Division of labor in post-emergence colonies of the primitively eusocial waspPolistes instabilis de Saussure (Hymenoptera: Vespidae)

Polyethism was quantified in post-emergence colonies of the primitively eusocial wasp,Polistes instabilis, and compared to polyethism in a sympatric advanced eusocial wasp,Polybia occidentalis. Like P.occidentalis, P. instabilis foragers collected food (nectar and prey) and nest materials (wood pulp and water).P. instabilis foragers showed some evidence of specialization with respect to which materials they gathered, but most foragers, divided their effort among food and nest materials, a pattern that is rarely seen inP. occidentalis. In colonies of both species, more foragers collected nectar than any other material; in contrast, most water foraging was performed by one or two workers. Upon returning to the nest,P. instabilis foragers gave up part or all of most nectar, prey, and pulp loads to nestmates, while water was rarely partitioned. Prey loads were most likely to be given up entirely.P. instabilis workers show evidence of conflict over the handling of materials at the nest. The frequency with which workers took portions of nectar loads from forgers was positively correlated with their frequency of aggressive dominant behavior, and with their frequency of taking other foraged materials. Compared to polyethism inP. occidentalis P. instabilis showed less individual specialization on foraging tasks and less partitioning of foraged materials with nestmates, suggesting that these characteristics of polyethism have been modified during the evolution of advanced insect societies.     

Abundance of wasps and prey consumption of paper wasps (Hymenoptera, Vespidae : Polistinae) in Northland, New Zealand

Polistine and vespine wasps were captured in Malaise traps in two fire-modified shrubland habitats of varying canopy height and composition at Lake Ohia, Northland, New Zealand. Prey consumption rates were calculated for the Asian paper wasp (Polistes chinensis antennalis) occupying these two areas of shrubland and a home garden in Whangarei, Northland. The sites were systematically searched for nests and wasp prey determined by intercepting foragers returning to nests. The Asian paper wasp predominated in the Malaise trap samples from the low- growing habitat while the German wasp (Vespula germanica) was more common in the taller vegetation type. The Asian paper wasp was more abundant than the German wasp in the samples in February and early March. Only four Australian paper wasps (Polistes humilis) and no common wasps (Vespula vulgaris) were caught. Asian paper wasps collected an estimated 15 000 prey loads per ha per season from one of the shrubland areas, and 478 000 prey loads per ha per season from the second area. These convert to estimates of 31 and 957 g per ha per season of invertebrate biomass removed by paper wasps from each habitat, respectively. The estimate for the garden site was 79 g per ha per season. Wasp nest densities varied between 20 and 210 nests per hectare. The biomass estimates are similar to average figures calculated for vespine wasps in scrubland and pasture. Both Asian paper wasps and Australian paper wasps preyed mainly on lepidopteran larvae. The cabbage white butterfly (Pieris rapae) was the most commonly collected species. Noctuid species were also well represented. Both male and female Asian paper wasps collected nectar in late March and early April.     

Introduction of a scented carbohydrate resource into the nest increases departure rate in Polybia occidentalis

Social wasps do not possess a sophisticated, signal-based mechanism for recruiting foragers to food resources. Instead, in some species na?ve potential foragers use cues, specifically the scent of a resource obtained from successful foragers, to help locate the resource in the field. Prior studies have focused on the effectiveness of this mechanism on increasing the number of foragers at an artificial resource in the field; the increase is typically modest. Here, we focus on the activity at the nest in Polybia occidentalis, a tropical social wasp, and quantify the magnitude of the effect an influx of a known amount of scented carbohydrate solution added directly to the nest has on activating foragers to leave the nest in search of the resource. Under our experimental conditions, adding a scented 2.0 M sucrose solution to the nest doubled the average rate of departure. No increase occurred when the same amount of water was added as a control. This mass activation of foragers may give colonies of this species a competitive edge by enhancing their ability to rapidly exploit new resources.     

Hunting behaviour,prey capture and ant avoidance in the tropical social waspPolybia sevicea (Hymenoptera: Vespidae)

Summary Polybia sericea (Hymenoptera: Vespidae) prey foraging was studied by following individual foragers as they hunted in the field, by observing how wasps handled prey once they had captured it, and by observing wasps as they returned to the nest with prey. Wasps were most likely to forage for prey between 0700 and 1300 hours and between 1600 and 1700 hours. The prey foraging sequence consisted of the behaviours high flight, search, touch, land, groom, walk, bite and malaxate. Captured small prey were malaxated and carried to the nest. Wasps removed the gut from large prey and dragged the meat up a twig or grass stem. A load of the meat was then bitten off and malaxated; the remainder was cached while the wasp made an orientation flight and returned to the nest. The forager returned within minutes for the remainder of the prey. Experiments demonstrated that caching the prey remains above the ground rather than close to the ground, where the prey are generally captured, reduces the chance that the prey will be found and expropriated by ants.     

Use of terrestrial chemical trails for nest orientation in an open nesting wasp, <Emphasis Type="Italic">Dolichovespula saxonica</Emphasis> F.

Summary The long-cheeked wasp Dolichovespula saxonica typically constructs exposed nests which can be reached by flying. Usually foragers do not walk on substrates in the close vicinity of the nests as cavity breeding wasps do (Steinmetz et al., 2002). Unexpectedly, when forced to walk outside the nest instead of flying in an artificial tunnel system, D. saxonica foragers lay a terrestrial trail and use it for orientation in the nest area in our experiments. 41% of the foragers followed the trail in a direction they were not accustomed to. We suggest that the foragers have employed the same orientation cues normally used for orientation in the close vicinity of the nest when approaching a free-hanging nest by flying, for example nest odour. Nest odour substances may have been transferred to the substrate as a trail as a consequence of foragers walking through the tunnels.Received 23 August 2002; revised 27 January 2003; accepted 30 April 2003.     

Temporal polyethism in Korean yellowjacket foragers,Vespula koreensis (Hymenoptera,Vespidae)

We examined the foraging behavior of the Korean yellowjacket, Vespula koreensis, to determine whether this species displays temporal polyethism. Using video-recordings of the entrances of artificial nest boxes installed in the field, we investigated the association between the tasks performed by workers and age. We identified three foraging tasks (pulp, nectar and prey foraging). Pulp foraging was performed by younger foragers, while nectar and prey foraging were performed by older foragers. We measured the foraging time (time spent outside of the nest during a single foraging bout) and the weight of the materials that foragers brought into the nest for each task to estimate the cost of the task. Pulp foraging was less costly than nectar or prey foraging by both measures. Taken together, the results suggest that yellowjacket foragers tend to perform low-cost task in their early foraging days and high-cost task later. Our results add to a growing literature showing temporal polyethism in social insects.     

Comparative characteristics of nests of Vespula germanica (F.) and Vespula vulgaris (L.) (Hymenoptera: Vespinae) from Christchurch City,New Zealand

Abstract

From July 1987 to June 1989, 141 nests and 365 nests, respectively, of the adventive social vespines the German wasp, Vespula germanica (F.), and the common wasp V. vulgaris (L.), from the City of Christchurch, New Zealand, were evaluated for seven major characteristics.

There were few, if any, differences in nest sites, and from spring to early summer in nest traffic, nest size, and numbers of combs. However, Getman wasps showed no preference for direction of nest entrances, while common wasp nests were more numerous in areas most exposed to the morning sun. Some Getman wasp nests survived the winter and began producing new worker cells by late June, but all common wasp nests died by June. Because common wasp nests have been reported surviving the winter in beech forest, which produces honey dew, and honey dew is not available in Christchurch City, common wasp nest survival over winter may be more dependent upon carbohydrates and/or prey dependent upon carbohydrates than the survival of German wasp nests. The presence of large, expanding wasp nests from early spring must impose localised predation pressures virtually unknown in the Northern Hemisphere where overwintering nests are rare. Wasp population dynamics, and impacts of wasps on fauna, are likely to vary between different geographical areas of New Zealand, but as targets for biological control, the two wasp species can generally be considered to be quite similar.     

An uneasy alliance:a nesting association between aggressive ants and equally fierce social wasps

Although the Neotropical territorially dominant arboreal ant Azteca chartifex Forel is very aggressive towards any intruder,its populous colonies tolerate the close presence of the fierce polistine wasp Polybia rejecta(F.).In French Guiana,83.33%of the 48 P.rejecta nests recorded were found side by side with those of A.chartifex.This nesting association results in mutual protection from predators(i.e.,the wasps protected from army ants;the ants protected from birds).We conducted field studies,laboratorybased behavioral experiments and chemical analyses to elucidate the mechanisms allowing the persistence of this association.Due to differences in the cuticular profiles of the two species,we eliminated the possibility of chemical mimicry.Also,analyses of the carton nests did not reveal traces of marking on the envelopes.Because ant forager flows were not perturbed by extracts from the wasps’Dufour’s and venom glands,we rejected any hypothetical action of repulsive chemicals.Nevertheless,we noted that the wasps"scraped"the surface of the upper part of their nest envelope using their mandibles,likely removing the ants'scent trails,and an experiment showed that ant foragers were perturbed by the removal of their scent trails.This leads us to use the term"erasure hypothesis."Thus,this nesting association persists thanks to a relative tolerance by the ants towards wasp presence and the behavior of the wasps that allows them to"contain"their associated ants through the elimination of their scent trails,direct attacks,"wing-buzzing"behavior and ejecting the ants.     

Effects of prey quantity and quality on predatory wasps

1. The simultaneous effects of prey quantity and prey quality on fitness correlates of the predatory wasp Polistes fuscatus were examined in a glasshouse study. Prey quantity was manipulated by providing prey in excess (high quantity) or one‐third of that (low quantity). Prey quality was manipulated by providing either palatable (Manduca sexta) caterpillars or unpalatable (Junonia coenia) caterpillars. 2. The effect of prey quality on wasp production depended on prey quantity. Nests given unpalatable prey produced few wasps whereas nests given palatable prey increased wasp production with increased prey. 3. The low production of nests given unpalatable prey reflected the low acceptability of those prey. The wasps preferred the palatable prey and learned to reject the unpalatable prey. With no choice of prey, they took only enough unpalatable prey to develop a small nest or colony. 4. A diet of unpalatable prey also resulted in smaller wasps and reduced the proportion of males produced, from about 40% to just 8–14%, depending on the year.     

The importance of location and visual cues during foraging in the German wasp (Vespula germanica F.) (Hymenoptera: Vespidae)

Abstract

The way in which foraging wasps use cues for prey location and choice appears to depend on both the context and on the type of prey. Vespula germanica is an opportunistic, generalist prey forager, and individual wasp foragers often return to hunt at sites of previous hunting success. In this paper, we studied which cues are used by this wasp when relocating a food source. Particularly we analysed the response to a displaced visual cue versus a foraging location at which either honey or cat food had been previously presented. We conclude that location is used over a displaced visual cue for directing wasp hovering, although the landing response is directed differently according to bait type. When wasps are exploiting cat food, location also elicits landing, but if they are exploiting honey, a displaced visual cue elicits landing more frequently than location.     

Changes in the temporal pattern of antennal drumming behavior across the <Emphasis Type="Italic">Polistes fuscatus</Emphasis> colony cycle (Hymenoptera,Vespidae)

Social wasps in several genera exhibit a diverse array of conspicuous vibrational behavior patterns closely associated with larval feeding. Polistes, as the only genus in which these substrate-borne mechanical signals have been studied in some detail, is a useful system for understanding their functions. Most Polistes species examined perform antennal drumming (AD) in the context of feeding prey liquid to larvae. Two existing hypotheses on the function of AD propose that it is a behavioral releaser signal that regulates the release of larval saliva, but with opposite effects. One proposes that AD stimulates larvae to release their saliva for the drumming adult to imbibe, whereas the other proposes that AD inhibits saliva release. A recently proposed third hypothesis argues instead that AD has a modulatory effect on development: exposure to high levels of AD biases larvae toward a worker phenotype as adults. While the larval-saliva-release hypothesis for AD function has little support, predictions made by both the inhibition hypothesis and the mechanical switch hypothesis are yet to be tested within the broader ontogenetic framework of the Polistes colony cycle. We investigated the contexts, rates of performance, and actors associated with AD across 13 weeks of the P. fuscatus colony cycle. Mean colony-wide rates of AD were high during pre-emergence and early post-emergence stages, but dropped dramatically following the third week after the first workers emerged. This variation in the temporal pattern of AD was correlated neither with the rate at which larvae were fed liquid, the number of larvae on the nest, nor with the adult-to-larva ratio, but was solely a function of colony stage. In contrast, rates of feeding liquid to larvae varied only as a function of the number of larvae in the nest. Queens drummed and fed liquid to larvae at much higher rates than did workers. Queen AD and feed-liquid rates decreased after the third week of worker emergence. During the same period, total feed-liquid rates of workers became as high as levels of queens during pre-emergence. Colony-wide AD rates dropped dramatically because workers seldom drummed while feeding liquid to larvae. The mean duration of AD bursts for queens also decreased after the second week of worker emergence. These results fail to support the salivary inhibition hypothesis, but provide indirect support for the mechanical switch hypothesis on AD function.     

Nest site competition between cavity nesting passerines and golden paper wasps Polistes fuscatus

Nest boxes provide sheltered nesting sites for both passerines and paper wasps. Although neither wasps nor birds appear to evict the other once one is fully established, it is unclear which is the dominant competitor at the onset of the breeding season. Using wire mesh, we excluded birds but not golden paper wasps Polistesfuscatus from alternating boxes along a transect through edge habitat in North Carolina from 2006 – 2008. If wasps dominate Carolina chickadees Poecile carolinensis and eastern bluebirds Sialia sialis during the early spring (all have similar nest initiation dates), we would expect wasps to settle in both box types at equal frequencies. However, if birds dominate wasps, we would expect wasp nests to be concentrated in “bird‐proof” boxes. We found wasps in bird‐proof boxes significantly more often than in bird‐accessible boxes, indicating that secondary‐cavity nesting birds are able to exclude wasps from available nest sites. Additionally, we found that during the period of nest initiation, birds usurp wasps more often than vice versa.     

Les recherches sur les lépidoptères foreurs des graminées et leurs antagonistes : bilan et perspectives

Ancistrocerus, Discoelius and Symmorphus are xylicolous predatory wasps, the former two genera hunting caterpillars, the last one using both caterpillars and larvae of Chrysomelidae beetles. One of the aims of this study was to compare the stinging pattern of species from these genera with the ancestral “complete four-sting pattern” (C4SP) and reduced “two-sting pattern” (C2SP). Another aim was to clarify whether stinging pattern of predatory wasps depends mostly on the prey type or on the systematic position of the wasp. Sting traces on 6123 paralysed prey specimens, taken from 409 fresh wasp nests made by 10 wasp species in reed (Phragmites australis) trap-nests in three localities of Lithuania, were studied. The total number of stings applied to a prey specimen, the number and the share of stings directed to its body segments and the probability of stinging the segments were considered. The main conclusions of the study are: (1) the total number of stings delivered to a prey is species-specific, in some cases it may be used to distinguish fresh nests of related wasp species, as those of Symmorphus crassicornis and S. murarius and those of Discoelius dufourii and D. zonalis. (2) The distribution of stings is significantly different from the basic patterns C4SP and C2SP in all ten studied wasps; however, four caterpillar-hunting species have the distribution close to the C2SP The stinging pattern of three Symmorphus species hunting Chrysomelidae larvae included regular stings to the throat, three thoracic and the first abdominal segment; it could be abbreviated as C5SP (a complete five-sting pattern). (3) The distribution of stings among the segments of prey was found to be more dependent on the taxonomic position of the wasp species than on the prey type, thus it might be used as a behavioural character in comparative and phylogenetic studies. (4) The stinging pattern depended on the prey type as well: in the caterpillar-hunting S. debilitatus, it was quite different from the stinging pattern of the Symmorphus that hunt Chrysomelidae larvae.     

Foraging decisions of individual workers vary with colony size in the greenhead ant Rhytidoponera metallica (Formicidae, Ectatomminae)

Summary. The ability of worker ants to adapt their behaviour depending on the social environment of the colony is imperative for colony growth and survival. In this study we use the greenhead ant Rhytidoponera metallica to test for a relationship between colony size and foraging behaviour. We controlled for possible confounding ontogenetic and age effects by splitting large colonies into small and large colony fragments. Large and small colonies differed in worker number but not worker relatedness or worker/brood ratios. Differences in foraging activity were tested in the context of single foraging cycles with and without the opportunity to retrieve food. We found that workers from large colonies foraged for longer distances and spent more time outside the nest than foragers from small colonies. However, foragers from large and small colonies retrieved the first prey item they contacted, irrespective of prey size. Our results show that in R. metallica, foraging decisions made outside the nest by individual workers are related to the size of their colony.Received 23 March 2004; revised 3 June 2004; accepted 4 June 2004.     

Distance effects on resource profitability and recruitment in the giant tropical ant,Paraponera clavata

Summary We examine how cost and benefit components of resource profitability affect recruitment in the giant tropical ant, Paraponera clavata. To vary resource profitability, we changed the quantity of artificial nectar baits presented to foragers and the distance of nectar baits from the nest. Both distance to and amount of resource affected quantitative aspects of recruitment. At increased distances foragers were less likely to recruit, and fewer workers were recruited to the resource area. The amount of nectar affected the tendency of foragers to recruit, but had no effect on the number of ants recruited. Variation in resource distance was also associated with qualitative changes in recruitment strategy. Foragers at distant sites recruited from the canopy rather than from the nest, and often transferred nectar to other workers for transport to the nest. Nectar transfer and extra-nidal recruitment significantly reduced the time required for resource collection. It may also have increased the ability of workers to specialize in specific foraging tasks. A portion of the colony's foraging force specialized spatially by remaining in distant foraging areas without returning to the nest. The flexible recruitment system of P. clavata increases colonial net energetic gain rates by concentrating foraging effort on resources yielding the highest net energetic rewards, and increases the competitive abilities of individual colonies at resource sites by decreasing collection times.     

A test of adaptive hypotheses for rapid nest construction in a swarm-founding wasp

Most social insect species enlarge their nests gradually and in close correlation with the growing need for space for brood and/or stored food. In contrast, some species of swarm-founding eusocial wasps construct the nest rapidly to a final size in the first two to three weeks of the founding stage. We considered four hypotheses on the functions of rapid nest construction in the wasp Polybia occidentalis and directly tested two of them. The first hypothesis is that rapid construction maximizes output of the worker force when there are few other work demands; it predicts that construction rate remains high until the first eggs begin to hatch, following which it declines as increasing amounts of worker effort are allocated to the feeding of larvae. The second says that rapid nest construction minimizes the time the adults in the swarm are exposed to predation and the elements; it predicts that nest-construction rate should drop steeply after the nest is large enough to house all the adults in the swarm. We measured pulp-foraging rates for the first 12 days of the founding stage in control colonies and in colonies whose nests we manipulated to prevent housing of the swarm. The treatment and control groups did not differ in construction rate for several days following the housing event, contradicting the adult-protection hypothesis. Late in nest construction, treatment colonies were building at significantly higher rates than were control colonies. If demand for brood care were a major factor in determining construction rate, both groups would have responded to the eclosion of larvae in the same way and shown a parallel decline in construction rate, but this did not happen. Instead, the patterns of nest construction rate we observed provided indirect support for the two remaining hypotheses. The first of these is that rapid construction minimizes exposure of the brood to natural enemies and desiccation. The second is that rapid construction promotes competition among queens by providing empty cells for oviposition, thereby facilitating the selecting out of the less fecund of the multiple reproductive females. Also consistent with this hypothesis is the apparent absence of explosive nest construction in monogynous, eusocial bees. Received 13 October 2007; revised 31 March 2008; accepted 6 April 2008.     

Social Learning in Vespula Germanica Wasps: Do They Use Collective Foraging Strategies?

Vespula germanica is a social wasp that has become established outside its native range in many regions of the world, becoming a major pest in the invaded areas. In the present work we analyze social communication processes used by V. germanica when exploiting un-depleted food sources. For this purpose, we investigated the arrival pattern of wasps at a protein bait and evaluated whether a forager recruited conspecifics in three different situations: foragers were able to return to the nest (full communication), foragers were removed on arrival (communication impeded), or only one forager was allowed to return to the nest (local enhancement restricted). Results demonstrated the existence of recruitment in V. germanica, given that very different patterns of wasp arrivals and a higher frequency of wasp visits to the resource were observed when communication flow between experienced and naive foragers was allowed. Our findings showed that recruitment takes place at a distance from the food source, in addition to local enhancement. When both local enhancement and distant recruitment were occurring simultaneously, the pattern of wasp arrival was exponential. When recruitment occurred only distant from the feeder, the arrival pattern was linear, but the number of wasps arriving was twice as many as when neither communication nor local enhancement was allowed. Moreover, when return to the nest was impeded, wasp arrival at the bait was regular and constant, indicating that naive wasps forage individually and are not spatially aggregated. In conclusion, this is the first study to demonstrate recruitment in V. germanica at a distance from the food source by modelling wasps’ arrival to a protein-based resource. In addition, the existence of correlations when communication was allowed and reflected in tandem arrivals indicates that we were not in the presence of random processes.     

Partial nectar loads as a cause of multiple nectar transfer in the honey bee (Apis mellifera): a simulation model

Honey bee foragers transfer their nectar loads to receiver bees within the nest. Surprisingly, they often transfer to more than one receiver (published values range from 1.9 to 2.7). Several adaptive hypotheses have been proposed to explain why multiple transfer occurs. One hypothesis, information improvement, states that multiple transfer arises as an adaptive forager-driven process. Foragers use the delay in finding a receiver to assess the relative work capacities of foragers and receivers, performing recruitment dances when appropriate. Multiple transferring improves their delay information. We used a stochastic simulation model to investigate the non-adaptive partial loads hypothesis. We determined the extent to which partial crop loads and receiver filling and emptying rules (i.e. how much nectar to accept before leaving the transfer area) can cause multiple transfer. As many as 1.9 nectar transfers per returning forager were generated within biologically realistic parameter space. We suggest that much multiple transfer arises as a non-adaptive consequence of partitioning nectar foraging between foragers and receivers, but that this will also result in foragers having better information about the relative work capacities of foragers and receivers as a useful consequence. We suggest that the number of transfers caused by partial loads could also be increased by an adaptive forager-driven effort to improve their information concerning the balance of foragers and receivers and we outline a framework wherein the information improvement hypothesis can be directly tested.     

Nectar distribution and its relation to food quality in honeybee (<Emphasis Type="Italic">Apis mellifera</Emphasis>) colonies

In honeybees (Apis mellifera), the process of nectar collection is considered a straightforward example of task partitioning with two subtasks or two intersecting cycles of activity: (1) foraging and (2) storing of nectar, linked via its transfer between foragers and food processors. Many observations suggest, however, that nectar collection and processing in honeybees is a complex process, involving workers of other sub-castes and depending on variables such as resource profitability or the amount of stored honey. It has been observed that food processor bees often distribute food to other hive bees after receiving it from incoming foragers, instead of storing it immediately in honey cells. While there is little information about the sub-caste affiliation and the behaviour of these second-order receivers, this stage may be important for the rapid distribution of nutrients and related information. To investigate the identity of these second-order receivers, we quantified behaviours following nectar transfer and compared these behaviours with the behaviour of average worker hive-bees. Furthermore, we tested whether food quality (sugar concentration) affects the behaviour of the second-order receivers. Of all identified second-order receivers, 59.3% performed nurse duties, 18.5% performed food-processor duties and 22.2% performed forager duties. After food intake, these bees were more active, had more trophallaxes (especially offering contacts) compared to average workers and they were found mainly in the brood area, independent of food quality. Our results show that the liquid food can be distributed rapidly among many bees of the three main worker sub-castes, without being stored in honey cells first. Furthermore, the results suggest that the rapid distribution of food partly depends on the high activity of second-order receivers. Received 31 August 2006; revised 8 December 2006; accepted 11 December 2006.     

Orchids mimic green-leaf volatiles to attract prey-hunting wasps for pollination

An outstanding feature of orchids is the diversity of their pollination systems [1]. Most remarkable are those species that employ chemical deceit for the attraction of pollinators [2]. The orchid Epipactis helleborine is a typical wasp flower, exhibiting physiological and morphological adaptations for the attraction of pollinating social wasps [3]. As noted by Darwin [1], this species is almost entirely overlooked by other potential pollinators, despite a large nectar reward. Therefore, the mechanism for the attraction of pollinating social wasps was something of a mystery. By using a combination of behavioral experiments, electrophysiological investigations, and chemical analyses, we demonstrate for the first time that the flowers of E. helleborine and E. purpurata emit green-leaf volatiles (GLVs), which are attractive to foragers of the social wasps Vespula germanica and V. vulgaris. GLVs, emitted by damaged plant tissues, are known to guide parasitic wasps to their hosts [4]. Several E. helleborine GLVs that induced response in the antennae of wasps were also emitted by cabbage leaves infested with caterpillars (Pieris brassicae), which are common prey items for wasps [5]. This is the first example in which GLVs have been implicated in chemical mimicry for the attraction of pollinating insects.