Differential vulnerability determines the diet of a slow-moving predatory stream insect

1. We investigated the diet and prey electivity of Rhyacophila obliterata, a slow‐moving invertebrate predator capable of hunting in high‐flow microhabitats, and quantified the components of the predation sequence of fifth‐instar larvae foraging on mobile (Baetis mayflies, Amphinemura stoneflies) versus semi‐sessile (larval blackflies) prey. 2. In the field, fifth‐instar Rhyacophila consistently took more larval blackflies than more mobile prey. In behavioural trials, the number of attacks by Rhyacophila differed significantly between prey types, mobile prey being attacked more often than blackflies. Capture success, by contrast, was highest for blackflies, whereas Amphinemura and Baetis were rarely captured. In mixed‐prey feeding trials, Rhyacophila showed strong preference for blackflies and equally strong avoidance of Amphinemura and Baetis. 3. For mobile prey, the risk of being captured by this sluggish predator is very low, so they can afford to be in close contact with it. Rhyacophila was almost unable to capture any other prey but blackflies, resulting in strong passive selection for blackflies. 4. Therefore, the diet of fifth‐instar Rhyacophila can be predicted from laboratory observations and prey behaviour is the major determinant of the diet of this invertebrate predator.     

Drift behaviour and microhabitat selection in the preimaginal stages of Simulium chutteri (Diptera Simuliidae)

The aquatic stages of the cattle biting pest, Simulium chutteri Lewis utilize river flow in their dispersal and colonization behaviour. Peaks of drifting activity in S. chutteri larvae occurred in the early morning and late afternoon. It is deduced that female flies scatter eggs onto slow flowing waters upstream of rapids. Larvae hatch from drifting eggs and colonize substrates in slower flowing regions upstream of rapids, while later stage instars move into faster flowing regions within rapids where they complete their development. This microhabitat selection by S. chutteri leads to rapid attainment of large population sizes in suitable sections of river and reduces competition between different stage larvae. In their utilization of a variety of microhabitats the larvae of this species differ from co-existing simuliid species which restrict developmental stages to single habitats.Catastrophic drift was recorded for S. chutteri larvae and could be a mechanism to regulate population size.Drift of simuliid larvae off rapids was not related to benthic population densities in the rapids and was therefore not due to excessive production. It is concluded that larval drift off rapids is related to habitat disturbances associated with water flow fluctuations and the activity of aquatic predators and other animals.The distribution of S. chutteri in the Vaal River is restricted by oviposition requirements of the adult female. Knowledge of drift behaviour and water flow requirements of Simuliidae have been applied to manage the population size of S. chutteri in the Vaal River.     

Estimation of preferred water flow parameters for four species of Simulium (Diptera: Simuliidae) in small clear streams in South Africa

Blackfly larvae typically occur in fast-flowing riffle sections of rivers, with different blackfly species showing preferences for different hydraulic conditions. Very little quantitative data exist on hydraulic conditions linked to the blackfly species occurring in South African streams. Stones-in-current biotopes (i.e. fast riffle flows over cobbles) were sampled from four sites in three small clear streams in the Eastern and Western Cape provinces of South Africa. Mean water column velocities at each sampled stone were measured using a mini current meter, while flow velocities closer to the boundary layer where blackfly larvae occurred were estimated using indirect techniques (standard hemispheres and aerating tablets). Standard hemispheres were also used to calculate more complex hydraulic parameters such as Froude and Reynolds numbers. Four species of Simuliid were sampled in sufficient numbers to show trends in flow velocity preferences. Simulium impukane and S. rutherfoordi both occurred at their highest densities at velocities of 0.3m s^?1, while S. merops preferred velocities of 0.7m s^?1. Simulium nigritarse SL attained the highest densities of all the blackfly species sampled, and its relative abundances were greatest at velocities of 0.8–0.9m s^?1. Within the streams surveyed, all blackfly species occurred in subcritical-turbulent flows — based on a classification using Froude and Reynolds numbers — although two of the species were also found in high densities in supercritical flows where these existed at the sites. Local hydraulics within the stones-in-current biotope are complex, but in the absence of fine-scale equipment for measuring micro-velocities, standard hemispheres are a useful, cost-effective technique for the initial quantification of hydraulic parameters in small, clear streams. Such an approach facilitates further understanding of links between hydraulics and aquatic invertebrates in South African streams.     

The costs of reduced feeding due to predator avoidance: potential effects on growth and fitness in Ischnura elegans larvae (Odonata: Zygoptera)

ABSTRACT.

• 1 This paper investigates the behaviour, in the laboratory, of a forager simultaneously confronted with the conflicting needs to feed and to avoid predators. The foragers were larvae of the damselfly Ischnura elegans Van der Linden, feeding on Daphnia magna Strauss. The predators were adult females of Notonecta glauca L.
• 2 Patch choice by Ischnura larvae was significantly modified by the presence of predators. Larvae moved to feed in patches of high prey density when predators were absent but preferred dense cover, even though virtually no prey were available, when predators were present. This behaviour was not altered by hunger, up to 12 days without food. In other words, Ischnura larvae were risk averse in their foraging behaviour.
• 3 In experiments with abundant prey available, the feeding rates of Ischnura larvae confined to a single patch were also significantly reduced by the presence of hydrodynamically and chemically detectable predators. Predators detectable only by vision had little effect.
• 4 Calculations made from published data show that reduced larval feeding rates can lead to slower growth and development and prolonged instar durations in Ischnura elegans larvae. This may have important consequences for larval survival and adult reproductive fitness.

     

Predator-prey interactions in a benthic stream community: a field test of flow-mediated refuges

Ecological theory suggests that the impact of predation can be strongly modified by the existence of regions of the environment in which prey are less accessible to predators, which underscores the need for empirical studies examining the factors influencing the availability and importance of such prey refuges. Our study tested whether benthic microhabitats with high flows provide suspension-feeding larval black flies (Simulium␣vittatum) with a spatial refuge in which the negative impact of predatory flatworms (Dugesia dorotocephala) is reduced. We conducted a short-term field experiment in Chester Creek (southeastern Pennsylvania, United States) to examine how the number of black fly larvae inhabiting tile substrates responded to manipulated variations in flatworm abundance and current speed. The abundance of flatworms declined with increasing current speed, thereby creating the potential for sites with high flows to provide larvae with a refuge from these predators. Multiple regression analysis revealed that the final abundance of larvae exhibited a significant negative relationship to flatworm abundance and a significant positive relationship to current speed. After adjusting for variations in elapsed time and initial larval abundance, flow and predators explained 38% of the variation in the rate of change in larval abundance. The positive correlation between larval abundance and flow had two components: a positive, direct effect of flow on larvae, which arises because these food-limited consumers prefer to reside within sites with faster flows where they can feed at higher rates; and a negative effect of flow on predators, and of predators on larvae, which combine to yield a positive indirect effect of flow on larvae. This indirect effect demonstrates the existence of flow-mediated refuges (i.e., microhabitats in which the impact of predation is reduced due to high flows), although the effect accounts for a small proportion of total variation in larval abundance. A consideration of biomechanical relationships suggests that microhabitats with high flows are likely to create prey refuges in a wide range of freshwater and marine benthic environments. In particular, predators will often experience greater dislodgement forces than prey because of their larger size and because they project farther above the bed where current speeds are faster. Moreover, the ability to resist a given dislodgement force may be greater for many prey, especially those that are sessile or semi- sessile. Received: 31 March 1997 / Accepted: 25 October 1997     

Blackfly (Diptera: Simuliidae) abundance in a lake outlet: test of a predictive model

A mathematical model based on assumptions of proportionality of filter-feeding insect larvae and their food supply and of the ability of the larvae to substantially reduce the sestonic food supply was developed. The predictions of the model were tested by censusing simuliid blackfly larvae in the outlet stream of a mesotrophic lake. Observed trends in simuliid density agree closely with the predictions. Two seston components (diatoms, detritus) increase downstream contrary to prediction. Potential predators of simuliids were censumed. Isoperla spp. decreased with increasing distance below the lake while the perlid stoneflies Calineuria and Hesperoperla seemed to be excluded from the outfall region.     

Balancing food availability and hydrodynamic constraint: phenotypic plasticity and growth in Simulium noelleri blackfly larvae

Organisms through phenotypic plasticity can cope with multiple changed environmental conditions. Theory predicts that animals in streams and rivers should be able to balance demands of the needs to obtain food efficiently and to adjust response to hydrodynamic variability. This study examined effects of variations in food availability and current velocity on the feeding structure and growth rate of Simulium noelleri blackfly larvae. The larvae developed larger labral fans and more rays under slow current and low food regimes than in fast current and high food conditions. In both fast and slow current regimes, growth rates were higher and development periods to the final-instar stage were shorter in high food treatments. The estimated flux rates of food particles through labral fans under high food treatments for both fast and slow current regimes were higher than those under low food treatments. Although both food and current velocity appeared to have selected for flexibility of feeding structure and growth rate, food availability was a more important factor for phenotypic and developmental plasticity than current velocity. The results indicate a strong link between environmental changes in food availability and current velocity, phenotypic plasticity, and growth rate of S. noelleri. This study suggests that plasticity of ecomorphs with macroevolutionary significance may play a role in the early evolutionary stages of blackfly larvae.     

The life cycle and growth ofIsoperla grammatica andI. difformis (Plecoptera) in southernmost Sweden: intra- and interspecific considerations

Studies on the life cycle and growth ofIsoperia grammatica andI. difformis from eight southern Swedish streams demonstrated that the former species shows greater between-stream differences than the latter species.I. grammatica larvae had typically two periods of rapid growth, autumn and spring, whileI. difformis larvae grew mainly in autumn-winter. In one of the streams, however,I grammatica grew rapidly throughout all winter. Maximum larval size differed between streams, and so did the timing of the first appearance of nymphs and nymphal maturation. Temperature was probably the most influential factor explaining all these differences. While sexual dimorphism inI. grammatica is slight but significant, it is very prominent inI. difformis. The sizes of larvae overlapped little between species as well as sexes suggesting that intra- and interspecific competition was, or had been, important. InI. grammatica this reduced overlap was expressed in several size-classes of males and females successively replacing each other, resulting in a meandering pattern of the size distribution charts. There were additional indications of possible interactions, such as a significant negative correlation between the densities of the two species.     

Unusual behaviour and morphology of some Rhyacophila Pictet, 1834 caddisfly (Trichoptera: Rhyacophilidae) larvae reflect their ability to use the hyporheic zone

We found that larvae of four Rhyacophila Pictet, 1834 species preferred the hyporheic biotope in rapids and glides, which appears to be an unusual habitat among species in this genus. These species, i.e., Rhyacophila nigrocephala Iwata, 1927, R. nipponica Navas, 1933, R. shikotsuensis Iwata, 1927 and R. kawamurae Tsuda, 1940, belonging to the nigrocephala species group, of which six species occur in Japan. To study the movement behaviour of these species in the hyporheic biotope, we introduced their larvae into a small aquarium with a sandy bottom substrate. The larvae of these species burrowed into the sand bed and moved smoothly through interstices using their forelegs and their highly flexible and elastic abdomens. The larval morphology of these six species differs from that of the representatives of the other species group of Japanese Rhyacophila. They have more slender and flatter head capsules, more elongate abdominal segments, shorter thoracic legs and more slender anal prolegs. These features are adaptations that allow these species to effectively use the hyporheic biotope by enabling them to burrow through the interstices.     

Predators of citrus blackfly [Hom.: Aleyrodidae]

A 2-year survey of known whitefly predators was conducted from winter 1976 to winter 1978 in Broward County, Florida. Also, the impact of predators on citrus blackfly populations was measured under field conditions using lifetable data and predator exclusion data. In addition, feeding experiments were conducted to determine specific predators on different citrus blackfly stages. In the survey, the total number of predators caught in 1977 was significantly greater than 1978 and reasons for this difference are discussed. Spiders and the coccinellid,Delphastes pusillus Lec. were the most abundant predators being ca. 90% of all predators caught. Results from the lifetable and predator exclusion technique were similar showing that predators caused a 52 to 66% reduction in citrus blackfly populations. Feeding tests showed that predator responses to feeding on citrus blackfly were variable depending on species of predator, stage of predator, and stage of citrus blackfly.

---

Résumé Une étude pendant 2 ans des prédateurs de l'aleurode des citrus a été réalisée de l'hiver 1976 à l'hiver 1978 dans le Comté de Broward en Floride. L'influence des prédateurs sur les populations d'aleurodes a été mesurée dans les conditions naturelles par l'élaboration d'une table de vie et par des essais d'exclusion des prédateurs. En outre des expériences ont été effectuées afin de déterminer les prédateurs spécifiques pour chacun des stades de l'aleurode. Le nombre total de prédateurs capturés en 1977 fut significativement plus élevé qu'en 1978; les causes de cette différence sont discutées. Des araignées et la coccinelleDelphastes pusillus Lec. ont été les plus abondants, représentant et iron 90% du total. Les résultats de la table de vie et de l'exclusion des prédateurs ont été analogues et montrent que ceux-ci provoquent une réduction de 52 à 66% des populations d'aleurodes. Les essais sur l'alimentation indiquent que les réponses ont été variables selon l'espèce de prédateur, le stade du prédateur et de l'aleurode.

---

---

Florida Agricultural Experiment Station Journal Series No. 1791. Partly supported by funds from Cooperative Agreement # 12-14-7001-1148 between U.S.D.A., and University of Florida.     

Effects of insect damage on photosynthesis,transpiration and SO2 uptake by sycamore

Summary The bilberry (Vaccinium myrtillus L.), is the dominant plant in the field layer of many boreal forests in northern Sweden. It is utilized by several herbivorous insect larvae as food (i.e. Lepidoptera: Geometridae, Tortricidae and Hymenoptera: Symphyta). Total density of insect larvae was 63% lower where birds had access to larvae compared to exclosures. Larvae with a feeding behaviour which exposes them to birds (geometrids and sawflies) showed a pronounced reduction in density when exposed to bird predators. Density of larvae with a concealed feeding behaviour (tortricids) was unaffected by birds. The frequency of larval damage to bilberry annual shoots was significantly higher in areas where larval density was high due to exclusion of birds. I concluded that birds have the potential to heavily modify the interaction between bilberry and insect larvae.     

Contrasting diel activity and feeding patterns of four species of carnivorous stoneflies

1. The hypothesis under test was that larvae of Dinocras cephalotes (Curtis), Perla bipunctata Pictet, Isoperla grammatica (Poda), and Perlodes microcephalus (Pictet) differed markedly in their diel activity and feeding patterns. Mature larvae collected about 1 month prior to adult emergence were used either for gut analyses or for observations of their activity and feeding patterns in three stream tanks with natural substrata and glass bottoms, so that activity could be observed above and below the substratum. A dull red light was used for observations in the dark. Food (larvae of Ephemeroptera, Simuliidae, and Chironomidae) was provided in excess. 2. Larvae for gut analyses were taken 1 h before dusk or dawn (n = 30 larvae per species for each day or night sample). The only species with food in the gut for the day samples was P. microcephalus. All species fed at night, the mean number of prey per larva being very similar for D. cephalotes, P. bipunctata, and I. grammatica but significantly higher for P. microcephalus. Most prey were insect larvae, especially Simuliidae and Chironomidae. 3. Diel activity patterns of single larvae differed interspecifically but not intraspecifically. Larvae of D. cephalotes and P. bipunctata were rarely active during the day; their activity increased at dusk and decreased at dawn, and was highest during the night; their success at prey capture was highest at dusk and dawn, with an ambush rather than a search strategy. Isoperla grammatica was rarely active during the day, most active at dusk and dawn when prey capture was highest, using a search strategy, and less active for the rest of the night. Perlodes microcephalus was active during the day, but only below the substratum, and very active from dusk to dawn with a high prey‐capture success, using a search strategy. These experiments provided support for the hypothesis under test. The discussion concludes that the results could also help to explain known differences in growth rate and the length of the life‐cycle in these four species.     

A comparative study of the functional response of four species of carnivorous stoneflies

SUMMARY 1. Comparisons were made of the functional responses of mature larvae of Perlodes microcephalus, Isoperla grammatica, Dinocras cephalotes and Perla bipunctata. Experiments were performed in stream tanks with natural substrata and glass bottoms, so that feeding could be observed above and below the substratum. There was one stonefly per tank and one of 10 prey densities between 20 and 200 larvae of either Chironomus sp. or Baetis rhodani per tank. Consumed prey were replaced in a first set of experiments but not in a second set. Additional experiments assessed intraspecific interference between larvae of each predator species (two to five predators per tank). 2. The number of prey eaten increased curvilinearly with prey density. The relationship was described by two models, a Type II instantaneous model and its integrated equivalent, for experiments with and without prey replacement, respectively. Handling time did not change significantly with prey density, and was the same for experiments with and without prey replacement. Estimates of attack rate were similar for the two models, but varied between prey type and predator species. Handling time varied considerably but was normally distributed for each prey type and predator species. Mean handling time varied for Chironomus from 39 s for Isoperla, which rarely ate a whole larva, to 57 s for Perlodes and for Baetis from 116 s for Perlodes to 167 s for Perla. All predators were more efficient at capturing Baetis, but the longer handling time for Baetis ensured that more Chironomus were eaten. It was concluded that these variations in attack rate and handling time were related to activity and growth differences between the predator species, and that experiments with and without prey replacement could both be relevant to the field, depending on how the predators searched for food. 3. In the interference experiments, mean handling time did not change with increasing predator density, but attack rate decreased curvilinearly, the decrease varying from negligible for Isoperla to marked for Perlodes. Prey capture decreased with decreasing attack rate. Therefore, interference reduced prey consumption, but this effect was negligible for Isoperla and increasingly severe in the order Dinocras, Perla and Perlodes.     

Evaluation of in situ enclosures during a study of the importance of starvation to the vulnerability of herring larvae to a piscine predator

Two competing hypotheses about the effect of starvation on the vulnerability of fish larvae to predators were tested in large enclosures (mesocosms). One hypothesis predicts greater vulnerability as starvation weakens larvae. The alternative recognizes that the behaviour of starving larvae may reduce encounters with predators or fail to elicit predatory attacks, thereby decreasing vulnerability. Results of four large-scale experiments, in which progressively starved Atlantic herring Clupea harengus L. larvae were exposed to larger herring as predators, were unable to support either hypothesis because of a large amount of variability. Much of the variability was thought to be due to erratic feeding behaviour of the predators. Additional experimental observations suggest that sunshine and predator satiation can affect these experiments.     

Effects of Bt maize on Frankliniella tenuicornis and exposure of thrips predators to prey-mediated Bt toxin

As a part of a risk assessment procedure, the impact of Bt maize expressing Cry1Ab toxin on the thrips Frankliniella tenuicornis (Uzel) (Thysanoptera: Thripidae) was investigated, and the potential risks for predators feeding on thrips on Bt maize were evaluated. The effects of Bt maize on F. tenuicornis were assessed by measuring life‐table parameters when reared on Bt and non‐Bt maize. The content of Cry1Ab toxin in different stages of F. tenuicornis reared on Bt maize and the persistence of the toxin in adults where determined in order to evaluate the possible exposure of predators when feeding on thrips. In addition, Chrysoperla carnea (Stephens) (Neuroptera: Chrysopidae) was used as a model predator to assess how the behaviour of prey and predator may influence the exposure of a natural enemy to the Bt toxin. Life‐table parameter results showed that F. tenuicornis was not affected when it was reared on Bt maize. This indicates that the potential for prey quality‐mediated effects on predators is low. Bt content was highest in thrips larvae and adults, and negligible in the non‐feeding prepupal and pupal stages. The persistence of the Cry1Ab toxin in adult F. tenuicornis was short, resulting in a decrease of 97% within the first 24 h. Predation success by young C. carnea larvae varied among the thrips stages, indicating that exposure of predators to Bt toxin can additionally depend on the prey stage. When combining the current knowledge of the susceptibility of major thrips predators with our findings showing no potential for prey quality‐mediated effects, relatively low toxin content in thrips as well as short persistence, it can be concluded that the risks for predators when feeding on thrips in or next to Bt maize fields are negligible.     

Alarm response in larval western toads,Bufo boreas: release of larval chemicals by a natural predator and its effect on predator capture efficiency

An alarm response in larvae of the western toad, Bufo boreas, was examined in the laboratory. A natural predator, the giant water bug, Lethocerus americanus, while capturing and feeding on a tadpole, can cause sufficient damage for the release of toad alarm substance and subsequent alarm response in prey conspecifics. Larvae increased their activity and avoided the side of the tank that had a predator feeding on a conspecific tadpole in a visually isolated but interconnected container. Larvae did not increase activity or avoid the side of the tank that had a predator feeding on a heterospecific tadpole. Performance of the alarm response in conspecifics increased survivorship. The capture efficiency of predators (naiads of the dragonfly Aeshna umbrosa) hunting larvae that had been exposed to the alarm substance of conspecifics was reduced: predators required more capture attempts per successful attack compared to control tests.     

Resident spiders as predators of the recently introduced light brown apple moth,Epiphyas postvittana

The invasive light brown apple moth, Epiphyas postvittana (Walker) (Lepidoptera: Tortricidae), has not reached damaging levels on crops in California (USA), suggesting that its populations and impact are limited by generalist natural enemies. In a series of laboratory experiments, we examined resident spiders as predators of E. postvittana larvae on two host plants, the ornamental Australian tea tree, Leptospermum laevigatum (Gaertn.) F. Muell (Myrtaceae), and the weed French broom, Genista monspessulana (L.) L.A.S. Johnson (Fabaceae). Of three abundant spider species in Australian tea tree, two hunting spiders measurably reduced the numbers of E. postvittana larvae and plant damage, whereas a web‐weaving spider had no detectable impact. The adult stage of the dominant hunting spider Anyphaena aperta Banks (Anyphaenidae) consumed E. postvittana larvae, but neither large nor small juveniles had statistically detectable effects on numbers of larvae. However, plant damage was reduced in the presence of large juvenile A. aperta, suggesting that A. aperta may also have non‐consumptive effects on the feeding behavior of E. postvittana larvae. Anyphaena aperta consumed larvae only when larval densities exceeded a low threshold in a functional response experiment, leading to a type III functional response. Adult A. aperta showed no preference for different E. postvittana instars, whereas Cheiracanthium mildei L. Koch (Miturgidae), an abundant hunting spider on French broom, showed a partial preference for late‐instar larvae. The generalist feeding habits of the spiders may have precluded strong prey preferences. Results show that hunting spiders may help limit E. postvittana populations in California, and that they may in turn reduce the impact of E. postvittana on its host plants.     

Movement and disappearance of mountain birch defoliators are influenced by the interactive effects of plant architecture and induced resistance

Abstract. 1. Changes in herbivore movement and feeding behaviour may determine the efficacy of induced plant resistance by affecting the location of damage within the foliage and by modifying the vulnerability of herbivores to predators. 2. Observations of larval feeding sites were used to test whether induced resistance increased the movement of free‐living Epirrita autumnata Borkh. (Lepidoptera, Geometridae) larvae feeding on mountain birch [Betula pubescens ssp. czerepanovii (Orlova) Hämet‐Ahti]. The amount of defoliation at different canopy parts was measured to test the associated changes in the spread of damage within the foliage. 3. The architectural complexity of trees was measured to test its association with the disappearance of larvae from their hosts. The underlying hypothesis was that the architectural traits of the host plant could affect disappearance by influencing the frequency of herbivores encountering predators. 4. Distance between the consecutive feeding positions, the number of leaves damaged, and consumption of long shoot leaves all increased in trees with induced resistance. 5. Disappearance of larvae depended on the architectural complexity of trees. The effect of complexity differed between defoliation treatments, and may depend on the activity and number of predators in relation to the canopy size. 6. Accordingly, this study suggested that the interactive effects of plant architecture, induced resistance, and herbivore behaviour can determine the performance of herbivores on their host plant.     

Assessment of trends in predation pressure on insects across temperate forest microhabitats

相似文献   

Different colonization patterns of two closely related stream insects (Austrosimulium spp.) following disturbance

SUMMARY.

• 1 In many streams, blackfly larvae rapidly colonize scoured surfaces, then decline in numbers as other species are still increasing. Such a colonization pattern may be generated by species that seek disturbed substrata because they prefer scoured surfaces having low epilithon cover (i.e. they are opportunists) or because of lowered faunal densities (i.e. they are fugitives), but this has rarely been tested.
• 2 In the Acheron River, south-eastern Australia, the larvae of the blackfly Austrosimulium torrentium show the typical pattern of increase and decline whereas A. victoriae shows a different pattern of more gradual increase.
• 3 An experiment showed that A. torrentium, consistent with its colonization pattern, responded strongly to treatments that mimicked the effects of disturbance; it colonized bricks with few other animals present regardless of epilithon cover and therefore is potentially a fugitive species. A. victoriae did not respond convincingly to treatments, which is also consistent with its colonization pattern.
• 4 Blackfly larvae of both species preferentially colonized bricks that were in fast-moving water. Hence, A. torrentium only responded to the densities of other fauna when bricks occurred in currents greater than approximately 30 cm s^?1.
• 5 Whilst epilithon cover did not affect abundance of blackfly larvae, it did affect attachment positions. In one experimental treatment, epilithon was reduced on one half of the top surfaces of bricks. More than 50% of larvae on these bricks were attached to the epilithon-reduced halves.
• 6 Our results show that functional feeding or taxonomic groups do not necessarily contain ecologically equivalent taxa; colonization patterns may provide a more useful way of categorizing species for the purposes of modelling these assemblages.