Survival in Dalbulus leafhopper vectors improves after exposure to maize stunting pathogens

Using D. maidis and pathogen isolates collected at three field sites along an altitudinal gradient in Mexico, we compared survival in leafhoppers exposed to healthy maize to those exposed to maize infected with one of four isolates of maize stunting pathogens: two isolates of the corn stunt spiroplasma (CSS, Spiroplasma kunkelii) and two of the maize busby stunt phytoplasma (MBSP). Survival improved after exposure to either plant pathogen under both the cooler and warmer environmental conditions D. maidis is likely to encounter during the dry season. Survival varied among leafhoppers from the different field sites, suggesting that gene flow between these populations is limited. The leafhoppers responded differently to the four isolates (i.e., we noted significant population by exposure interactions), but we found no difference between MBSP and CSS exposure. Finally, we found evidence of local adaptation in one leafhopper population to sympatric, as compared to allopatric, plant pathogens. We have shown with this and our earlier study that aspects of the interaction phenotype in the association between D. maidis and the plant pathogens are mutualistic and that this association has considerable potential as a model for studies of local adaptation.     

An attachment tip and pili-like structures in insect- and plant-pathogenic spiroplasmas of the class Mollicutes

Ultrastructural studies using scanning electron microscopy (SEM), negative-staining transmission electron microscopy (TEM), and thin-sectioning TEM on four species of Spiroplasma, in vitro and/or in vivo, indicated that their helices commonly possess one tapered end (tip structure) and one blunt or round end. These tip structures appeared morphologically different from the rest of the helix, exhibiting an electron-dense conical or rod-shaped core. In thin sections of the midgut of the leafhopper Dalbulus elimatus, the tip structures of Spiroplasma kunkelii in the midgut lumen were mostly aligned between microvilli, perpendicular to the apical plasma membrane of epithelial cells. These tip structures appeared frequently attached or closely apposed to the plasma membrane, in which cup-shaped invaginations close to the tips were observed. Pleomorphic forms of spiroplasma, enclosed in membranous vesicles, were found in the cytoplasm of the midgut epithelial cells. These findings suggest that the tip structure may be involved in the orientation and attachment of spiroplasma helices in relation to their host cells, and thus may be functionally comparable to the attachment organelle of mycoplasmas. Additionally, pili-like structures were observed by negative-staining TEM on the surface of Spiroplasma melliferum, and in thin sections of S. kunkelii infecting the leafhopper vector Dalbulus gelbus. Abbreviations CSS Corn stunt spiroplasma - SEM Scanning electron microscopy - TBS Tris-buffered saline - TEM Transmission electron microscopy     

Leafhopper response to genetically diverse maize stands

At a site in Nicaragua with high population densities of the leafhopper Dalbulus maidis Delong & Wolcott, leafhopper densities were significantly lower in mixed stands of maize (Zea mays mays L.) varieties than would be expected by averaging the densities found in the pure stands of the component varieties. This response to genetic diversity appears to be due to a behavioral response during the period of colonization or establishment. The reduction in leafhopper abundance was not clearly reflected in a reduction in the incidence of the corn stunt pathogen transmitted by the leafhopper, probably because of increased leafhopper movement in mixed stands.

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Résumé Deux champs du Nicaragua, très différents quant aux densités moyennes de la cicadelle, Dalbulus maidis, et à la fréquence du nanisme par spiroplasme du maïs, — dont la cicadelle est le vecteur —, ont servi à l'étude de l'influence de l'hétérogénéité génétique sur l'abondance de Dalbulus maidis Delong & Wolcott. La maïs a été semé en parcelles pures d'une seule variété ou en parcelles génétiquement hétérogènes avec mélange de 5 variétés. Dans la zone où D. maidis est très abondant, la densité de la cicadelle était significativement plus faible dans les parcelles hétérogènes que ne le laissaient prévoir les densités moyennes observées dans les parcelles des variétés pures intervenant dans le mélange. Cette réponse à l'hétérogénéité génétique semble due aux réactions comportementales pendant la période de colonisation ou d'installation. La diminution de l'abondance de D. maidis ne se traduit pas nettement par une réduction de la fréquence du nanisme du maïs, peutêtre par suite d'un accroissement des mouvements de cicadelles dans les parcelles hétérogènes.

     

Oviposition behavior of the cabbage root fly,Delia radicum (L.), influenced by host plant extracts

An ethanolic extract of cabbage leaves (Brassica oleraceavar. capitata,Golden Acre)and derived fractions were tested on natural and surrogate leaves in order to study the oviposition behavior of the cabbage root fly Delia radicum(Diptera: Anthomyiidae). On surrogate plastic leaves coated with a thin layer of paraffin wax and treated with 0.1 gram leaf equivalent (gle) of an ethanolic raw cabbage extract, the females displayed the same sequence of behavioral patterns as on a natural host plant. The quantified oviposition behavior correlated well with the actual number of eggs laid. The extract-treated surrogate leaves received at least as many eggs as natural leaves with a similar surface area. Previous exposure to surrogate or natural leaves seemed not to induce a specific preference. The three fractions (hexane, butanol, and water) of the raw extract stimulated the oviposition behavior. This was taken as evidence for the presence of a multicomponent mixture or a chemical pattern in the cabbage leaves stimulating oviposition. At the tested concentration each fraction alone could stimulate in some individuals the complete behavior. No effect of volatile components of the raw extract could be detected. This is thought to be due to the extraction procedure, which limits the production of attractive, volatile compounds, such as isothiocyanates.     

Oviposition behavior of Edovum puttleri, reared on two hosts, Leptinotarsa decemlineata and L. texana

Female Edovum puttleri Grissell [Hymenoptera: Eulophidae], reared from eggs of Leptinotarsa decemlineata (Say) or Leptinotarsa texana Schaeffer [Coleoptera: Chrysomelidae], were videotaped as they attacked egg masses of L. decemlineata containing 20 host eggs. We identified 15 components of ovipositional behavior. Parasitoids reared on L. texana attacked and oviposited in significantly more host eggs than did females reared on L. decemlineata. Ethometric analyses of behavioral transitions and a clustering analysis of 34 behavioral parameters showed that females reared on L. texana attacked the host egg mass in a different manner than those reared from L. decemlineata. It was concluded that differences were associated with the host species upon which they were reared. Contrary to previous reports, mortality of unparasitized hosts was caused by an ovipositor probe of short duration, which was not related to host-feeding.     

Consequences of mixed species infestation on the searching behavior and parasitism success of a larval parasitoid

When two herbivore pest species are potential hosts of a single parasitoid species, two questions arise. Firstly, which host is preferable for mass rearing in terms of later parasitoid performance, and secondly, how do parasitoids perform in mixed herbivore situations after colony establishment? We tested Hyssopus pallidus, a gregarious parasitoid of two major pests of apple, Cydia (Grapholita) molesta and Cydia pomonella, before and after landing on apples infested by one of the two Cydia species. Pre-alighting host preference was tested in a Y-tube olfactometer setup, and parasitism success in a contact bioassay. To gain information on parasitoid performance throughout the growing season, different fruit growth stages were used. Irrespective of the host they had developed on, the parasitoids showed similar olfactory preferences when given a dual choice between infested and healthy fruits, and they did not discriminate between fruits infested by C. molesta and C. pomonella. Responsiveness was generally high, especially late in the season close to harvest. Both hosts are parasitized regardless of the host the parasitoid female had developed on, and no differences in parasitism rates or number of offspring were noted for the two hosts offered. Results were consistent for all apple growth stages tested. In conclusion, mass rearing of this parasitoid can be carried out on either host, without limiting the future efficacy of the bio-control agent. Similarly, established colonies are expected to develop further on both hosts without any bias in host preference.     

Release and establishment in Nigeria of Epidinocarsis lopezi, a parasitoid of the cassava mealybug, Phenacoccus manihoti

The encyrtid wasp Epidinocarsis (= Apoanagyrus) lopezi (De Santis) was imported from Paraguay into Nigeria for the biological control of the cassava mealybug, Phenacoccus manihoti Matile-Ferrero. It was mass-reared and released at four localities in Nigeria. The parasitoid is now established and it is dispersing throughout cassava growing areas of Nigeria.

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Libération et installation au Nigéria d'Epidinocarsis lopezi, parasitoïde de la cochenille du manioc Phenococcus manihoti

Résumé Epidinocarsis lopezi (Apoanagyrus) lopezi a été introduit du Paraguay au Nigéria pour lutter contre la cochenille du manioc, Phenacoccus manihoti. Il a été lâché dans quatre champs de manioc pour étudier son acclimatation et son installation au Nigéria. Trois ans après les lâchers, les résultats ont permis de conclure que E. lopezi s'est établi avec succès et se disperse dans la plupart des zones de culture du manioc au Nigeria; il a aussi survécu à trois saisons pluvieuses pendant lesquelles les populations de P. manihoti ont été très faibles. Quatorze mois après les premiers lâchers, cet encyrtide a été obtenu à environ 150 km du lieu de libération.

     

Interactions of Russian wheat aphid, a hymenopterous parasitoid and resistant and susceptible slender wheatgrasses

Interactions among three trophic levels of resistant and susceptible slenderwheat grasses, Elymus trachycaulum (Link) Goule ex Shinners ex. H.F. Lewis, Russian wheat aphid, Diuraphis noxia (Mordvilko), and a hymenopterous parasitoid were studied in the laboratory and greenhouse. These relationships were compared with a commercial susceptible wheat Triticum aestivum L. variety. Aphids reared on the resistant entries showed significantly lower weights and numbers. Significant reduction of parasitoid mummy weight and adult size was positively correlated with the effects on the aphids. Resistant entries also induced a longer prereproductive period for both the aphids and parasitoids. Numbers of aphids and aphid damage were significantly modified by the addition of parasitoids. Parasitism was higher on plants that did not have leaf rolling. These findings may indicate that antibiosis resistance studied here is not the most desirable because it decreases natural enemy vitality.     

Oviposition and reproductive performance of a generalist parasitoid (Trichogramma pretiosum) exposed to host species that differ in their physical characteristics

The response of generalist egg parasitoids to alternative natural hosts that are present simultaneously is not well known. We investigated the behavior of Trichogramma pretiosum Riley (Hymenoptera: Trichogrammatidae) in relation to two field hosts Helicoverpa armigera Hübner and Spodoptera litura Fabricius, in choice and no choice tests. We quantified the effects of natal host species and post-emergence adult age on the oviposition preference of the parasitoids. H. armigera eggs were consistently preferred over S. litura eggs, regardless of the natal host and adult age. When only S. litura eggs were available as hosts, they were parasitized at statistically similar rates to H. armigera eggs (average of 17 ± 2.7 vs. 13 ± 3.0, H. armigera to S. litura). The adult lifespan and lifetime fecundity of T. pretiosum were variable but were affected by natal host species and/or host species to which they were exposed. Mean lifespan and fecundity of parasitoids that had developed in H. armigera eggs and were exposed to H. armigera eggs for oviposition were 13.9 ± 1.8 days and 98.7 ± 11.0 adult offspring. By contrast, those that developed in S. litura eggs and were exposed to S. litura eggs for oviposition lived for 7 ± 0.9 days and produced 53.8 ± 8.0 adult offspring. The ovigeny index (OI) was significantly lower in the parasitoids exposed to H. armigera eggs than in those exposed to S. litura eggs, regardless of the natal host, indicating that H. armigera eggs sustain the adult parasitoids better than S. litura eggs. These results are used to predict parasitoid behavior in the field when both hosts are available.     

The use of molecular assays to identify plant pathogenic organisms vectored by biological control agents

The planthopper Prokelisia marginata VanDuzee (Homoptera: Delphacidae) has beenconsidered for the biological control of theweed Spartina alterniflora Loisel(Poaceae) in Willapa Bay, Washington, U.S.A. Prokelisia marginata is a stenophagousphloem-feeding insect with the potential totransmit bacterial plant diseases that could bemoved by less-specific vectors to other plantspecies. Initial assays with PCR primers thatare putatively specific for phytoplasmas gavepositive results in Spartina. However,subsequent analyses did not indicate thetransmission of the pathogen by theplanthopper. We sequenced the 16S ribosomalRNA (rRNA) gene of the bacterial species thatgave positive results in PCR. Comparisons withsequences available in GenBank suggested thatthe positive results using the putativelyspecific PCR primers were due to the presenceof such bacteria as Pseudomonas, Holomonas, Vibrio, and Acinetobacter. We did not find phytoplasmasin either Spartina or the planthopperP. marginata.     

Oviposition experience on a host-infested plant affects flight and antennal searching behavior of Cotesia kariyai toward the host-plant complex

The flight response of Cotesia kariyaiWatanabe (Hymenoptera: Braconidae), a parasitoid of the polyphagous herbivore, Mythimna separataWalker (Lepidoptera: Noctuidae), to pairs of different plant species infested by M. separatalarvae was tested under a dual choice condition in the laboratory. The oviposition-inexperienced (naive) wasps showed preference in the order: corn > kidney bean > Japanese radish. Wasps that had previously oviposited on the less preferred plant in a pair were found to have shifted their preference to this plant at 2 h after oviposition. However, this shift became indistinct at 17 h after oviposition. Prior oviposition on a plant species other than those being compared also affected the preference. These data suggest that learning is involved in the wasp's flight response. Prior oviposition was also observed to have an effect on the antennal searching behavior of the wasp on corn leaves. Such behavioral plasticity may enhance the efficiency by which C. kariyaisearches for polyphagous hosts.     

Life-table study of Anagrus atomus, an egg parasitoid of the green leafhopper Empoasca decipiens, at four different temperatures

The objective of our study was to assess thepotential of the egg parasitoid Anagrusatomus L. (Hymenoptera: Mymaridae) for controlof the greenhouse leafhopper Empoascadecipiens Paoli (Homoptera: Cicadellidae). Theegg-adult development time, survivorship andreproduction of A. atomus were evaluatedat four constant temperatures (16, 20, 24 and28°C). Developmental time ranged from33.6 days at 16°C to 13.3 days at 28°C. Based on a linear regression ofdevelopment rate on temperature the lowerthreshold was estimated at 8.39°C. Anagrus atomus required 263.2 degree-days tocomplete its development from egg to adult. Theegg-adult survival rate and the sex ratio weresignificantly lower at 28°C than at theother three temperatures tested. The intrinsicrate of increase (r _m) variedsignificantly between all four temperatures.The potential of A. atomus to attackdifferent host ages was additionallyinvestigated. Host eggs were parasitizedthroughout their development but rate ofparasitism was reduced in host eggs older thansix days. The number of eggs parasitized waspositively density dependent but the rate ofparasitism decreased with increasing hostdensity. A maximum rate of parasitism of 62.5%was recorded. The potential impact of the eggparasitoid on the population dynamics of E. decipiens is discussed.     

Evaluation of an improved method for mass-rearing of thrips and a thrips parasitoid

Improved laboratory methods are described in detail for mass rearing of various thrips species, such as Frankliniella occidentalis, Frankliniella intonsa, Thrips palmi, Thrips tabaci (Thysanoptera: Thripidae) and a thrips parasitoid, Ceranisus menes (Hymenoptera: Eulophidae), using various foods. In one method, plant pollen and honey solution are used as food sources. In a second method, germinated broad bean seeds are used. Eggs, produced in large numbers in water, are collected by a suction funnel onto a filter paper and incubated in a Petri dish. Large numbers of larvae that hatch are collected by using food traps (plant pollen). Larvae can be reared on pollen or on germinated broad bean seeds until adult emergence without additional water and food. This method has been found useful for producing even-aged thrips at different densities (up to 500 larvae in a cage of 80 mm diameter) with relatively low mortality rates. Evaluation of this rearing method for F. intonsa, shows that during 2 weeks at 20 °C per 100 females more than 4000 females could be produced in the next generation. About 5 min per day is required to achieve this productivity of mass production. The method is also suitable for producing large numbers of the solitary endoparasitoid of thrips larvae, C. menes.     

Bionomics of Alloxysta pleuralis, a cynipoid hyperparasitoid of an aphidiid parasitoid Trioxys indicus

Alloxysta pleuralis parasitizes the larvae of Trioxys indicus, a parasitoid of Aphis craccivora. Either of the sexes of this hyperparasitoid (A. pleuralis) was ready to mate shortly after emergence. During courtship the females carried the males on the top of her back for 2–3 min. Although the hyperparasitoid is obligatory, a similar, typical oviposition exploration pattern takes place on healthy aphids, however, then the eggs are laid only in parasitized aphids in the haemocoel of the larvae of the parasitoid. The life-cycle takes about a month. Superhyperparasitism was commonly observed. Adults survived for 4–13 days.

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Résumé Alloxysta pleuralis, est un endohyperparasitoïde des larves de Trioxys indicus, parasite du puceron Aphis craccivora. Les deux sexes d'A. pleuralis sont aptes à l'accouplement peu après l'émergence (10–30 min). La femelle est passive pendant la pariade. Elle porte le mâle sur son dos pendant 2 à 3 min. La copulation dure 30 sec. Bien que l'espèce soit hyperparasite obligatoire, le comportement exploratoire de ponte s'observe sur les pucerons non-parasités. Cependant les oeufs ne sont pondus qu'à l'intérieur des pucerons parasités, dans l'haemocoele des larves de troisième stade de T. indicus. Les adultes ne s'alimentent pas sur l'hôte, mais sur le miellat des pucerons. Le cycle dure environ un mois. On observe souvent du superhyperparasitisme. Les adultes vivent de 4 à 13 jours.

     

Oviposition response and development of the egg-pupal parasitoid Fopius arisanus on Bactrocera oleae, a tephritid fruit fly pest of olive in the Mediterranean basin

To date, information is wanting with regard to the use of new exotic parasitoids against olive fruit fly, Bactrocera (=Dacus) oleae (Gmelin) (Diptera: Tephritidae), a serious pest of olives Olea europaea L., in the Mediterranean basin. We investigated the oviposition response and developmental biology on B. oleae of Fopius (=Biosteres) arisanus (Sonan) (=Opius oophilus Fullaway) (Hymenoptera: Braconidae), an egg-pupal parasitoid of tephritid fruit flies, never tested before as a potential parasitoid of this host. Our results showed that olive fruits infested with B. oleae eggs exerted a relevant attraction to gravid F. arisanus and represented a stimulus for oviposition. Nevertheless they were not as attractive to female parasitoids as the Mediterranean fruit fly, Ceratitis capitata Wiedemann (Diptera: Tephritidae), eggs infested papaya fruits (Carica papaya L.). In our experimental conditions, F. arisanus completed development in B. oleae within 33 ± 1.7 days (males) and 35 ± 1.6 (females). Increases in host egg to female parasitoid ratios of 1:1, 5:1, 10:1 and 20:1 corresponded with decreases in the percentage of B. oleae parasitisation and host killing but corresponded also with increases in absolute parasitisation. Our findings are discussed in light of possibilities of utilising F. arisanus for biological control of olive fruit fly.     

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.     

Biology, rearing and field release on Guam of Euplectrus maternus, a parasitoid of the fruit-piercing moth, Eudocima fullonia

The eulophid parasitoid, Euplectrus maternus, is a gregarious ectoparasitoid of the larvae of thefruit-piercing moth, Eudocima (Othreis) fullonia. This parasitoid is indigenous to India and was released into Guam in 1998–99 to aid in the biological control of E. fullonia, an important pest of ripeguava, banana, mango, pomegranate, litchi,papaya, tomato, orange and other fruit crops.In the laboratory, biology and rearing of E. maternus on the larvae of E. fulloniawere studied. In no-choice tests, ovipositionby E. maternus was significantly greateron first and second instar larvae than on thirdinstar larvae. However, in free-choice tests,oviposition by E. maternus wassignificantly greater on second instar larvae,followed by first and third instar larvae. Thisparasitoid did not lay eggs on later instars,either in no-choice or free-choice tests.Females laid a significantly greater number ofeggs (>80%) on the dorsal surface of thelarvae than on either lateral or ventralsurfaces (1 to 8%). In general, more eggs werelaid on abdominal segments 1–3 on larvalinstars one to three. The survival of theimmature stages (from egg to adult) of E. maternus on second instar E. fullonialarvae was 61%, which was significantlygreater than the survival rates on first andthird instar larvae (32% and 26%), indicatingthat second instar host larvae are ideal formass rearing of the parasitoid. Mated femaleE. maternus continuously laid eggs on thesecond instar larvae of E. fullonia forup to 30 days, but the greatest number of eggswere laid during the first week after exposure.When E. maternus laid more than two eggson host larva, more female progeny wereproduced, indicating female-biasedreproduction. Adult parasitoids lived longerwhen fed with a honey: water (50% w/v)solution than with pure honey. Similarly, thefecundity of females increased significantlywhen fed with the honey-water solution whencompared to feeding with pure honey. To date,E. maternus has not established in Guam.     

Comparison of oviposition behavior of a solitary and a gregarious parasitoid (Hymenoptera: Mymaridae)

Anaphes victus Huber andAnaphes listronoti Huber (Hymenoptera: Mymaridae) are respectively solitary and gregarious egg parasitoids of the carrot weevil,Listronotus oregonensis (LeConte) (Coleoptera: Curculionidae). We made detailed ethograms of the oviposition behavior on unparasitized and parasitized hosts for the two species. We then compared the behavior of virgin and mated females for the oviposition of male and female progenies. The two species did not always oviposit after insertion of the ovipositor, but these punctures without oviposition could be readily differentiated from oviposition.A. victus oviposited only once by puncture, whileA. listronoti deposited one to three eggs during the same sequence. The variability of the duration of the various components was generally lower for a given female than between females. Two components, the abdominal vibrations and the pause, were significantly shorter in ovipositions that resulted in male progency for the two species. However, an important overlap in duration prevents using these differences to sex the progeny at oviposition. Virgin females of both species, although capable of producing only males, exhibited both behaviors. Parasitized hosts were recognized through internal and external markings that were used in host discrimination.     

Flight responses in a wind tunnel of the parasitoid Microplitis croceipes to three spring and three summer host plants of Heliothis spp.

Flight responses of the parasitoid, Microplitis croceipes (Cresson) (Hymenoptera: Braconidae) to two phenological stages of spring host plants (crimson clover, Trifolium incarnatum L., hairy vetch, Vicia villosa Corbiere, and cutleaf geranium, Geranium dissectum L.) and summer host plants (cotton, Gossypium hirsutum L., sorghum, Sorghum bicolor L., and soybean Glycine max (L.) Merr.) of Heliothis virescens (F.), were measured in a dual choice situation in a wind tunnel. Preflowering clover was more attractive than preflowering geranium, followed by preflowering vetch. Clover was preferred over geranium in flowering stages as well. No within species differences were detected for the two phenological stages. Preflowering and flowering cotton and soybean were preferred over preflowering and flowering sorghum. The findings could be important in pre-determining release sites for M. croceipes in the field.     

Host-handling behavior of the parasitoid,Diglyphus minoeus [Hym.: Eulophidae], parasitizing the honeysuckle leaf-miner,Phytomyza lonicerae [Dipt.: Agromyzidae]

The host-handling behavior of individual female waspsDiglyphys minoeus which attack the larvae of the honeysuckle leaf-miner,Phytomyza lonicerae, include 4 types of behavior: probing, ovipositor insertion, host-feeding, and resting. Ovipositor insertion may either be injection of venom, probing of the host by the sensilla on the ovipositor's tip or egg laying. Three types of attacks were distinguished: oviposition attack for unparasitized larvae, host-feeding attack for unparasitized host larvae and host rejection for previously parasitized host larvae. Oviposition attack was characterized by frequent alternation between probing and ovipositor insertion, long duration of ovipositor insertion and resting, and the long duration of host-handling. Resting behavior is thought to protect the progeny against superparasitism, host-feeding by other wasps or hosts' recovery from paralysis. Host-feeding attack was characterized by frequent alternation between host-feeding and ovipositor insertion and long duration of host-feeding. Host rejection was composed mainly of probing and ovipositor insertion and short handling time.