The Role of Tactile and Chemical Stimuli in the Formation and Maintenance of the Processions of the Social Caterpillar Hylesia lineata (Lepidoptera: Saturniidae)

Colonies of the social caterpillar Hylesia lineata (Lepidoptera: Satumiidae) form long, single-file, head-to-tail processions as they move between their shelters and distant feeding sites. Although investigations of other processionary species have implicated a silk trail in the processionary process, silk plays little or no role in initiating or maintaining processions in H. lineata. Studies we report here implicate both tactile stimuli and a trail pheromone in the establishment and maintenance of processions. Processionaries elicit locomotion in the individual preceding them in line by brushing their heads against prominent sulci that project from the tips of their abdomens. Caterpillars mark their pathways with a pheromone deposited by brushing the ventral surfaces of their last abdominal segments against the substrate. The persistent pheromone is soluble in hexanes and appears to be secreted from glandular setae found on the proximal regions of the anal prolegs and the venter. In Y-choice tests, caterpillars selected newer trails over older trails and stronger trails over weaker trails. They did not distinguish between trials deposited by newly fed caterpillars and those deposited by starved caterpillars. Despite the unidirectional nature of processions, there is no indication that caterpillars can determine from the trail alone the direction in which the procession advanced. The significance of these findings to the foraging ecology of the caterpillars is discussed.     

Testing the host‐finding ability of a monophagous caterpillar in the field

1. Relatively few studies of the host‐finding ability of specialised, phytophagous insects involve direct observations of individual insects moving among intact hosts and non‐hosts. Information from such studies can inform the design of restoration programmes for species of conservation concern. 2. The movement of caterpillars of the threatened Oregon silverspot butterfly, Speyeria zerene hippolyta (Edwards) (Lepidoptera: Nymphalidae) was studied in the field in cleared arenas with 10 cm radii. Caterpillars were placed in the centre, surrounded by three individuals of their host, Viola adunca, and three different non‐host individuals, separated by bare ground. In a second experiment, second instars were placed between a host and a non‐host, 3–6 cm away. Caterpillars were observed to determine if they walked to their host more often than expected by chance. 3. Caterpillars walked to vegetation significantly more often than expected by chance. They did not, however, reach their hosts more often than expected, based on plant availability. 4. It is concluded that S. z. hippolyta caterpillars can distinguish vegetation from bare ground from 10 cm away. There is no evidence that they can distinguish their host plant from other herbaceous species at distances of 3 cm.     

Spatial orientation of social caterpillars is influenced by polarized light

Processionary caterpillars of Thaumetopoea pityocampa (in Europe) and Ochrogaster lunifer (in Australia) (Lepidoptera: Notodontidae) form single files of larvae crawling head-to-tail when moving to feeding and pupation sites. We investigated if the processions are guided by polarization vision. The heading orientation of processions could be manipulated with linear polarizing filters held above the leading caterpillar. Exposure to changes in the angle of polarization around the caterpillars resulted in corresponding changes in heading angles. Anatomical analysis indicated specializations for polarization vision of stemma I in both species. Stemma I has a rhabdom with orthogonal and aligned microvilli, and an opaque and rugged surface, which are optimizations for skylight polarization vision, similar to the dorsal rim of adult insects. Stemmata II-VI have a smooth and shiny surface and lobed rhabdoms with non-orthogonal and non-aligned microvilli; they are thus optimized for general vision with minimal polarization sensitivity. Behavioural and anatomical evidence reveal that polarized light cues are important for larval orientation and can be robustly detected with a simple visual system.     

Acoustic stimuli from predators trigger behavioural responses in aggregate caterpillars

Prey detect their predators through predator signals and cues and, consequently, respond with anti‐predatory behaviours to inhibit the action of their aggressors. Lepidopterans can intercept signals emitted by predators and may defend themselves through chemical, morphological or behavioural responses. In this study, we investigated the effect of acoustic stimuli of different predators on defensive behaviour of gregarious caterpillars. Our results demonstrated that Hylesia nigricans (Lepidoptera, Saturniidae) caterpillars alter their behaviour (i.e. abruptly raising the head) in response to the acoustic stimulus of the predators (i.e. predation risk signals from birds and wasps). The magnitude of this response depended on predator identity and caterpillar body size. Larger caterpillars responded more strongly to predatory stimuli than smaller caterpillars. However, regardless of the size of the caterpillars, they responded more strongly to the stimuli of wasps. In addition, we identified that H. nigricans caterpillars emit ultrasonic noise after detecting the stimuli of the predators – this noise seems to function as an alert about the risk of predation during the early stages of development (second and fifth instars). The duration of ultrasonic emission (i.e. milliseconds) increases with the number of repetitions of the stimuli (i.e. wing‐beat sounds of the wasps and insectivorous birds). These results provide novel information about predation risk in interactions among caterpillars and their predators, and indicate possible communication among invertebrates mediated by the risk of predation.     

Silk drives aggregation and following in the neotropical caterpillar Mechanitis menapis (Nymphalidae: Ithomiini)

Gregarious larvae that use chemical communication to feed and move together are widespread among folivorous insects, although social behaviour has been studied almost exclusively in a few temperate zone genera. The Menapis (or variable) tigerwing butterfly Mechanitis menapis mantineus Hewitson (Lepidoptera, Nymphalidae, Danainae, Ithomiini) is a neotropical species whose larvae feed gregariously on Solanaceae host plants. In laboratory experiments conducted in the Ecuador cloud forest, M. menapis caterpillars are attracted to silk produced by conspecifics and show no evidence of pheromone production. Indeed, caterpillars consistently choose arenas with silk over bare arenas but do not show a preference for arenas marked with abdominal cuticular surface residues. Mechanitis menapis caterpillars on silk‐coated plants are both more mobile and more cohesive than those on control plants. Nonetheless, caterpillars move independently over unmarked surfaces and groups do not make rapid collective choices between two food sources. Collective behaviour in M. menapis thus appears to be based on aggregation on collectively produced silk to facilitate feeding, as well as using this silk to maintain cohesion. Silk production is common in caterpillars, although M. menapis appears to be unique among species studied so far in using silk to maintain group cohesion.     

Life on a deadly trap: Buckleria paludum,a specialist herbivore of carnivorous sundew plants,licks mucilage from glands for defense

Caterpillars of Buckleria spp. (Lepidoptera: Pterophoridae) have a unique feeding habit of eating trap leaves of carnivorous sundew plants (Drosera spp.). We observed the foraging behavior of Buckleria paludum on trap leaves of Drosera spp. and discussed how the moth species avoided being caught by trap leaves. In 81.5% (66/81) of encounters with glandular hairs on adaxial surfaces of Drosera trap leaves, B. paludum larvae licked mucilage and crawled on the processed hairs. The frequency of licking mucilage was significantly higher than the frequency of other behaviors such as eating glandular hairs, chewing bases of them without eating and ignoring when encountering secreted mucilage. Licking mucilage enables the caterpillars to move safely on trap leaves and prevents bending of glandular hairs.     

Diet mixing enhances the performance of a generalist caterpillar,Platyprepia virginalis

1. Platyprepia virginalis caterpillars are dietary generalists and feed on multiple host species within a single day. We conducted field experiments to evaluate their performance on diets consisting of only their primary food, Lupinus arboreus, or diets consisting of L. arboreus plus other acceptable host species. 2. We found that relative growth rates and rates of survival were higher when they fed on mixed diets compared to lupine only. These results were consistent with hypotheses that mixed diets provided balanced nutrition, diluted toxins, and/or allowed recovery from parasitoids, although our data did not allow us to separate these non‐exclusive explanations. 3. We assayed alkaloids in their host foliage, in the caterpillars themselves, in parasitoids within caterpillars, in food boluses passing through their guts, and in frass that they excreted. We consistently found positive assays for alkaloids in foliage and in frass but negative assays in caterpillars, parasitoids, and food boluses. This suggests that the alkaloids that they ingest are metabolised or rendered non‐reactive by unknown means during passage through the gut. We found no support for the hypothesis that mixed diets prevented caterpillars from exhausting food supplies or allowed them to sequester chemicals from their alkaloid‐containing hosts. 4. Behavioural observations revealed that previous experience influenced a caterpillar's likelihood of moving to a different host. Caterpillars that had previously fed on other hosts were more likely to move to lupine while caterpillars previously collected on lupine were equally likely to choose more lupine or a different host. 5. These results are unusual in providing a clear and consistent benefit of diet mixing in a natural field setting where multiple ecological factors act upon the caterpillars.     

Behavioural ecology of defence in a risky environment: caterpillars versus ants in a Neotropical savanna

1. Predatory ants may reduce infestation by herbivorous insects, and slow‐moving Lepidopteran larvae are often vulnerable on foliage. We investigate whether caterpillars with morphological or behavioural defences have decreased risk of falling prey to ants, and if defence traits mediate host plant use in ant‐rich cerrado savanna. 2. Caterpillars were surveyed in four cerrado localities in southeast Brazil (70–460 km apart). The efficacy of caterpillar defensive traits against predation by two common ant species (Camponotus crassus, C. renggeri) was assessed through experimental trials using caterpillars of different species and captive ant colonies. 3. Although ant presence can reduce caterpillar infestation, the ants' predatory effects depend on caterpillar defence traits. Shelter construction and morphological defences can prevent ant attacks (primary defence), but once exposed or discovered by ants, caterpillars rely on their size and/or behaviour to survive (secondary defence). 4. Defence efficiency depends on ant identity: C. renggeri was more aggressive and lethal to caterpillars than C. crassus. Caterpillars without morphological defences or inside open shelters were found on plants with decreased ant numbers. No unsheltered caterpillar was found on plants with extrafloral nectaries (EFNs). Caterpillars using EFN‐bearing plants lived in closed shelters or presented morphological defences (hairs, spines), and were less frequently attacked by ants during trials. 5. The efficiency of defences against ants is thus crucial for caterpillar survival and determines host plant use by lepidopterans in cerrado. Our study highlights the effect of EFN‐mediated ant‐plant interactions on host plant use by insect herbivores, emphasizing the importance of a tritrophic viewpoint in risky environments.     

Foliage‐age mixing within balsam fir increases the fitness of a generalist caterpillar

1. Manipulative field studies were carried out to evaluate the foliage age preference–performance relationship for an extreme generalist herbivore, the whitemarked tussock moth (Orygia leucostigma Smith) (Lepidoptera: Lymantriidae), within balsam fir [Abies balsamea (L.) Mill]. 2. Field surveys indicated that early instar caterpillars fed almost exclusively on young (i.e. current‐year) foliage, whereas late instars caterpillars fed on both young and mature (i.e. 1‐ and 2‐year‐old) foliage. 3. Survival of early instar caterpillars was highest in treatments where current‐year and/or 1‐year old foliage were available, but decreased significantly on older foliage. In contrast, late instar caterpillars had the highest survival when allowed to feed on all age classes of foliage, whereas potential fecundity was highest for late instars that fed on young foliage. 4. Overall, caterpillars had 32–65% higher fitness when able to feed on all rather than just one age class of foliage. 5. These results support both the ‘complementary diet' hypothesis, which states that dietary mixing of different‐aged foliage can increase nutrient uptake and/or dilute harmful secondary plant chemicals, and the ‘ontogeny’ hypothesis, which attributes changes in diet to changes in the nutritional needs and/or tolerance to plant defences of juvenile insects as they develop.     

Response of Brassica oleracea to temporal variation in attack by two herbivores affects preference and performance of a third herbivore

1. Plants are frequently under attack by multiple insect herbivores, which may interact indirectly through herbivore‐induced changes in the plant's phenotype. The identity, order, and timing of herbivore arrivals may influence the outcome of interactions between two herbivores. How these aspects affect, in turn, subsequently arriving herbivores that feed on double herbivore‐induced plants has not been widely investigated. 2. This study tested whether the order and timing of arrival of two inducing herbivores from different feeding guilds affected the preference and performance of a subsequently arriving third herbivore, caterpillars of Mamestra brassicae L. (Lepidoptera: Noctuidae). Aphids [Brevicoryne brassicae L. (Hemiptera: Aphididae)] and caterpillars [Plutella xylostella L. (Lepidoptera: Yponomeutidae)] were introduced onto wild Brassica oleracea L. (Brassicaceae) plants in different sequences and with different arrival times. The effects of these plant treatments on M. brassicae caterpillars were assessed in pairwise preference tests and no‐choice performance tests. 3. The caterpillars of M. brassicae preferred to feed from undamaged plants rather than double herbivore‐induced plants. Compared with undamaged plants, they preferred plant material on which aphids had arrived first followed by caterpillars, whereas they avoided plant material with the reverse order of herbivore arrival. Performance of the caterpillars increased with increasing arrival time between herbivore infestations in double herbivore‐induced plants. Although M. brassicae grew faster on plants induced by aphids than on those induced by caterpillars alone, its performance was not affected by the order of previous herbivore arrival. 4. These results imply that the timing of colonisation by multiple herbivores determines the outcome of plant‐mediated herbivore–herbivore interactions.     

Effects of trichomes on the behavior and distribution of Platyprepia virginalis caterpillars

Trichomes are an important physical resistance mechanism of plants, as they reduce insect herbivore movement, feeding, and digestion. However, we know little about how trichomes influence herbivore distributions and populations. We conducted laboratory and field experiments to evaluate the preferences of Platyprepia virginalis (Boisduval) (Lepidoptera: Arctiidae) caterpillars to natural and manipulated densities of trichomes on their primary food, Lupinus arboreus Sims (Fabaceae). We then conducted field surveys to determine whether variation in trichome density among lupine bushes affected caterpillar spatial distribution on the landscape. Platyprepia virginalis caterpillars preferred lupine leaves with fewer trichomes in choice and no‐choice experiments. In the field, caterpillar feeding damage was found more often on leaves with fewer trichomes. These preferences scaled up to the level of bushes in the landscape such that more caterpillars were found on bushes with lower trichome densities than on bushes with higher trichome densities. This is one of few studies to show the potential for trichome density to influence herbivore population size and distribution in a natural system at a landscape level. The results are consistent with trichomes functioning as a resistance mechanism with consequences for herbivore choice, performance, and distribution.     

Where to from here? The mechanisms enabling the movement of first instar caterpillars on whole plants using Helicoverpa armigera (Hübner)

Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae) is an economically-important, polyphagous herbivore in Old World countries. The distribution of larvae within various host plants has been described, but few studies have tried to determine the behavioural mechanisms by which the given distributions arose. Our aim was to determine the mechanisms which enable larval movement on pea plants, starting with first instars. Observations and bioassays determined larval movement in response to light and angled surfaces, as well as the effect of feeding and plant volatiles on these responses. The majority (68–72%) of 1st instars were positively phototactic towards blue, green and white light and 42% towards UV light. In the dark, larvae showed negative geotaxis. The angle of their substrate also had a kinetic effect on larvae; the steeper the angle from horizontal the more larvae moved under all conditions. Phenylacetaldehyde (a flower volatile) suppressed larval movement except at 90°. (Z)-3-Hexenyl acetate (a green leaf volatile) reversed the direction of movement at the flattest angle. Feeding lessened the probability of moving. We suggest that phototaxis and geotaxis are behaviours common to larval lepidopterans (caterpillars), and that these basic behaviours are modulated by environmental, larval, and plant factors to give observed distributions. Using a multinomial model approach, we created a flow chart to qualitatively and quantitatively represent the decision-making process of first instar H. armigera in response to the factors influencing movement.     

Lack of susceptibility of soil‐inhabiting Platyprepia virginalis caterpillars,a native arctiid,to entomopathogenic nematodes in nature

Entomopathogenic nematodes (EPNs) can kill and regulate populations of soil‐inhabiting insects, but studies evaluating these interactions in native ecosystems are rare. The objective of this study was to examine the effects of EPNs on a non‐agricultural caterpillar, Platyprepia virginalis (Boisduval) (Lepidoptera: Arctiidae), under natural conditions. Platyprepia virginalis caterpillars live in litter on the soil surface feeding beneath bush lupine during summer, autumn, and winter. Initial laboratory assays revealed that the caterpillars were vulnerable to at least two species of EPNs with which they co‐occur in the coastal prairie in northern California (USA). In contrast to laboratory assays, caterpillars survived exposure to prairie soil containing EPNs under natural conditions in field assays. To better understand the divergence between laboratory and field results for this native caterpillar, we used sentinel insects [Galleria mellonella L. (Lepidoptera: Pyralidae)] to identify particular locations where EPNs were present in the field. Platyprepia virginalis caterpillars were caged at these sites but again showed no evidence of susceptibility to EPNs. Platyprepia virginalis caterpillars reduce their exposure to EPNs by spending their time in and above the litter rather than contacting the soil when given the choice in nature. We conclude that P. virginalis is unlikely to serve as a reservoir for EPNs and that nematodes are unlikely to be important mortality factors for P. virginalis in this natural system.     

A single lycaenid caterpillar gets an ant‐constructed shelter and uninterrupted ant attendance

Ant‐lycaenid associations range from mutualism to parasitism and the caterpillars of some species of lycaenids are reported to enter ant nests for shelter, diapause, or pupation. The present study aimed to examine the nature of the association between Euchrysops cnejus (Fabricius) (Lepidoptera: Lycaenidae) and Camponotus compressus (Fabricius) (Hymenoptera: Formicidae) worker ants on the extrafloral nectary‐bearing cowpea plant, Vigna unguiculata (L.) Walp. (Fabaceae). The abundance patterns of the ants and the lycaenid caterpillars together with the spatial patrolling patterns of the ants on the plants revealed that ant abundance increased with the occurrence of the lycaenid caterpillars and the ants preferred the lycaenids over the extrafloral nectar. Camponotus compressus worker ants constructed a shelter at the cowpea plant base after interaction with one or more lycaenid caterpillar(s) and tended the caterpillars and pupae till the emergence of the butterfly. The ant‐constructed shelters (ACSs) inhabited by the minor caste workers (13 ± 1.3 ants per ACS), were utilized by the caterpillars to undergo pupation. The ants confined their activities predominantly to tending the pod‐feeding caterpillars and the solitary pupa within each ACS. It appears that the behavior of the tending worker ants is modulated by the lycaenid vulnerable stages.     

Influence of chorion ingestion on the performance of Ascia monuste and its association with cannibalism

1. In some lepidopterans, the newly hatched caterpillars feed on chorion (animal protein) as their first food. This is also a frequent behaviour of newly hatched caterpillars of Ascia monuste. 2. According to some parameters tested (time for pupation, number of adults, male imago weight, and fifth‐instar ingestion), chorion ingestion by first‐instar larvae affects adult performance positively. The ingestion of ultraviolet‐sterilised chorion provided the same positive effect on performance. It is thus suggested that young caterpillars may be benefiting from chorion nutritionally, and that chorion ingestion is a chain of events that leads to positive effects on insect performance. 3. Cannibalism in A. monuste was observed in newly hatched caterpillars and is related to the chorion ingestion behaviour. A condition for this to occur was the interval of time of hatching, which means that, if a group of caterpillars hatches very much before another group, once the caterpillars have ingested the chorion of their own eggs, there is a tendency for them to ingest the chorion of other eggs (including unhatched eggs) and, consequently, practice cannibalism. 4. Ascia monuste immatures are considered to be herbivorous, however it is important to know that they eat animal tissue (chorion and conspecific eggs).     

Social Behavior of Larvae of the Neotropical Processionary Weevil Phelypera distigma (Boheman) (Coleoptera: Curculionidae: Hyperinae)

Socially gregarious behavior among free‐living leaf‐eating insect larvae occurs mostly among Lepidoptera, Symphyta, and a few Chyrsomelidae (Coleoptera). However, the Neotropical hyperine curculionid Phelypera distigma has also evolved this lifestyle, exhibiting a suite of social behaviors unique among beetles. The larvae are nomadic processionary foragers that punctuate foraging bouts with rosette‐shaped resting formations (cycloalexy). Larvae also vibrate or bob their heads rapidly when moving, especially when in contact with conspecifics, and this suggests acoustic or vibrational communication. In this study we used observational and experimental approaches to investigate the basis of processionary, cycloalexic,and head‐vibration behavior of this species. Larvae used both trail pheromones and thigmotactic signals to organize themselves into head‐to‐tail processionary columns. The trail pheromone, produced from the center of the abdomen, remains active for up to 4 h. Processions are not consistently led by particular individuals, but dynamically change over time and often temporarily break into two or more subprocessions. Subprocessions reunite through use of the trail pheromone. We found no evidence that head‐bobbing generates attraction through substrate‐borne or acoustic signals, but this behavior functions in direct contact to excite group activity. Time‐lapse videography used to analyze cycloalexic group formation showed that larvae transition from feeding in a line along the leaf margin to cycloalexic formations on the upper leaf surface via a coordinated back‐up movement that brings the posterior tip of their abdomens into contact. We identify three phases of cycloalexic formation: line‐up, back‐up, and an adjustment phase. Complete assembly can be achieved in as little as 5 s, but often the two phases establishing the basic rosette lasts 5–10 min, while the adjustment phase slowly tightens the group over a period of up to an hour. Collectively these studies present the first documented case of chemical trail marking in a beetle, and provide insight into a remarkable social‐behavioral repertoire convergent in key respects with the better‐studied social caterpillars and sawflies.     

Multisegmental motor activity in the segmentally restricted gin trap behavior in Manduca sexta pupae

Stimulation of sensory neurons innervating hairs in the gin traps on the abdomen of Manduca sexta pupae evokes a rapid bending of the abdomen that is restricted to one or more of the three articulating posterior segments. However, electrical stimulation of the gin trap sensory nerve in an isolated abdominal nerve cord evokes characteristic motor neuron activity in every abdominal segment. To determine if the segmentally distributed motor activity also occurred in intact animals and how it contributed to the segmentally restricted reflex movement, mechanical stimulation of the sensory hairs in intact animals was used to evoke reflex responses that were recorded as electromyograms synchronized with video recordings of the behavior. Motor activity was monitored during movements to determine if there was activity in many segments when the movement was restricted to one segment. Coordinated muscle activity was evoked throughout the abdomen in response to stimulation of any of the three gin traps, even when movement was restricted to one segment. Differences in the timing of ipsilateral and contralateral motor activity among segments allowed the closing of gin traps to be segmentally restricted. These findings suggest that the neural circuit underlying the gin trap reflex is distributed throughout the abdominal nerve cord. This network generates a complex, yet coordinated, motor pattern with muscular activity in many abdominal segments that produces a localized bending reflex. Accepted: 10 January 1997     

Evolution of the cercal sensory system in a tropical cricket clade (Orthoptera: Grylloidea: Eneopterinae): a phylogenetic approach

The diversity of sensory systems in animals has poorly been explored on a phylogenetic basis at the species level. We addressed this issue using cricket cerci, comprising abdominal appendages covered with touch‐ and air‐sensitive hairs. Scanning electron microscopy measurements and spatial analyses of hair positioning were used to quantify the structural diversity of cercal structures. Eighteen Eneopterinae and two Gryllidae (outgroups) were studied from a phylogenetic perspective. Cerci were revealed to be complex, diverse, and variable between cricket species. Based on maximum likelihood estimations, the ancestral Eneopterinae cercus had a small size, and its hair equipment allowed the use of both air and touch mechanoreception. The evolution of Eneopterinae cerci was mainly unconstrained by the phylogeny; it was rather a punctuated process, involving apical transformations, and was mostly unrelated to environmental patterns. All studied species have enhanced their overall perceptive capacities compared to the ancestor. Most have longer cerci with more and/or longer hairs. Sensory abilities have improved either in the direction of touch or air movement detection, or both, without discarding the potential for any sensory capacity that was already present ancestrally. This pattern is consistent with the hypothesis of an evolutionary trade‐off for sensory performances. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 614–631.     

A bodyguard or a tastier meal? Dying caterpillar indirectly protects parasitoid cocoons by offering alternate prey to a generalist predator

In some parasitic Hymenoptera the dying caterpillars remain attached or close to the parasitoid cocoons. It has been suggested that the caterpillars act as ‘bodyguards’ for the vulnerable cocoons and therefore protect them against predators and/or hyperparasitoids (the ‘usurpation hypothesis’). This hypothesis has been demonstrated in associations where the caterpillars remain active and/or aggressive after parasitism. However, in other associations the caterpillars are so physiologically depleted after parasitism that they are unable to physically defend the cocoons and instead sit atop them in a moribund state. In this study a generalist predator, the spined soldier bug, Podisus maculiventris Say (Hemiptera: Pentatomidae), was provided with cocoons of the gregarious endoparasitoid Cotesia glomerata L. and the solitary endoparasitoid Microplitis mediator Haliday (both Hymenoptera: Braconidae), in turn attended by their hosts, Pieris brassicae L. (Lepidoptera: Pieridae) and Mamestra brassicae L. (Lepidoptera: Noctuidae), respectively. Cotesia glomerata produces broods of up to 40 cocoons and the dying caterpillars sit atop the cocoons where they exhibit little response to physical stimuli. Previous studies reported that dying P. brassicae caterpillars were ineffective bodyguards against two species of hyperparasitoids. In both associations, the dying host caterpillars were significantly preferred as food by P. maculiventris over the parasitoid cocoons. However, in absence of caterpillars, the bugs readily attacked the C. glomerata cocoons. Alternatively, the survival of M. mediator was very low, irrespective of whether a caterpillar was present or not. Caterpillars attacked by M. mediator are several times smaller than those attacked by C. glomerata. Consequently, the predators ran out of food much more quickly in the former and switched from one prey to the other. We show that in some host–parasitoid associations the dying caterpillars provide more visually apparent or nutritionally superior prey, rather than acting as bodyguards.     

Restructuring of Lepidoptera communities by introduced Vespula wasps in a New Zealand beech forest

Introduced social wasps (Vespula vulgaris) reach high densities in some New Zealand beech forests, because honeydew provides an abundant high-energy food source. We manipulated wasp density to estimate an “ecological damage threshold” for large, free-living Lepidoptera larvae. There will be a continuum of ecological damage thresholds for wasp density depending on the prey species or habitat. Experimentally placed small caterpillars had a significantly higher survival rate than large caterpillars, and the survival rate of both groups decreased with increasing wasp density. Spring-occurring caterpillars have a probability of surviving of 0.90–0.95, assuming wasps are the only source of mortality. However, at the peak of the wasp season we predict caterpillars would have virtually no chance (probability of 10⁻⁷⁸ to 10⁻⁴⁰) of surviving to adults. Wasp abundance must be reduced by at least 88% to conserve the more vulnerable species of free-living caterpillars at wasp densities similar to those observed in our study sites. This equates to a damage threshold of 2.7 wasps per Malaise trap per day. It was exceeded for about 5 months of the year in non-poisoned sites. There are currently no biological or chemical control techniques available in New Zealand that will reduce wasp abundance below this damage threshold throughout the year. Our models show that most Lepidoptera with spring caterpillars will be able to persist, but species with caterpillars occurring in the peak wasp season will be eliminated. Received: 5 January 1998 / Accepted: 10 February 1999