Host location and behavioural response patterns of the parasitoid,Tachinaephagus zealandicus Ashmead (Hymenoptera: Encyrtidae), to host and host‐habitat odours

Abstract 1. To overcome the challenge of host location in patchy and complex environments, many parasitoids exploit host‐habitat derived odour cues. This study investigated the role of odour cues used during host location by the generalist parasitoid, Tachinaephagus zealandicus, a common parasitoid of Dipteran larvae found in association with decomposing carrion. 2. A Y‐tube olfactometer was used to test aspects of host searching behaviour of T. zealandicus females under different choice scenarios: combinations of host–liver complex, eaten liver, unwashed host larvae, washed host larvae, liver in six stages of decay, and control (no odour). 3. T. zealandicus females demonstrated an innate response to, and preference for, odours arising from liver substrates following interaction with the host. However, this was dependent on the stage of meat decay. Females were not attracted to liver that had never been in contact with a host, regardless of the stage of decay. There were discernible differences in parasitoid movement and behaviour under different choice scenarios. The absence of odour stimulus elicited limited movement about the olfactometer. In the presence of any odour stimuli, females were more active within the olfactometer, and when presented with two odour options T. zealandicus females demonstrate a heightened investigation response, moving across all regions of the olfactometer. 4. Knowledge of the cues that mediate host location for T. zealandicus can contribute to fields such as forensic entomology, where the presence of parasitoids on decomposing remains can be used to estimate time since death.     

The role of host- and host-plant odours in the attraction of a parasitoid, Epidinocarsis lopezi, to the habitat of its host, the cassava mealybug, Phenacoccus manihoti

In the field, the encyrtid wasp, Epidinocarsis lopezi (DeSantis) can find its host, the cassava mealybug (Phenacoccus manihoti Matile-Ferrero) on cassave plants when the host is scarce. As a step towards understanding the parasitoid's effectiveness we investigated the cues with which it locates its host-habitat. Using a fourarmed olfactometer, we determined the attactivity of various host-and host-plant odours to female E. lopezi which had been previously exposed to infested plants. Attractivity or preference of an odour was determined from the proportion of 50 choices made between the odour and a blank, or between two odours. Female E. lopezi were attracted by mealybug-infested cassava leaves (CML), but not by the odour of cassava mealybugs (CMB) alone or by uninfested plants (ULU). An artificial combination of CMB and ULU was also not attractive. CML was preferred over ULU and CMB + ULU. These results suggest that the attractive element arises from the cassava plant itself in response to CMB infestation. We therefore tested the attractivity of uninfested leaves from partly infested plants, and found that these were indeed attractive. E. lopezi probably uses the odour which is emitted from infested cassava plants to guide it to its host in the field.

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Zusammenfassung Epidinocarsis lopezi ist imstande, seine Wirte auf den Cassavapflanzen zu finden, auch wenn die Wirte nur in geringer Dichte vorkommen. Die Reaktion der Schlupfwespe auf Gerüche der Wirte und der Wirtspflanzen wurde untersucht. Weibchen der Schlupfwespe, die vorher mit Schmierläusen auf Cassavapflanzen zusammengebracht worden waren, wurden individuell in einem vierarmigen Olfaktometer verschiedenen Kombinationen dieser Gerüche ausgesetzt. Die relative Attraktivität jedes Geruchs wurde bestimmt anhand von 50 Entscheiden zwischen dem Geruch und einer Kontrolle oder zwischen zwei Gerüchen. E. lopezi Weibchen wurden von schmierlausinfizierten Blättern angezogen, aber nicht von Cassavaschmierläusen (CMB), oder von uninfizierten Cassavapflanzen (ULU). Die künstliche Mischung von CMN und ULU wurde auch nicht anlockend. CML war relativ mehr attraktiv als ULU und als die Mischung. Die Vermutung liegt vor dass E. lopezi von einem Pflanzenduft angezogen wird, der von mit Schmierläusen infizierten Cassavapflanzen ausgeschieden wird. Die Parasitoide wurden auch von uninfizierte Blättern teilweise infizierter Pflanzen angezogen. Wahrscheinlich wird im Freiland der Geruch infizierter Cassavapflanzen durch E. lopezi zur Auffindung weiterer wirtstragenden Cassavapflanzen gebraucht.

     

Behavioral responses of the small hive beetle,Aethina tumida,to odors of three meliponine bee species and honey bees,Apis mellifera scutellata

Recent studies have shown that honey bees, bumble bees, and some meliponine bee species of the genera Trigona, Meliponula, and Dactylurina are hosts of the small hive beetle (SHB) Aethina tumidaMurray (Coleoptera: Nitidulidae), a pest of honey bee colonies in various regions of the world. Olfaction has been implicated in SHB infestations of honey bee and bumble bee colonies. We used olfactometer bioassays to investigate responses of adult male and female SHBs to odors from intact colonies and separate hive components (pot honey, pot pollen, cerumen, and propolis) of three African meliponine bee species, Meliponula ferruginea (Lepeletier) (black morphospecies), M. ferruginea (reddish brown morphospecies), and Meliponula bocandei (Spinola) (Hymenoptera: Apidae). Although both sexes of the beetle strongly preferred intact colony, pot honey, and pot pollen odors, there was no evidence of attraction to propolis and cerumen odors from the three meliponine bee species. Both sexes of SHB also strongly preferred odors from honey bees, Apis mellifera L. (Hymenoptera: Apidae), over odors from the three meliponine bee species. Our results provide substantial evidence of the host potential of African meliponine bees for the SHB, and we discuss this complex association of the SHB with species within the Apidae family.     

Orientation behaviour of Helicoverpa armigera egg parasitoid,Trichogramma chilonis and the larval parasitoid,Campoletis chlorideae towards different groundnut genotypes

Behavioural responses of Helicoverpa armigera egg parasitoid, Trichogramma chilonis and the larval parasitoid, Campoletis chlorideae towards the leaves of groundnut (Arachis hypogaea) genotypes (ICGV 86699, ICGV 86031, ICG 2271, and ICG 1697-resistant, and the susceptible check-JL 24) were studied by using a Y-tube olfactometer. Orientation was studied in comparison to clean air, to insect resistant genotypes in relation to JL 24 and towards H. armigera damaged and undamaged leaves. Leaves of ICGV 86699, ICGV 86031 and ICG 2271 were more attractive to T. chilonis adults than to the clean air. They were strongly attracted to the leaves of ICGV 86699, ICGV 86031 and ICG 1697 than of JL 24. Insect damaged leaves of ICGV 86699, ICGV 86031 and ICG 1697 were more attractive than the respective uninfested leaves. C. chlorideae showed greater attraction towards leaves of ICGV 86699, ICG 2271 and ICG 1697 than the clean air, and were more attracted towards leaves of ICGV 86699 and ICGV 86031 than those of JL 24. The damaged leaves of ICGV 86699, ICGV 86031 and ICG 2271 were more attractive to C. chlorideae than the respective uninfested leaves. Thus insect resistant genotypes exhibited greater compatibility with the natural enemies in groundnut.     

Olfactory cues used in host‐habitat location and host location by the parasitoid Cotesia urabae

There is a growing body of evidence that many hymenopteran parasitoids make use of olfaction as the primary mechanism to detect and locate hosts. In this study, a series of bioassays was conducted to investigate the orientation behaviour of the gum leaf skeletonizer larval parasitoid Cotesia urabae Austin & Allen (Hymenoptera: Braconidae) in both Y‐tube and four‐arm olfactometers. In a Y‐tube olfactometer, male C. urabae were attracted only to virgin conspecific females. Host‐plant leaves, damaged leaves, host larvae, and host larvae feeding on leaves were highly attractive to female C. urabae, whereas host frass and conspecific males were not. The multiple‐comparison bioassay conducted in a four‐arm olfactometer clearly indicates that C. urabae females were significantly more attracted to the host Uraba lugens Walker (Lepidoptera: Nolidae) larvae feeding on Eucalyptus fastigata H Deane & Maiden (Myrtaceae) leaves than to any other of the odour sources tested. The results of this study show that C. urabae individuals responded to chemical cues specific to the host plant and target host insect, and support hypotheses that unreliable cues are not utilized for host location by specific natural enemies.     

Behavioural responses of the aphid parasitoid Diaeretiella rapae to volatiles from Arabidopsis thaliana induced by Myzus persicae

In response to herbivory by insects, several plant species have been shown to produce volatiles that attract the natural enemies of those herbivores. Using a Y‐tube olfactometer, we investigated responses of the aphid parasitoid Diaeretiella rapae MacIntosh (Hymenoptera: Aphidiidae) to volatiles from Arabidopsis thaliana Columbia (Brassicaceae) plants that were either undamaged, infested by the peach‐potato aphid, Myzus persicae Sulzer (Homoptera: Aphididae), or mechanically damaged, as well as to volatiles from just the aphid or its honeydew. In dual‐choice experiments, female D. rapae given oviposition experience on A. thaliana infested with M. persicae were significantly attracted to volatiles from A. thaliana infested with M. persicae over volatiles from undamaged A. thaliana and similarly were significantly attracted to plants that had been previously infested by M. persicae, but from which the aphids were removed, over undamaged plants. Diaeretiella rapae did not respond to volatiles from M. persicae alone, their honeydew, or plants mechanically damaged with either a pin or scissors. We conclude that an interaction between the plant and the aphid induces A. thaliana to produce volatiles, which D. rapae can learn and respond to. Poor responses of D. rapae to volatiles from an A. thaliana plant that had two leaves infested with M. persicae, with the two infested leaves being removed before testing, suggested the possibility that, at this stage of infestation, the majority of volatile production induced by M. persicae may be localized to the infested tissues of the plant. We conclude that this tritrophic interaction is a suitable model system for future investigations of the biochemical pathways involved in the production of aphid‐induced volatiles attractive to natural enemies.     

Attraction of male pear psylla, Cacopsylla pyricola, to female-infested pear shoots

Post-diapause winterform pear psylla, Cacopsylla pyricola (Förster) (Homoptera: Psyllidae), exhibit a highly clumped distribution in late winter in pear orchards. The behaviors leading to clumped distributions in this species are unknown, but could include aggregation for mating activities. Choice tests and assays with an olfactometer were done to test whether male psylla of the overwintering morphotype are attracted to pear shoots infested by post-diapause females and to shoots previously occupied by females. Paired choice tests in small arenas showed that males accumulated on pear shoots currently occupied or previously occupied by females if those shoots were paired with uninfested shoots or shoots previously occupied only by males. Assays with an olfactometer showed that males were attracted to volatile odors from female-infested or previously infested shoots. The exact source of the attractants (i.e., the female psylla, the pear shoot, or a combination of these sources) remains to be determined.     

Sugar in moderation: sugar diets affect short‐term parasitoid behaviour

The biological control potential of parasitic wasps in the field is expected to increase with the provisioning of sugar sources, which increase longevity and replenish carbohydrate reserves. Apanteles aristoteliae is an important parasitoid of Argyrotaenia franciscana, the orange tortrix, an economic pest in fruit crops. In the present study, the effect of sugar diet on the physiological status of A. aristoteliae is investigated in the laboratory, as well as the effects of nutritional status on short‐term olfactory orientation and parasitism behaviour, and the association between olfactory orientation and immediate parasitism activity. Levels of glycogen, fructose, total sugars, proportional weight gain and volume consumed are higher among females fed 25% sucrose solution than 10% sucrose solution, and lowest for those fed water. Sugar feeding also affects behaviour: wasps with higher levels of fructose or weight gain have a lower probability of making a choice in the olfactometer. Wasps with intermediate fructose levels or weight gain have a higher probability of orienting towards hosts than wasps with low or high levels. Among wasps that make a choice in the olfactometer, wasps fed 10% or 25% sucrose significantly prefer host versus food cues, whereas starved wasps are just as likely to select food cues as host cues.     

Behavioural and electrophysiological responses of the European corn borer Ostrinia nubilalis to host‐plant volatiles and related chemicals

The European corn borer Ostrinia nubilalis (Hübner) (Lepidoptera: Crambidae) is a worldwide pest of maize (Zea mays L.) and other crops. The semiochemicals released by maize plants and structurally‐related compounds can be used by adult female O. nubilalis for host‐plant location and oviposition. Headspace volatile compounds emitted by watered and water‐deprived maize plants are collected and identified by their retention indices and mass spectra. The most abundant compounds from watered plants are limonene, linalool, benzoic acid, indole, β‐caryophyllene and acetophenone, whereas, in water‐deprived plants, limonene, acetophenone, hexanoic acid, benzoic acid and indole are dominant. In addition, (E)‐4,8‐dimethyl‐1,3,7‐nonatriene, 6‐methyl‐5‐hepten‐2‐one, anisole and 1‐carvone are undetected in the water‐deprived plants. Some of the identified compounds show electrophysiological activity (electroantennogram) in the antennae of both sexes, with the responses elicited by tridecane, tetradecane, dodecane, nonanal, decanal and 2‐ethylhexanol on males being particularly noteworthy. In a dual‐choice olfactometer, adult females show a preference for 2‐hexanol, heptanal, methyl salicylate, hexyl acetate, nonanal, methyl dodecanoate, β‐pinene and (E)‐2‐hexenyl acetate over hexane controls. Tetradecane, linalool, methyl hexanoate, methyl nonanoate, (Z)‐3‐hexenyl benzoate, tridecane, 2‐cyclopentylcyclopentanone, 3‐methylbutyl acetate, β‐myrcene and (Z)‐3‐hexenyl butanoate result in fewer females in the test arm compared with the control arm. No single compound displays an activity similar to watered maize plants, supporting the hypothesis that blends of volatiles in specific ratios are more effective than single volatile chemicals. The results of the present study suggest that methyl salicylate, which elicits also one of the highest electrophysiological responses in female antennae, plays a role in host preference by O. nubilalis females.     

Modification of Macrosiphum euphorbiae colonisation behaviour and reproduction on potato plants treated by mineral oil

Although mineral oil spray is one of the most effective ways to control the transmission of non‐persistent aphid‐borne viruses in the field, its mode of action is poorly understood. In this study, the effects of mineral oil treatment of potato plants on host selection behaviour, growth, and reproduction of potato aphids, Macrosiphum euphorbiae (Thomas) (Hemiptera: Aphididae), were investigated. The effects were assessed 30 min, 1 day, and 7 days after treatment, (1) on aphid orientation behaviour by using a Y‐tube olfactometer, and (2) on aphid feeding behaviour by using the electrical penetration graph (EPG) technique. Olfactory experiments showed that the oil had a repulsive effect only 30 min after spraying. EPG experiments showed a slight modification of the aphid feeding behaviour mainly 7 days after treatment. The number of both salivation and sap ingestion events during the phloem phases were increased 7 days after treatment. In addition, irrespective of the time after treatment, xylem ingestion time was increased. Clip cage experiments were set up to assess potential effects of the oil treatment on aphid survival and population parameters. Nymphal mortality was increased on treated plants, whereas fecundity of surviving insects was enhanced. The antagonistic effects of oil treatment on aphids are discussed in a plant protection context.