The consequences of larval aggregation in the butterfly Chlosyne lacinia

1. Females of Chlosyne lacinia (Geyer) (Lepidoptera: Nymphalidae, Melitaenae), the bordered patch butterfly, clump eggs in a few large clusters on their host plant, Helianthus annuus. Resulting larvae form sibling aggregations to at least the third instar.
2. The effect of group size on survival and development of C. lacinia larvae was tested experimentally in the field. Larvae developed faster and survived better in larger groups.
3. The effects of various predator guilds (ground-dwelling arthropods, aerial arthropods and avian predators) on survival of larvae was then tested while controlling group size. Ground-dwelling arthropods, mainly fire ants Solenopsis xyloni , reduced larval survival greatly but other solitary invertebrate and avian predators did not alter survival. Group defences and aposematism of C. lacinia larvae are probably ineffective against predatory ants that attack en masse and recruit other colony members.
4. In laboratory experiments, two possible mechanisms underlying faster development of larvae in larger groups were tested: (i) overcoming the physical toughness of host plant leaves, and (ii) social stimulus to feed. Results support the physical toughness hypothesis but not the social stimulus hypothesis.
5. Feeding in large groups by C. lacinia larvae confers multiple advantages, including protection from solitary predators and increased feeding efficiency because grouped, early-instar larvae can initiate feeding wounds on tough sunflower leaves. These advantages of larval gregariousness, coupled with reduced desiccation at the egg stage, apparently outweigh disadvantages of aggregation, such as interference and exploitative competition among larvae.     

Group size effects on survivorship and adult development in the gregarious larvae of Euselasia chrysippe (Lepidoptera, Riodinidae)

Caterpillars living in aggregations may derive several benefits that outweigh the costs, including better survivorship and improved growth rates. I tested whether larval group size had an effect on these two vital rates in Euselasia chrysippe. These caterpillars feed gregariously during all instars and move in processionary form over the host plant and even pupate together. There was a positive relationship between group size and larval survivorship in the field, although genetic variability was not taken into account in this experiment. Under laboratory conditions, there was also a positive relationship between group size, and larval growth rate and adult weight. This supports the hypothesis that aggregations facilitate feeding and larval growth. Single sixth instar larvae in the laboratory also had a lower survivorship than larvae in groups. These results provide further evidence of the benefits of group living for gregarious caterpillars.     

The effect of larval aggregation behaviour on larval growth of the spruce bark beetle Dendroctonus micans

1. The great spruce bark beetle Dendroctonus micans is a primary pest of spruce in Europe. It is unusual among Eurasian scolytids in that apparently healthy trees are attacked by solitary adults, but larvae feed en masse , in response to a larval aggregation pheromone.
2. The effect of brood size on larval growth was determined in experiments on detached bark, logs and living trees. A positive relationship was found between brood size and larval growth in experiments started with either eggs or larvae up to fifth instar. The relationship appeared to be independent of the effects of both preformed and induced defences in bark.
3. No evidence was found to support the hypothesis that larvae feeding in groups spend a greater proportion of time feeding. Addition of resin to egg chambers in detached bark pieces resulted in high mortality of first-instar larvae. A possible role for larval aggregation in minimizing the effects of host defences is discussed.     

Between-clutch interactions affect a benefit of group feeding for pipevine swallowtail larvae

Abstract.  1. In California, early instar larvae of the pipevine swallowtail ( Battus philenor ) develop at an accelerated rate when feeding in large groups compared with small groups due to a plant-mediated response to feeding group size. Larvae benefit from accelerated growth because the time larvae remain in early stages, where mortality is highest, is reduced. Occasionally, multiple clutches are laid on the same plant stem. Clutch size modification by females ovipositing on plant with previously laid clutches and the effect of kinship and group size on larval behaviour was examined. The direct and indirect interactions between clutches were investigated to determine if group size and time between clutch establishment affects the performance of early instar larvae.
2. Larger groups consume the young foliage more quickly and develop at an accelerated rate compared with smaller groups. Older foliage available to later clutches is an inferior food resource compared with younger foliage.
3. There was no evidence that females adjust clutch size in response to the presence of conspecific clutches.
4. Second groups of larvae readily joined previously established feeding groups. There were no observed behavioural differences between sibling and mixed-family groups.
5. The effect of a second group on the growth of the initial group was dependent on the size of both groups and the time interval between the arrival of the two groups.
6. Accelerated growth associated with larger feeding aggregations was absent when these groups were introduced to plants with previously established groups.
7. It is beneficial for ovipositing females to avoid plants with previously laid clutches because direct and indirect interactions with established clutches compromises larval performance.     

Some beneficial effects of aggregation in young larvae ofPryeria sinica moore (Lepidoptera: Zygaenidae)

The effects of group size on the survival and development of young larvae of Pryeria sinicaMoore were investigated by laboratory and field experiments. Under laboratory conditions, about 20% of isolated larvae died of unsuccessful feeding in the first instar, however, larvae survived successfully in aggregations of four or more individuals. In the field, larvae emerge in early spring and wait for new leaves to open before feeding. In this period, the larger the group size of hatchlings the survival rate became higher. The nest-web spun by hatchlings was considered to play an important role in protecting them from desiccation. In the period that larvae began to feed on leaves, more than 36 larvae are necessary to aggregate for the successful establishment of feeding groups. The nest-web played an important role also in the establishment of feeding group. However, the natural group size of the first instar larvae was larger than the minimum group size to spin a sound nest-web in the field experiment. On the other hand, in later stage, larvae in a large group did not have an excess advantage in survival or developmental rate over larvae in a small group. It was found that the experiments on survival and developmental rates could not explain the reason that this species maintain large compact groups in the most part of larval period.     

Larval gregariousness and neonate establishment of the eucalypt-feeding beetle Chrysophtharta agricola (Coleoptera: Chrysomelidae: Paropsini)

The selective advantage offered to individuals living within groups may relate to natural enemy defence, but in leaf feeding insects may also relate to overcoming plant defences, especially with respect to feeding establishment. We conducted a series of experiments focusing on neonate larval survival, examining the effect of group size and leaf age on the survival of a eucalypt-feeding beetle, Chrysophtharta agricola , which formed groups of up to 43 larvae on the foliage of Eucalyptus nitens in the field. In the laboratory, in the absence of natural enemies, we found that initial density, leaf age and damage to the leaf margin significantly affected larval survival. Survival of solitary first-instar larvae on young foliage was around 80% whereas on older foliage it was around 11%. Prior damage to the leaf margin significantly increased survival on older leaves to around 61%. Initial larval density also affected survival, although mortality was always significantly higher on older leaves. On older leaves the larval group size above which mortality increased no further was over two-fold that on young leaves. Observations of group feeding behaviour at each instar showed that the majority of larvae (75.7%) were aligned facing away from the feeding site and that only around 7.5%, or just 1–2 larvae per group, fed at any one time. Feeding larvae chewed the leaf edge by straddling the leaf margin. Measurements of leaf margins showed that older leaves had significantly thicker leaf margins and 'thickness' ratios (leaf margin to leaf lamina proper). In the field, approximately 85% of all larvae occurred on the first two expanded leaf pairs, and larval mortality was highest between eclosion and establishment of the first instar. However, beetles apparently did not adjust clutch size according to leaf age.     

Effect of group size on parasitism in a natural population of the Baltimore checkerspot Euphydryas phaeton

Summary The effect of group size of early instars on parasitism of Euphydryas phaeton (Nymphalidae) was examined. Different numbers of larvae were stocked per web to determine the effect of group size on parasitism. Larval aggregations of moderate size (the size occurring naturally) had the least parasitism. Larger larval groups had a disproportionately high rate of parasitism. The major larval parasitoids located vulnerable larvae within webs, instead of attacking larvae available on the outside of webs. Parasitism rates were similar for larvae of damaged and undamaged webs, a consequence of the behavior and location of larvae in the webs. Lower limit to group size was a function of facilitation of larval numbers in reaching the first feeding site, the top of the host plant. Feeding facilitation by larval aggregations was not a factor in larval survival or growth.     

Rainfall, nitrogen and host plant condition: consequences for the processionary caterpillar, Ochrogaster lunifer

1. At the end of November in subtropical areas of Australia, second-instar larvae of the processionary caterpillar Ochrogaster lunifer (Lepidoptera: Thaumetopoeidae) initiate feeding on the peripheral shoots of acacias (first-instar larvae do not feed). Field surveys at ten localities in south-east Queensland showed that larval survival was highly variable both among and within localities. Within-locality variation in larval growth was low compared with variation among localities. Larval growth and survival rates were higher at coastal and island localities, where November rainfall was high, than at drier inland localities.
2. Potted Acacia concurrens were grown in the greenhouse under high and low watering regimes, with and without nitrogen-rich fertilizer. Plant vigour (height, foliar water content and quantity of flush growth) was significantly greater in high-water treatments than in low-water treatments. Watering also affected foliar nitrogen, with plants in the high-water/no fertilizer treatment having similar nitrogen levels to those in fertilized treatments. Fertilizer increased foliar nitrogen levels of plants in low-water treatments and increased the number of shoots in high-water treatments. Different treatments had no effect on leaf toughness. After the first 3 weeks of feeding, size and survival of larvae were significantly reduced on the small, less vigorous plants in low water treatments. These results do not support the plant stress hypothesis.
3. Early-instar larvae (instar II–IV) developed more quickly and grew larger when reared on flush leaves than when reared on senescent leaves of A. concurrens . As water uptake affects the quantity of flush growth available to early stage larvae as well as foliar quality, rainfall and water availability may have important consequences for the distribution and population dynamics of the moth at local and regional scales.     

Understanding gregariousness in a larval Lepidopteran: the roles of host plant, predation, and microclimate

Abstract.  1. Many moth and butterfly larvae are gregarious early in development, but become solitary in late instars. This ontogenetic variation in behaviour is probably the result of temporal changes in the costs and benefits associated with gregariousness. This study provides observational and experimental evidence that, in one particular moth species, a series of different ecological factors influence larval behaviour at different times during development.
2. Field observations show that young caterpillars of the limocodid Doratifera casta form large aggregations while foraging, but that mature larvae are largely solitary.
3. A field experiment revealed that individual first to third instar larvae in larger groups develop more rapidly, but that group size had no detectable influence on survival. The developmental advantage associated with gregariousness is affected by host plant species, but not by predator exclusion, suggesting that group living in these cryptic early instar larvae promotes feeding facilitation, but does not provide individuals with protection from natural enemies.
4. Laboratory experiments revealed that aposematic fourth instar caterpillars in large groups were less likely to be attacked by a generalist insect predator than those in small groups.
5. Field observations provided no evidence that group living affects body temperature, suggesting that microclimatic factors do not favour gregariousness in this species.
6. It is concluded that gregariousness in D. casta confers at least two different advantages on larvae at different stages early in development, but that these advantages disappear, or are outweighed by costs associated with intraspecific competition, in final instars.     

Enemy‐free space and the host range of a lichenivorous moth: a field experiment

According to the enemy‐free space hypothesis (EFS), parasites and predators create a selective force for a specialization on a host that assures better protection against natural enemies than other potential hosts. Few studies have found support for EFS and none of them have covered the whole larval period in natural conditions. I studied the growth and survival of lichen‐feeding moth larvae on five lichen species with and without their natural enemies in natural conditions covering the whole larval period. All the three following EFS predictions gained support. First, natural enemies caused significant mortality of larvae. Second, when natural enemies were present, larval survival was highest on preferred Ramalina lichens. Third, larvae attained higher mass on non‐preferred Parmelia sulcata than on Ramalina species, indicating fecundity cost to feed on Ramalina species instead of P. sulcata. EFS for C. lichenaria larvae on Ramalina species is likely a consequence of shrubby appearance of Ramalina species which provide better larval protection from predation than other hosts.     

Does group feeding by toxic prey confer a defensive benefit? Aristolochic acid content,group size and survival of first‐instar pipevine swallowtail (Battus philenor L.) larvae

1. Aggregative feeding of larvae is widespread in the Lepidoptera, and many hypotheses have been proposed to explain the adaptive significance of this trait. Group feeding occurs disproportionately more in species with aposematic, chemically defended larvae compared with species with cryptic, non‐chemically defended larvae, consistent with the hypothesis that group feeding provides an enhanced aposematic signal to natural enemies. Most species characterised as having chemically defended larvae are cryptic during the first instar, when they are most highly aggregated and most vulnerable to predation. 2. The benefits of group feeding in terms of decreased predation were explored for first‐instar larvae of the pipevine swallowtail, Battus philenor, a species that sequesters aristolochic acids from its Aristolochia host plant and exhibits aposematism in later instars and as adults. We found that groups of larvae with experimentally enhanced aristolochic acid content had significantly lower survivorship due to predation both in the field and in laboratory experiments compared with groups of larvae without enhanced chemical defence. 3. A laboratory experiment found that aristolochic acid does not deter the generalist predator Hippodamia convergens. 4. No evidence was found that was consistent with the hypothesis that group feeding and increased sequestered chemical defence interact to decrease rates of larval mortality in non‐aposematic, first‐instar larvae. Future research on chemical defence, aposematism, and aggregative feeding should continue to appreciate that particular chemical defences and feeding behaviours are not universally effective against all natural enemies.     

Allee effect in larval resource exploitation in Drosophila: an interaction among density of adults, larvae, and micro-organisms

Abstract 1. Aggregation pheromones can evolve when individuals benefit from clustering. Such a situation can arise with an Allee effect, i.e. a positive relationship between individual fitness and density of conspecifics. Aggregation pheromone in Drosophila induces aggregated oviposition. The aim of the work reported here was to identify an Allee effect in the larval resource exploitation by Drosophila melanogaster, which could explain the evolution of aggregation pheromone in this species. 2. It is hypothesised that an Allee effect in D. melanogaster larvae arises from an increased efficiency of a group of larvae to temper fungal growth on their feeding substrate. To test this hypothesis, standard apple substrates were infested with specified numbers of larvae, and their survival and development were monitored. A potential beneficial effect of the presence of adult flies was also investigated by incubating a varying number of adults on the substrate before introducing the larvae. Adults inoculate substrates with yeast, on which the larvae feed. 3. Fungal growth was related negatively to larval survival and the size of the emerging flies. Although the fungal growth on the substrate was largely reduced at increased larval densities, the measurements of fitness components indicated no Allee effect between larval densities and larval fitness, but rather indicated larval competition. 4. In contrast, increased adult densities on the substrates prior to larval development yielded higher survival of the larvae, larger emerging flies, and also reduced fungal growth on the substrates. Hence, adults enhanced the quality of the larval substrate and significant benefits of aggregated oviposition in fruit flies were shown. Experiments with synthetic pheromone indicated that the aggregation pheromone itself did not contribute directly to the quality of the larval resource. 5. The interaction among adults, micro‐organisms, and larval growth is discussed in relation to the consequences for total fitness.     

Selection of large host plants for oviposition by a monophagous leaf beetle: nutritional quality or enemy‐free space?

Abstract.  1. Oviposition site selection is crucial for the reproductive success of herbivorous insects. According to the preference–performance hypothesis, females should oviposit on host plants that enhance the performance of their offspring. More specifically, the plant vigour hypothesis predicts that females should prefer large and vigorously growing host plants for oviposition and that larvae should perform best on these plants.
2. The present study examined whether females of the monophagous leaf beetle Cassida canaliculata Laich. (Coleoptera: Chrysomelidae) prefer to oviposit on large host plant individuals of the meadow clary and whether large host plants are of higher nutritional quality than small host plants. Subsequently, it was tested whether the female preference correlates with offspring performance and survival.
3. In the field, females preferred large host plant individuals for oviposition and host plant quality, i.e. leaf nitrogen content, was significantly higher in leaves of large than of small host plants.
4. In the laboratory, larval development time was shorter on leaves of large host plant individuals than on small host plant individuals, but this could not be shown in the field.
5. However, a predator-exclusion experiment in the field resulted in a higher survival of larvae on large host plants than on small host plants when all predators had free access to the plants. On caged host plants there was no difference in survival of larvae between plant size categories.
6. It is concluded that females of C. canaliculata select oviposition sites that enhance both performance and survival of their offspring, which meets the predictions of the plant vigour hypothesis.     

Group feeding as a strategy for exploiting food resources in the burnet moth Pryeria sinica

Summary Pryeria sinica (Zygaenidae) larvae feed on young and growing leaves of Euonymus japonicus in groups. The larvae often defoliate their host plant. Hence, the larvae are occasionally subject to serious shortage in food resources. We hypothesize that larval aggregation is an adaptation for the economical utilization of limited food resources. To test this hypothesis, the patterns of resource utilization were studied on larvae settled on shoots of host plant in various group sizes. The amount of food resources in a shoot was affected by the following three factors; (1) shoot growth, (2) food consumption of larvae, and (3) inhibition of growth or degeneration of shoot by larval activity on it. These factors were measured and the efficiency in resource utilization was compared among groups of varied sizes. The loss in resources caused by the third factor was found to be decreased with group size, i.e., the larger the group size, the larvae utilized the food resources more economically. The advantages of group feeding in survival and reproduction of this species were discussed using a simple graphical model.     

Do exotic generalist predators alter host plant preference of a native willow beetle?

1 Selection can favour herbivores that choose host plants benefitting their offspring either by enhancing growth rates or by increasing larval defences against native predators. For exotic predator species that feed on herbivores, their success with invading new habitats may depend upon overcoming defences used by native prey. Whether exotic predators can alter herbivore host choice has remained unexamined. Therefore, we compared the efficacy of larval defence by Chrysomela knabi (a native beetle species) that had fed on two native willow hosts: Salix sericea (a phenolic glycoside (PG)-rich species) and Salix eriocephala (a PG-poor species), when attacked by exotic generalist predators. In addition, the preference and performance of C. knabi on S. sericea and S. eriocephala was examined.
2  Chrysomela knabi preferred and performed better on S. sericea. In a common garden, adult C. knabi were nine-fold more common and oviposited five-fold more frequently on S. sericea than on S. eriocephala . In the laboratory, adult feeding preference on leaf discs and survival rates of larvae were both greater on S. sericea , and time to pupation was shorter.
3  Chrysomela knabi larvae produced significantly more salicylaldehyde when fed S. sericea leaves than when fed S. eriocephala leaves. Additionally, those larvae with greater salicylaldehyde had reduced predation by two exotic generalist predators, Harmonia axyridis larvae and juvenile Tenodera aridifolia sinensis .
4 The results obtained in the present study suggest that selection favoured the preference of C. knabi for PG-rich willow plants because larvae grew and survived better and that selection by common exotic generalist predators would reinforce this preference.     

A recently acquired host plant provides an oligophagous insect herbivore with enemy-free space

Enemy-free space (EFS) is a potentially important factor affecting host plant use by phytophagous insects. Yet only a few field studies have demonstrated that natural enemy activity is the sole mechanism underlying use of novel host plants by herbivorous insects. This may be due to the fact that in earlier studies, both herbivores and natural enemies had the opportunity to adapt to the new host plant. Here we studied the possibility that EFS underlies the recently recorded increase in Phthorimaea operculella densities on tomato plants in a few areas within its geographical range. Through field experiments in Ethiopia, we show that all three conditions proposed by Berdegue et al. to demonstrate EFS are fulfilled. First, a significantly higher proportion of larvae survive on caged than on exposed potato plants, showing that natural enemies are an important mortality factor on the original host, potato. Second, larval survival was significantly higher on exposed tomato than potato plants, implying greater protection for the herbivore from its natural enemies on tomato than on potato plants. Thus tomato plants provide P. operculella with an EFS. Finally, larval survival was significantly higher on caged potato than on caged tomato plants at the preblossom stage, indicating that, in the absence of natural enemies, there is a fitness cost when larvae feed on the sub-optimal tomato plants. Fulfillment of this third condition points to the importance of natural enemy activity relative to that of other unidentified factors, such as food quality and competition. An intensive field survey provides further support for this conclusion.     

Effects of natural enemies on relative fitness of Heliothis virescens genotypes adapted and not adapted to resistant host plants

We investigated the potential of two natural enemies of Heliothis virescens (F.) (Noctuidae) to affect its rate of adaptation to tobacco containing Bacillus thuringiensis Berliner toxin. Larval fitness of two laboratory strains of H. virescens, one adapted to B. thuringiensis toxin and one not adapted, was compared on toxic and nontoxic plants, in the presence of the parasitoid Campoletis sonorensis (Cameron) (Ichneumonidae) or the entomopathogenic fungus Nomuraea rileyi (Farlow) Samson. By exposing larvae to plants and enemies for no more than 24 h, we focussed on the behavioral rather than physiological component of their interaction with toxic plants and natural enemies. Parasitism of H. virescens larvae by C. sonorensis during exposure periods of 1–4 h was lower on toxic plants than nontoxic plants and was lower for nonadapted larvae than for toxin-adapted larvae. Decreased larval feeding damage on toxic versus nontoxic plants, and by nonadapted versus adapted larvae, may explain differences in parasitism, because C. sonorensis locates host larvae using cues from damaged plants. Effects of plant toxicity and larval strain on H. virescens survival were numerically consistent with effects on parasitism, but they were not statistically significant. When mean larval survival is used to estimate fitness of the nonadapted genotype relative to the toxin-adapted genotype, we find that C. sonorensis is expected to delay adaptation to toxic plants. Percent infection by N. rileyi of H. virescens larvae exposed to fungus-treated plants for 24 h was greater when plants were toxic, and was greater for nonadapted larvae than toxin-adapted larvae. There were corresponding decreases in larval survival on toxic compared to nontoxic plants, and of nonadapted compared to adapted larvae. Interaction of effects of plant line and larval strain on survival was significant in the presence of fungus, but not in the absence of fungus, which indicates that the effect of toxic plants on the relative fitness of toxin-adapted and nonadapted larvae was mediated by fungus. As in the interaction with C. sonorensis, behavior of larvae on plants may explain differences in susceptibility to N. rileyi. Because nonadapted larvae moved more than toxin-adapted larvae on toxic plants, nonadapted larvae may have been more likely to encounter a lethal dose of conidia. In contrast with C. sonorensis, N. rileyi, which decreased the fitness of the nonadapted genotype relative to the adapted genotype, is expected to accelerate adaptation to toxic plants.     

Communication of leaf suitability by gregarious eastern tent caterpillars (Malacosoma americanum)

ABSTRACT. 1. Previous work has shown that leaf age affects recruitment trail marking by eastern tent caterpillars ( Malacosoma americanum Fabr.). Young leaves of host plants elicit trail marking to a greater degree than mature leaves.
2. Experiments were conducted to establish the relationship between the differential behavioural responses of larvae to young and mature leaves and the suitability of foliage for larval growth and survival. Foliage of black cherry ( Prunus serotina Ehrh.), a typical rosaceous host plant, was used for this comparison.
3. Larvae preferred young leaves to mature leaves in choice tests, and marked more to young leaves than to mature leaves in no-choice tests.
4. Mature leaves supported adequate growth through two larval instars of rearing, but thereafter were unsuitable for growth. Larvae fed mature leaves had lower pupal weight, poorer survival, and grew less efficiently than larvae fed young leaves.
5. The results support the hypothesis that the trail communication system of eastern tent caterpillars is an adaptation to efficiently locate leaves which are favourable for larval growth and survival.     

Slow larval growth on a suboptimal willow results in high predation mortality in the leaf beetle Galerucella lineola

The slow growth/high mortality hypothesis predicts that herbivorous insects feeding on suboptimal host plants are subjected to higher predation mortality owing to the longer time spent in the vulnerable juvenile stages compared with conspecifics feeding on optimal plants. We tested this hypothesis for the willow-feeding leaf beetle Galerucella lineola raised on one suitable (Salix viminalis) and one unsuitable (S. dasyclados) willow species as well as on plants from an interspecific cross between the two species. Cohorts of larvae raised on caged plants (protected from enemies) and uncaged plants (exposed to enemies) were monitored daily throughout larval development in two consecutive years. Larvae raised on S. viminalis developed faster, grew larger and survived better than those raised on S. dasyclados. The suitability of the hybrid plants was intermediate to that of their parents. Our results strongly support the slow growth/high mortality hypothesis. In both years, total predation during the larval period was higher on S. dasyclados than on S. viminalis. Furthermore, the daily predation rate (i.e. the proportion of larvae preyed upon per day) was higher on S. dasyclados than on S. viminalis. When hybrid plants were included in the analysis total predation was positively correlated with both larval development time and daily predation rate. We suggest that high predation on beetles on low-quality plants is the combined result of their longer development time and elevated daily predation rate. The results are discussed in relation to the evolution of host plant selection and the paradox of sublethal plant defenses.     

Asymmetric resource competition in a filter-feeding stream insect (Hydropsyche siltalai; Trichoptera)

SUMMARY. 1. Field experiments in a lake outlet stream were carried out on filter-feeding larvae of the caddisfly Hydropsyche siltalai Döhler, to assess whether microhabitat quality, establishment and growth rate is affected by position (upstream—downstream) within a dense aggregation of larvae.
2. Artificial substrates holding high-density aggregations of hydropsychid larvae were used. Marked III-instar H. siltalai larvae were implanted either in the upstream end or in the downstream end of an aggregation and recovered after 17 days.
3. Larvae recovered from the downstream end of an aggregation were smaller than those that had been implanted in the upstream end.
4. Fewer larvae had settled in the downstream end of aggregations compared to the exposed upstream part.
5. Naturally colonizing H. sitalai larvae were, in contrast to implanted larvae, more numerous behind aggregations. Small (first and second instar) Hydropsyche angustipennis (Curtis) and/or H. pellucidula (Curtis) larvae were more numerous in front of aggregations.
6. Current velocities, measured with a microcurrent velocity meter, and density of drifting prey, were significantly lower behind aggregations of hydropsychid larvae. Food availability decreased by about 50% behind aggregations.