Parasitoid behaviour: predicting field from laboratory

Abstract.  1. Most of what is known about parasitoid behaviour comes from laboratory observations: field quantitative observations on searching parasitoids are extremely difficult to do and are rare. The basic components of Aphytis melinus 's response to California red scale ( Aonidiella aurantii ) were studied in the laboratory: encounter, rejection, drumming, probing, oviposition, and host-feeding. It was then asked whether these observations provided a reliable guide to behaviour in the field in a situation that was very different from the laboratory.
2. Field observations were carried out on bark on the trunk and interior branches of trees where live scale density is extremely high in patches, dead scale make up 90% of all scale, and could be expected to interfere with Aphytis search.
3. The laboratory observations predicted well the time taken in the field for each basic event (drumming or probing) and average times spent on a scale. Also well predicted were the distributions of times spent on drumming, probing, and total time on a scale. Rejection rates were much higher in the field. Thus, the laboratory studies predicted foraging behaviour in the field with variable success; potential explanations for observed mismatch between laboratory and field and its possible larger implications are discussed.     

Gene Flow in the Exotic Colonizing LadybeetleHarmonia axyridisin North America

The introduced parasitoid,Aphytis melinusDeBach (Hymenoptera: Aphelinidae), is used for augmentative biological control of California red scale,Aonidiella aurantii(Maskell) (Homoptera: Diaspididae). Commercially reared wasps are reared on oleander scale,Aspidiotus neriiBouché (Homoptera: Diaspididae). Oleander scale covers lack the chemical,O-caffeoyltyrosine, a kairomone mediating host selection byA. melinus.Wasps reared on oleander scale but individually exposed, or primed, toO-caffeoyltyrosine more readily accepted California red scale for probing in laboratory bioassays and parasitized a greater proportion of available California red scale in the field than wasps reared similarly but not exposed toO-caffeoyltyrosine. Thus, it may be possible to improve host recognition of commercial, insectary-rearedA. melinusby exposing them toO-caffeoyltyrosine prior to release. The goal of this study was to develop an inexpensive but effective means of priming thousands of wasps simultaneously toO-caffeoyltyrosine. The most effective method, but potentially the most expensive, was simply to spray parasitized oleander scale on their host plant with diluteO-caffeoyltyrosine prior to wasp emergence. In additional experiments, using controlled doses of syntheticO-caffeoyltyrosine applied to scale covers, we showed that primed wasps require both a lower minimum dose ofO-caffeoyltyrosine for recognition and also respond to measuredO-caffeoyltyrosine doses more consistently than unprimed wasps. The ability to mass-prime thousands of wasps prior to release is a crucial step toward realizing the concept of behavioral improvement of host selection of parasitoids on a commercial scale.     

红圆蚧的空间分布及其自然种群生命表

调查了红圆蚧在不同定居地（果、叶、枝条和树干）的密度分布。结果表明,红圆蚧密度在果和树干上较高,在枝条和叶片上较低。红圆蚧的生命表研究说明,红圆蚧以各种虫态越冬,但低温对其存活影响很大。越冬成蚧存活率仅有１８．３％,各世代累积存活率在０．５４％￣３．８％之间变化。     

Primary parasitoids of red scale (Aonidiella aurantii) in Australia and a review of their introductions from Asia

We used morphological and molecular differences to confirm the identities of red scale (Aonidiella aurantii) and yellow scale (A. citrina), and their primary parasitoids, in Australia. An extension to the distribution of yellow scale was confirmed. Six primary parasitoids of red scale were identified: Aphytis chrysomphali, A. lingnanensis, A. melinus, Comperiella bifasciata, Encarsia citrina, and E. perniciosi. With the exception of A. lingnanensis, these parasitoids, and a species of Aphelinus, were detected in association with red scale during studies in citrus orchards in coastal New South Wales between 2009 and 2012. Two races of A. melinus were recorded: one from the Indian Subcontinent, the other previously only recorded in China. The studies, and reviews of historical records, led us to conclude that 4 parasitoids, A. lingnanensis, C. bifasciata, and both species of Encarsia, were present in Australia before successful or unsuccessful formal introductions between 1902 and 1970. The A. melinus race previously recorded in China may also have been present before the Indian Subcontinent race was formally introduced in 1961. We suggest the possibility that the natural distribution of some of the parasitoids may include East and Southeast Asia, and parts of Australasia. We found no reports of native armored scales being recorded on species and hybrids of Citrus introduced to Australia, and no reports of introduced armored scales being recorded on native Rutaceae, including 6 species of Citrus. However, we subsequently recorded yellow scale on Geijera parviflora, a native rutaceous tree.     

Effects of carbonyl sulfide, methyl iodide, and sulfuryl fluoride on fruit phytotoxicity and insect mortality

Three potential chemical fumigants: carbonyl sulfide (COS), methyl iodide (MI) and sulfuryl fluoride (SF) were tested at selected dosages on lemons against California red scale (Aonidiella aurantii) and MI and COS were tested on nectarines against codling moth (Cydia pomonella). In nectarines, COS was tested at 0, 20, 40, 60 and 80 mg litre^?1, MI at 0, 10, 15, 20 and 25 mg litre^?1. Both fumigants intensified nectarine peel color, delayed fruit softening, but did not alter overall fruit quality. COS at 80 mg litre^?1 resulted in 87% codling moth mortality, but the fumigant dosage was insufficient to reach the desired probits 9 level (99.9968%). MI gave 100% codling moth mortality at 25 mg litre^?1. Lemons were treated with MI at 0,10,20,40,60 mg litre^?1, SF at 0,10,20,40, 80 mg litre^?1 and COS at 0,20,40, 60 and 80 mg litre^?1. MI gave 100% red scale mortality at ≥40 mg litre^?1 but caused significant fruit injury. Conditioning lemons at 15°C for 3 days before MI fumigation lessened lemon phytotoxicity. Forced aeration at 3.5 standard litres per minute of lemons for 24 h following MI fumigation at 20 mg litre^?1 significantly reduced phytotoxicity compared to 2 h postfumigation aeration after MI treatment. SF at ≥40 mg litre^?1 gave 100% red scale mortality but resulted in commodity phytotoxicity. Lemons treated with the highest selected dose of 80 mg litre^?1 COS gave only 87% kill of red scale, but failed to reach the desired probit 9 level.     

Effect of three diaspididae prey species on development and fecundity of the ladybeetle Chilocorus bipustulatus in the laboratory

The purpose of this investigation was to determine the most suitable prey for the development and fecundity of the predatory coccinellid, Chilocorus bipustulatus (L.) on three different diaspidid species: Aspidiotus nerii Bouché, Aonidiella aurantii (Maskell), and Pseudaulacaspis pentagona (Targioni-Tozzetti). Life tables were constructed at constant 25°C and the developmental time, longevity, fecundity, and the sex ratio were determined. Individuals fed with A. nerii displayed the shortest larval and pupal developmental time (26.0 days), lowest immature mortality rate (16.6%), highest net reproduction rate (264.7 females/ female), shortest generation time (72.9 days), and the highest intrinsic rate of increase (0.077 females/female/day). The results showed that under laboratory conditions C. bipustulatus developed best on A. nerii.     

Host recognition and selection by Aphytis species: Response to California red, oleander, and cactus scale cover extracts

One or more chemicals associated with the host's cover appear to play a major role in host recognition by Aphytis species. Aphytis melinus DeBach, A. lingnanensis Compère, A. coheni DeBach and Comperiella bifasciata Howard, all parasitoids of diaspidid scale-insects, were found to respond to water extracts of California red scale covers, Aonidiella aurantii (Mask.); A. melinus and A. lingnanensis responded to water extracts of cactus scale covers, Diaspis echinocacti (Bouché); and A. chilensis Howard to water extracts of oleander scale covers, Aspidiotus nerii Bouché. A. lingnanensis responded to water extracts of cactus scale covers only if it had been reared on this scale. A. melinus also responded to ethanol extracts of California red scale covers while A. lingnanensis did not. A. melinus failed to respond to water extracts of California red scale covers after a single ovipositional experience on California red scale. Such kairomones might prove useful in screening natural enemies as potential biological control agents for specific hosts as well as in elucidating the mechanisms by which parasitoids of scale-insects differentiate among potential hosts.

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Résumé Aphytis lingnanensis, A. melinus, A. coheni et Comperiella bifasciata réagissent tous à des extraits aqueux d'Aonidiella aurantii, tandis qu'A. chiliensis réagit aux extraits aqueux d'Aspidiotus nérii. La réaction de A. coheni aux extraits aqueux d'A. aurantii confirme les écrits de Quednau et Hübsche (1964). Nous n'avons pas été capables d'observer une réaction de A. chrysomphali aux extraits aqueux d'A. aurantii. Comme nous n'avons pas essayé de traiter les boucliers avec d'autres solvants, nous ne pouvons pas affirmer l'absence de réponse de cet aphélinide aux kairomones de cette cochenille.Que A. melinus et A. lingnanensis aient été élevés sur Aspidiotus nerii ou sur Aonidiella aurantii, ils ne réagissent pas aux extraits aqueux d'A. nerii; qui est accepté comme hôte par ces deux aphélinides au laboratoire. Ces résultats suggèrent que d'autres éléments, comme la forme ou la texture, jouent aussi un rôle important dans la détermination des hôtes. A. melinus et A. lingnanensis réagissent tous deux aux extraits aqueux de boucliers de Diaspis echinocacti; cependant, après élevage sur D. echinocacti, seul A. lingnanensis réagit à l'extrait. La réaction de A. melinus à l'extrait de D. echinocacti suggère que cet aphélinide est préadapté chimiosensoriellement à déceler cette cochenille, dont la distribution est néotropicale tandis que celle de A. melinus orientale. A. melinus a été élevé à partir de D. echinocacti récoltés dans la nature. La réaction de A. lingnanensis aux extraits de D. echinocacti uniquement lorsqu'il a élévé dessus, suggère qu'un conditionnement préimaginal peut aussi jouer un rôle dans la détermination de l'hôte.

     

Volatile cues used by the parasitoid, Aphytis melinus, for host location: California red scale revisited

Aphytis melinus DeBach (Hymenoptera: Aphelinidae) is a biological control agent of diaspidid scale insects. The parasitoid has a narrow host range but its hosts are polyphagous. We determined the source of volatile cues the wasp uses to locate its few host species when those hosts occur on more than one host plant species. We addressed four questions in regard to the use of volatile cues in host location of California red scale, Aonidiella aurantii (Maskell) (Homoptera: Diaspididae): (1) Does A. melinus use volatile cues to assist in host location? (2) Are these cues innately recognized or learned? (3) Are cues produced by female California red scale, or from other sources? (4) Are the cues specific to the host or host plant? These questions were tested through the use of a Y-tube olfactometer. Female A. melinus used volatile cues to orient toward both infested and uninfested host plant material. Wasps learned these cues by associating odors from the host plant with host presence. They had no innate preferences for scale insect or host plant volatile stimuli. Contrary to previous studies, we found no evidence of orientation toward the female-produced sex pheromone of California red scale, nor to volatile cues from the attacked host stage. Wasps given experience with scale insects growing on lemon fruit subsequently oriented toward lemon and orange fruit and leaves. The scale species with which the wasp was given experience did not affect this preference. Wasps given experience with California red scale growing on squash did not orient toward infested lemon fruit. The host ranges of the parasitoid and its hosts are used to explain the adaptive value for the evolution of learned rather than fixed responses to cues used in foraging behavior.     

三种寄生物与一种寄主的系统模型研究

建立了红圆蚧与３种寄生蜂,盾蚧长缨蚜小蜂、岭地蚜小蜂和印黄蚜小蜂相互作用的系统模型,按红圆 的面积大小分成３种,ｓ（０．５５ｍｍ＾２以上）,ｔ（０３９－０．５５ｍｍ＾２）、ｕ（０．３９ｍｍ＾２以下）,分析三维系统（２种寄生物与１种寄主）和四维系统３种寄生物与１种寄主）在不同红圆蚧大小分布（ｓ,ｔ,ｕ）条件下系统的动态变化,确定了寄生物相互排扩和边界条件,为红介的生物防治决防治决策提供理论指导。     

Discovery of a CI-inducing Wolbachia and its associated fitness costs in the biological control agent Aphytis melinus DeBach (Hymenoptera: Aphelinidae)

Wolbachia is an endosymbiotic bacterium that infects a large percentage of arthropods and can affect the fitness of its host. Here we verified for the first time that the biological control agent Aphytis melinus DeBach is infected with a Wolbachia that causes complete cytoplasmic incompatibility, and conducted an insectary and field survey to determine the infection frequency. A. melinus appears to suffer fitness costs associated with infection based on measurements of longevity and fecundity. We also quantified the Wolbachia titers of A. melinus reared at different temperatures and found that, although not completely cured, increased temperature resulted in a significant reduction in the number of Wolbachia copies found in an individual wasp. Implications of our results for biological control are discussed.