Behavioral Attraction of Two Parasitoids,Venturia canescens and Bracon hebetor,to Silk Extracts of a Host Plodia interpunctella

Kairomonal activities of silk extracts of host Plodia interpunctella were determined by measuring the rates of behavioral responses of two parasitic wasps, Venturia canescens and Bracon hebetor. Silk of P. interpunctella larvae attracted both parasitic wasps but the cocoon silk of silkworm, Bombyx mori and the web silk of two-spotted spider mite, Tetranychus urticae did not. Silk components of P. interpunctella were extracted by using either hexane or methanol, and tested the rates of three serial responses of wasps; host location, antennal drumming and ovipositor probing behaviors. The patterns of each behavioral response were similar in two wasps. The rates of each response were increased at the higher concentrations of both extracts. Antennal drumming behavior was much more responsive to lower concentrations of both extracts than ovipositor probing behavior was. Furthermore, the rate of antennal drumming response was higher in hexane-extracts rather than methanol-extract in both wasps; V. canescens and B. hebetor for 20 and 17 times, respectively. However, ovipositor probing response was similar in two different extracts. Both extracts elicited 100% of antennal drumming response but ovipositor probing response was only 60 to 80% of all tested individuals. Our results were shown that silk extracts of host larvae elicited strong behavioral responses of two parasitic wasps and could be applied for practical application of parasitoids attraction in the biological control of agricultural pests.     

Chemical signals mediating interactions betweenGaleruca tanaceti L. (Coleoptera, Chrysomelidae) and its egg parasitoidOomyzus galerucivorus (Hedqvits) (Hymenoptera, Eulophidae)

Chemical signals mediating interactions betweenGaleruca tanaceti and its egg parasitoidOomyzus galerucivorus (Hymenoptera, Eulophidae) were studied. Neither odor of gravid females ofG. tanaceti nor volatiles of their feces were attractive to the parasitoid. However, the presence of the beetles’ feces on a substrate arrested the parasitoid and elicited frequent antennal drumming. Thus, feces may contain a kairomone important for host finding. Odors of damaged and undamaged host plants had no effect on the parasitoids.O. galerucivorus did not detect its host eggs at close range but encountered them by chance. Neither the structure nor the dark color of the egg surface play a key role in host recognition, but chemicals of the extrachorion which could be isolated by dichloromethane. Fractionation of the dichloromethane extract by TLC revealed a single active fraction which induced host recognition behavior. Since the eggs ofG. tanaceti contain anthraquinones and anthrones which are active as feeding deterrents against predators, we hypothesized that reproductive success ofO. galerucivorus is due to sequestration of these protective compounds. However, GC-MS analyses revealed that there was no transfer of them from the host egg into the adult parasitoid.     

Role of the cocoon ofAcrolepiopsis assectella (Lep., Hyponomeutoidae) in host recognition by the parasitoidDiadromus pulchellus (Hym., Ichneumonidae)

The study of various behavioural criteria of femaleDiadromus pulchellus parasitoids in the presence of theirAcrolepiopsis assectella hosts has shown the essentially chemical nature of the stimulant determining host recognition. The physical stimuli of the cocoon seem not to be implicated. Thus cocoons whose original texture has been completely altered, either mechanically or chemically, as well as the silk excreted by the host caterpillars significantly stimulates the female parasitoids. The cocoon contact kairomones are detected in testing the aqueous extracts ofA. assectella which provoke a positive behavioural response from the females in a threshold concentration of 1 cocoon-equivalent. The comparison of aqueous extracts ofA. assectella host cocoons, and of non-host species:Bombyx mori, Ephestia kuehniella andCacoecimorpha pronubana demonstrate the kairomone specificity of the silk, the extracts of non-host cocoons being ignored by the parasitoid, as were the silk threads left byE. kuehniella caterpillars. Finally the contact kairomones linked to the silk seem to be independent of the host plant and of the nutrient diet of the host caterpillars. The cocoons spun by the caterpillars reared on leeks or on artificial diets with or without powdered leek provoke similar responses in the parasitoids.     

Specificity of chemical cues used by a specialist egg parasitoid during host location

Plant synomones and host kairomones are known to guide the egg parasitoid Oomyzus gallerucae to its specific host, the elm leaf beetle Xanthogaleruca luteola (= Pyrrhalta) (Muller) feeding upon elm leaves (Ulmus spp.). In this study, we investigated whether the activities of these plant synomones and kairomones are specific for the plant and herbivore species, respectively. Olfactometer and contact bioassays were used. In habitat location, O. gallerucae (Fonscolombe) is known to use synomones from Ulmus minor (Miller) that are induced by egg depositions of X. luteola. The attractiveness of such induced volatiles was shown to be specific both for the Ulmus species and the herbivore species depositing eggs. Neither leaves of U. glabra Hudson (= U. montana) carrying eggs of X. luteola nor leaves of U. minor (= U. campestris = U. procera) carrying eggs of the chrysomelid species Galeruca tanaceti L. emitted attractive synomones. O. gallerucae is also known to be attracted by volatile kairomones from faeces of X. luteola feeding on U. minor and to show prolonged antennal drumming when contacting substrates contaminated with these faeces. The kairomonal activity of the faeces was proved to be independent of the Ulmus species, since also faeces from elm leaf beetles feeding upon U. glabra emitted attractive volatiles. However, the faecal kairomones were specific for the herbivorous species, since faeces from a lepidopteran larva (Opisthograptis luteolata L.) feeding upon elm hardly elicited any antennal drumming in O. gallerucae. The egg parasitoid studied is known to recognize host eggs of X. luteola by contact kairomones extractable from the egg shell. O. gallerucae clearly differentiated between host eggs and eggs of another closely related chrysomelid species, Galerucella lineola L., as was shown by comparing duration of antennal drumming on host eggs and eggs of G. lineola.     

Gustatory sensilla sensitive to protein kairomones trigger host acceptance by an endoparasitoid

Proteins isolated from the host cocoon of Acrolepiopsis assectella (Lepidoptera: Yponomeutoidea) act as kairomones for host acceptance by the endoparasitoid wasp Diadromus pulchellus Wesmael (Hymenoptera: Ichneumonidae). In this study, morphological, ultrastructural and electrophysiological studies were carried out in order to identify the contact chemoreceptive sensilla on the parasitoid antennae that perceive the protein kairomones. Three types of sensillum on the antennae of the females were found to have a chemosensory function. The receptor cell(s) of one sensillar type were shown to give a positive electrophysiological response to protein kairomones. This sensillar type is apically multiporous and female specific. Consequently, this sensillum could be the one implicated in the perception of the protein kairomone that triggers the host-acceptance behaviour of D. pulchellus females.     

Role of Caterpillar Silk Thread in Location of Host Pupae by the Parasitoid <Emphasis Type="Italic">Diadromus pulchellus</Emphasis>

Diadromus pulchellus is a solitary ichneumonid parasitoid. Its only known host is the pupa of Acrolepiopsis assectella, a specialist herbivore of Allium species. D. pulchellus females parasitize A. assectella pupae within 48 h after the caterpillars spin their cocoon and begin to pupate. Having observed that the cocoon produced by the leek moth caterpillar stimulates parasitoid egg-laying and that caterpillar leaves a silk thread, we studied the hypothesis that silk thread might be involved in host-finding by the parasitoid. Behavioral tests showed that when D. pulchellus females encounter a host silk thread, they change directions, follow the thread, and quickly locate the host. These findings show that pupal parasitoids can use signals produced by their hosts at the developmental instar preceding the one that they parasitize.     

Behavioural and electrophysiological responses of Pantomorus cervinus (Boheman) (Coleoptera: Curculionidae) to host plant volatiles

Abstract  Fuller's rose weevil (FRW; Pantomorus cervinus ) (Coleoptera: Curculionidae), a polyphagous pest of citrus and other horticultural plants, was studied in laboratory assays designed to identify potential semiochemicals from host plants that might be exploited for weevil pest management. Using still-air bioassays, weevils were found to be most attracted to fresh whole lemon leaves compared with cut and/or dried lemon leaves. White clover, an understorey plant in kiwifruit orchards, was also found to be attractive to weevils in the dual-choice tests. Coupled gas chromatography-electrophysiological recording of weevil antennal responses to commercial extracts of lemon leaves indicated that weevils detected at least eight monoterpene components of the oil, i.e. linalool, terpinen-4-ol, nerol, neral, geraniol, geranial, neryl acetate and geranyl acetate. Significant antennal dose–responses were evident to lemon leaf oil, lemon constituent odours and two major green leaf volatiles detected from clover ( Z )-3-hexenol and ( Z )-3-hexenyl acetate. Still-air dual-choice tests on individual chemicals showed significant repellency from seven of the lemon leaf compounds at 500 µg/100 µL (all except geranyl acetate). Weevils were attracted to a synthetic blend consisting of the green leaf volatiles over a range of concentrations (1, 10 and 100 mg/100 µL), as well as to clover leaves. These plant kairomone components may be potentially useful as repellents or attractants for FRW control and management programme.     

Why does the larval integument of some sawfly species disrupt so easily? The harmful hemolymph hypothesis

The larvae of several sawfly species belonging to the Tenthredinidae (Hymenoptera) have such a low mechanical resistance in the integument that slight mechanical damage to the integument is enough to provoke the release of hemolymph at a given spot. We quantified this phenomenon, which we call "easy bleeding", by measuring the pressure needed to pierce dissected sawfly integument. We also investigated the feeding deterrance of ethanolic extracts of the hemolymph by laboratory bioassays using Myrmica rubra ant workers. These traits, integument resistance and hemolymph deterrence, were inversely related, considering 22 tenthredinid species. A negative correlation was obtained by only taking into account the species of one tenthredinid tribe, namely the Phymatocerini (nine species studied). Our results support the "harmful hemolymph hypothesis" that we present here and that assumes a functional link between these morphological and chemical traits, jointly acting as a chemical defense strategy. We suspect hemolymph deterrence to be often due to sequestration of plant secondary metabolites. We discuss the role of easy bleeding, considering the fact that sawfly larvae are frequently the prey of invertebrate and vertebrate predators. It is suggested that invertebrates such as ants were more important than vertebrates in the evolution of easy bleeding.     

Short Distance Cues Used by the Adult Parasitoid <Emphasis Type="Italic">Microctonus hyperodae</Emphasis> Loan (Hymenoptera: Braconidae,Euphorinae) for Host Selection of a Novel Host <Emphasis Type="Italic">Listronotus oregonensis</Emphasis> (Coleoptera: Curculionidae)

Insect parasitoids use host kairomone to detect their hosts. However, in parasitoid species that attack adult hosts, the mobility of adult insect may mean that the host can move away for kairomone sources. The effect of Listronotus oregonensis LeConte (Coleoptera: Curculionidae) adult sex, feces and movement on host selection behavior by Microctonus’’ hyperodae Loan (Hymenoptera: Braconidae; Euphorinae) females was evaluated in the laboratory. We hypothesized that, in addition of using host kairomones, parasitoids of adult stage should use host movement for host selection. The sex of L. oregonensis did not affect the host selection behavior of M. hyperodae. However, host feces decreased the number of weevil antennations done by M. hyperodae. Microctonus hyperodae stopped less frequently near immobile L. oregonensis than near walking ones and these latter were frequently pursued by M. hyperodae. Host movement was the stimulus that elicited oviposition by M. hyperodae. The adaptive implications of these results are discussed.     

Increased host acceptance in experienced females of the parasitoid Brachymeria intermedia: which types of oviposition behaviour contribute to experience?

Brachymeria intermedia (Nees) (Hymenoptera: Chalcididae) is a solitary endoparasitoid of the gypsy moth, Lymantria dispar (L.) (Lepidoptera: Lymantriidae). The probability that a wasp will parasitise this host is increased if she had a previous oviposition experience. We investigated which types of pre-oviposition and oviposition behaviour induced this change by comparing the acceptance behaviour of wasps with zero to five oviposition experiences as well as wasps with various partial host handling experiences (from no contact with a host to one complete oviposition sequence). The percentage of females accepting the host increased gradually with both the number of previous oviposition experiences and the amount of handling of a single host. Furthermore, the naive females were less likely (44%) to walk to the host than females with a variety of experiences (72–100%). A single antennal contact with the host was sufficient to increase the probability of walking to the host. Additional handling and additional oviposition experiences further elevated the propensity to oviposit. Thus, the modification of the acceptance behaviour through experience was essentially a gradual process in which antennal contact with a pupa was a major element. It is suggested that naive wasps learned the host odour when they first antennated a pupa. As a result, they were attracted to the host odour in subsequent encounters.     

Behavioral responses of the parasitoid Melittobia digitata to volatiles emitted by its natural and laboratory hosts

Responses of macropterous females of the ectoparasitoid Melittobia digitata Dahms (Hymenoptera: Eulophidae) to direct and indirect cues emitted by its natural hosts as well as laboratory hosts were investigated using a Y‐tube olfactometer. To locate the nest of mud dauber wasps, Trypoxylon politum Say (Hymenoptera: Crabronidae), and one of their inquilines, Anthrax spec., parasitoids exploit volatiles from the freshly built nest mud and the empty cocoon constructed by the wasps, as well as their meconium. However, the parasitoids did not respond to odors emitted by older nest mud or by the host stages that are attacked (T. politum prepupae and Anthrax spec. larvae). Melittobia digitata was not attracted to direct volatiles released by the dipteran hosts Anastrepha ludens Loew (Diptera: Tephritidae) (a natural host) and Sarcophaga bullata (Parker) (Diptera: Sarcophagidae) (a laboratory host). Based on our results, we suggest that M. digitata adopts a ‘sit and wait’ strategy to locate mud dauber wasps, relying mainly on indirect host‐related cues: females search for nests that are under construction and once found, they wait inside the cell until the host completes its cocoon and releases meconium, an indicator that is associated with host suitability. No attraction was found to dipteran hosts, suggesting that parasitization of these hosts may be incidental, due to the broad host plasticity of Melittobia wasps.     

Chemical information from the fungus Amylostereum areolatum and host-foraging behaviour in the parasitoid Ibalia leucospoides

Abstract.  Parasitoids locate hosts using reliable and predictable cues such as smells derived from host plants or from the hosts themselves. For host species that live with mutualistic organisms, such as several wood boring insects, cues derived from the symbionts are likely to be exploited by specific parasitoids. Through a set of bioassays, the behaviour of the parasitoid Ibalia leucospoides Hochenwarth (Hymenoptera: Ibaliidae) is studied in response to the fungus Amylostereum areolatum Boidin (Basidiomycotina: Corticiaceae), a symbiont of its host, the wood wasp Sirex noctilio Fabricius (Hymenoptera: Siricidae). The results show that parasitoids are attracted to the fungus when growing naturally within pine logs, and also when growing on an artificial medium. Fungal volatiles also elicit increased parasitoid activity and may provide information on relative densities of hosts available for parasitization. It is speculated that the the chemical information derived from the host fungal symbiont comprises reliable and detectable host-locating cues used by parasitoids to search for concealed hosts.     

Haemolymph Defense Capacity of the Neotropical sawfly <Emphasis Type="Italic">Aneugmenus merida</Emphasis> against Ant Predation

The defensive characteristics of the sawflies have received special attention due to the involvement of toxic compounds obtained from host plants. In this context, the haemolymph-based defense is one of the strategies known in sawflies to dissuade the attack of predators. Aneugmenus merida is a neotropical sawfly whose larvae are herbivorous on the toxic bracken fern Pteridium spp. The present study examines the defensive properties of the A. merida larval haemolymph and its possible link with the chemistry of its host plant. We report the behavior of the solitary hunter ant Odontomachus chelifer towards A. merida larvae under laboratory conditions. In addition, we studied the liquid intake behavior of the ants provided with solutions of crude haemolymph, bracken extracts, and its fractions. A. merida larvae showed a marked defensive capacity against the ants. The inhibition of the attack was observed during the stages of antennal contact and mandibular blow, suggesting that larval defensive capacity is due to factors present in the integument and haemolymph. Aqueous and methanolic fractions of haemolymph and bracken also deterred feeding. Although some common compounds were detected in the haemolymph and bracken fractions, they were in very small quantities, suggesting that they are not responsible for the bioactivity. Therefore, the hypothetical connection between the host plant chemistry and larval defensive capacity could not be evidenced. We suggest that the deterrent compounds present in the haemolymph and integument could be jointly acting in the sawfly’s defensive strategy.     

Physical and chemical cues influencing the oviposition behaviour of Aphidius ervi

The oviposition behaviour of the aphid parasitoid Aphidius ervi Haliday is influenced by both chemical and physical cues. Oviposition attack responses were elicited by paint pigments sealed into the tip of a glass capillary tube. Parasitoids reacted to yellow pigments with repeated oviposition attack responses, but they did not react to green pigments. The spectrum of reflected light from the yellow pigments was very similar to that from the `green' natural host Acyrthosiphon pisum (Harris), with a high proportion of the total radiation energy being emitted in the yellow-orange wavebands (580–660 nm). Pea aphid cornicle secretion also elicited oviposition attack responses, which were not exclusively induced by its pale yellow-green colour. In fact, the oviposition attack response to capillary tips coated with cornicle secretion remained evident under red light conditions, which, in contrast, nearly completely suppressed the response to yellow pigments. Chemical compounds from cornicle secretion do not appear to be involved in parasitoid orientation, even though they stimulate intense oviposition attack responses. Olfactometer experiments showed that the putative kairomone involved acts only at very short range or on contact. Host exuviae, which also elicited strong and persistent oviposition reactions from A. ervi females, appear to be a good alternative source of ovipositional kairomone(s). This work confirms the existence of an aphid cuticular kairomone.     

Learning and the role of host-specific volatiles during in-flight host-finding in the specialist parasitoid Microplitis croceipes

Abstract The effect of antennal contact with host-related stimuli on the subsequent flight response to host-specific volatiles was examined in the specialist parasitoid, Microplitis croceipes (Cresson) (Hymenoptera: Braconidae). In a choice test between two species of caterpillars feeding on cowpea, free-flying female M.croceipes lacking prior contact with host cues landed more often on sources of odour from its host, Heliothis tea (Boddie), than from non-host Spodoptera frugiperda (Smith). After female parasitoids contacted host cues with their antennae, more landings on sources containing hosts were observed only when caterpillars used in the pre-flight treatment and in the odour sources fed on the same plant species. The degree to which host-experienced wasps landed more often on hosts as compared with non-hosts varied with host-instar and plant species. The role of learning in host finding in specialist parasitoids is discussed.     

Role of chemical cues and natal rearing effect on host recognition by the parasitic wasp Melittobia digitata

Parasitoids are expected to have the ability to find, recognize, and perhaps to discern potential hosts that can best support the development of their progeny. Melittobia Westwood (Hymenoptera: Eulophidae) are gregarious ectoparasitoids, which primarily attack mud daubers (Hymenoptera: Sphecidae). How Melittobia females locate their host is not well known, but the process may involve host‐related chemical signals. In this study, we investigated the roles of host chemical cues and natal rearing effect in host recognition by Mel. digitata Dahms. In an olfactometer that contained prepupae of Trypoxylon politum Say (Hymenoptera: Sphecidae), Megachile rotundata (Fabricius) (Hymenoptera: Megachilidae), puparia of Neobellieria bullata (Parker) (Diptera: Sarcophagidae), empty or intact host cocoons, or nest mud, Mel. digitata females spent significantly more time in air fields that contained T. politum (prepupae + cocoon) and Meg. rotundata (prepupae + cocoon) than in N. bullata and control fields. Nest mud and natal host had no attraction for parasitoid host choice. Most first and last choices of Mel. digitata females in the olfactometer were not consistent, suggesting an initial random dispersion, although they responded positively towards hosts in cocoons.     

Effect of forest stand type on host plant quality and direct and indirect effects on pine sawfly performance

1. Observed lower levels of herbivory in mixed compared with monoculture stands have been hypothesized to depend on top-down forces, through higher predation pressure by natural enemies or through bottom-up mechanisms through plant quality effects on herbivore performance.
2. In this study, we compared the performance measured as host plant induced mortality, cocoon weight, and predation mortality of the European pine sawfly Neodiprion sertifer (Geoffroy) (Hymenoptera, Diprionidae) in mixed and monoculture forest stands.
3. We did not observe a difference in host plant induced mortality, cocoon weight, or predation mortality between mixed and monoculture forest stands. We did find an effect of local conditions around each experimental tree on pine sawfly performance. For example, the nitrogen content of pine needles is negatively affected by the proportion of pine around the experimental tree, which in turn increases the survival of sawfly larvae.
4. The results suggest that local conditions around individual trees are more important for the performance of the European pine sawfly than stand type, i.e. mixed or monoculture plant stands.
5. We conclude that the ongoing trend for diversification within commercial forestry calls for more research where the effects of both bottom-up and top-down effects are studied at several spatial scales.

     

Host handling behavior of the egg parasitoid Telenomus triptus to the egg mass of the stink bug Piezodorus hybneri

Host handling behavior of Telenomus triptus Nixon (Hymenoptera: Scelionidae) to an egg mass of Piezodorus hybneri Gmelin (Heteroptera: Pentatomidae) was studied in the laboratory. Five distinct behavioral events could be distinguished in the host handling behavior: drumming, ovipositor-insertion, marking, walking, and resting. Female wasps showed two types of behavioral sequence in an ovipositional bout. One consisted of drumming, ovipositor-insertion, and marking, and the other drumming, ovipositor-insertion, walking, and resting. Females did not seem to lay an egg when ovipositor-insertion was not followed by marking. This was observed frequently in the early oviposition bout, on average 2.9 times per female. Females finally succeeded in parasitizing all the eggs in a host egg mass in most cases. The durations of drumming, ovipositor-insertion, and marking on an egg mass were nearly constant, while the total time spent by a female on and beside a host egg mass varied considerably as a result of variable durations of walking and resting. Self-superparasitism occurred when almost all the eggs in a host egg mass were parasitized. Females laid the first male egg within the first four eggs; this could be an adaptation to small egg masses or single egg.     

Searching and oviposition behaviour of Amitus fuscipennis, a parasitoid of the greenhouse whitefly

Abstract: Amitus fuscipennis MacGown & Nebeker (Hym., Platygasteridae) is a parasitoid of the greenhouse whitefly, Trialeurodes vaporariorum (Westwood) (Hom., Aleyrodidae) on some crops as bean and snap bean (both Phaseolus vulgaris L.) in Colombia. The searching and oviposition behaviour of A. fuscipennis was studied in the laboratory, using T. vaporariorum as a host on bean leaves. The parasitoid's basic search pathway consisted of walking, encountering the host, and drumming and probing it. While walking, the parasitoid stopped for short periods of time, partly to preen itself. Walking while searching comprised 61% of the adults' time budget and probing hosts represented 16%. After a host nymph was parasitized by A. fuscipennis , the parasitoid remained close by and continued searching for new hosts in the immediate vicinity. Such behaviour suggests area-restricted searching. The parasitoid preferred first instars of T. vaporariorum for oviposition, resulting in long developmental times. Amitus fuscipennis had a high percentage of host acceptance resulting in a high percentage parasitism (60%) of all encountered hosts. Amitus fuscipennis , on average, walked faster before an oviposition (1.4 mm/s) than other whitefly parasitoids. The implications of these findings for the control of T. vaporariorum are discussed.     

Host recognition in the pupal parasitoid Trichopria drosophilae: a morpho-functional approach

The diapriid wasp Trichopria drosophilae Perkins (Hymenoptera: Diapriidae) attacks and develops in puparia of the common fruit fly, Drosophila melanogaster Meigen (Diptera: Drosophilidae). Host recognition of T. drosophilae was studied using both a morphological and behavioural approach. Scanning and electron microscopical observations of female parasitoid antennae showed the presence of two types of sensilla, which we named MGS1 and MGS2. The former are present on the ventral side of both the apical (A11) and sub‐apical (A12) antennomeres, while the latter occur only on A12. Ultrastructural features suggest a gustatory function for these sensilla. Arena bioassays using intact or antennaectomised females and intact host puparia showed that MGS2 are necessary for achieving host acceptance. Further bioassays, where the host's anterior spiracles were covered with wax, led to a very low level of host acceptance. We suggest that the secretion produced by glands associated with the anterior spiracles act as a contact kairomone, which has to be perceived by MGS2 in order to elicit host recognition. The removal of both the female apical antennomeres (A12) led to the failure of the parasitoid to recognize its host.