting that this phenomenon should be common, it has been reported in only a few host–parasite–predator systems. Using a system of gregarine endosymbionts in host mosquitoes, we designed experiments to compare the vulnerability of parasitized and unparasitized mosquito larvae to predation by obligate predatory mosquito larvae and then compared behavioral features known to change in the presence of predatory cues. We exposed Aedes triseriatus larvae to the parasite Ascogregarina barretti and the predator Toxohrynchites rutilus and assessed larval mortality rate under each treatment condition. Further, we assessed behavioral differences in larvae due to infection and predation stimuli by recording larvae and scoring behaviors and positions within microcosms. Infection with gregarines reduced cohort mortality in the presence of the predator, but the parasite did not affect mortality alone. Further, infection by parasites altered behavior such that infected hosts thrashed less frequently than uninfected hosts and were found more frequently on or in a refuge within the microcosm. By reducing predation on their host, gregarines may be acting as mutualists in the presence of predation on their hosts. These results illustrate a higher‐order interaction, in which a relationship between a species pair (host–endosymbiont or predator–prey) is altered by the presence of a third species.     

Environmental variation and behaviour: resource availability during ontogeny and feeding performance in salamander larvae (Ambystoma texanum)

1. The effects of resource availability during ontogeny on subsequent feeding performance were investigated in larvae of the small-mouthed salamander ( Ambystoma texanum ).
2. Salamander larvae were reared individually in either high or low prey density treatments for 7 weeks prior to intermediate prey density foraging trials. Larvae from the low prey density treatment were on average 35% smaller in body size than individuals from the high prey density treatment.
3. Resource availability during development influenced larval feeding rates and altered the relationship between body size and three feeding performance measures (attack rates, capture success and feeding rates). Feeding rates in predation trials were also positively correlated with growth rate early in the larval period (until the end of week 5).
4. These results suggest that the environment to which developing organisms are exposed can have significant effects on subsequent behaviour, and that small-mouthed salamander larvae may show state-dependent changes in feeding behaviour in response to differences in long-term feeding history. Additionally, differences in feeding performance may influence the probability of survival to the adult stage for organisms that utilize ephemeral habitats.     

Who eats whom in a pool? A comparative study of prey selectivity by predatory aquatic insects

Predatory aquatic insects are a diverse group comprising top predators in small fishless water bodies. Knowledge of their diet composition is fragmentary, which hinders the understanding of mechanisms maintaining their high local diversity and of their impacts on local food web structure and dynamics. We conducted multiple-choice predation experiments using nine common species of predatory aquatic insects, including adult and larval Coleoptera, adult Heteroptera and larval Odonata, and complemented them with literature survey of similar experiments. All predators in our experiments fed selectively on the seven prey species offered, and vulnerability to predation varied strongly between the prey. The predators most often preferred dipteran larvae; previous studies further reported preferences for cladocerans. Diet overlaps between all predator pairs and predator overlaps between all prey pairs were non-zero. Modularity analysis separated all primarily nectonic predator and prey species from two groups of large and small benthic predators and their prey. These results, together with limited evidence from the literature, suggest a highly interconnected food web with several modules, in which similarly sized predators from the same microhabitat are likely to compete strongly for resources in the field (observed Pianka's diet overlap indices >0.85). Our experiments further imply that ontogenetic diet shifts are common in predatory aquatic insects, although we observed higher diet overlaps than previously reported. Hence, individuals may or may not shift between food web modules during ontogeny.