Behavioural study of plasticity in host preferences of Trichogramma maidis (Hym.: Trichogrammatidae)

Abstract. To test the relative importance of genetic and environmental factors in host preferences of Trichogramma maidis Pint. & Voeg., a polyphagous parasitoid, the oviposition behaviour was analysed on two host species: Anagasta kuehniella used for mass rearing) and Ostrinia nubilalis a target pest). Females were compared according to the species on which they had been reared, and to that on which they had an oviposition experience. To quantify an oviposition sequence, the most frequent behavioural succession was established for each experimental situation, individual sequences were then compared with this standard one to count the number of additional behaviours. An affinity coefficient positively related to host acceptance was then calculated from these additional behaviours. A preference for O.nubilalis versus A kuehniella appeared, but rearing on A. kuehniella enhanced the level of spontaneous affinity for this unnatural host. These initial preferences were modified by an oviposition experience: the affinity for a given host was higher after an oviposition on the same species, which suggests a female's ability to learn. This learning ability was higher on the non-preferred host, which led to an equivalent final parasitic performance on both hosts. Moreover, the learning ability was specific to a given host, since the oviposition on a given species did not provide any improvement in the level of affinity for the other species.     

Oviposition and offspring survival within eggs of European corn borer: discrimination of the host embryo by female Trichogramma nubilale

Oviposition preference and offspring survival relative to the location of the host embryo was tested in the egg parasitoid Trichogramma nubilale on European corn borer host eggs. Females preferentially oviposited near to the embryo on hosts about 24 h old. Survival of Trichogramma was lower when females oviposited far from the embryo, and a higher proportion of host embryos hatched. Females did not show preferences for drilling or oviposition within very young hosts relative to where the embryo would develop, and these eggs did not produce wasps or host larvae. Female behavior was apparently adaptive in day-old hosts, not to very young hosts.     

Factors influencing parasitism of Trichogramma dendrolimi on eggs of the Asian corn borer, Ostrinia furnacalis

Laboratory studies were made to determine the capacity of Trichogramma dendrolimi to parasitize eggs of Ostrinia furnacalis, as affected by the rearing host species, substrate of host eggs, host age, original locality of host populations, and cold storage of host eggs. Wasps reared from eggs of Antheraea pernyi showed parasitic capacity on eggs of O. furnacalis on average twice as high as that of the wasps reared from eggs of Corcyra cephalonica. When the age of O. furnacalis eggs at 26 °C increased from 0–6 h to 18–24 h, the proportion of wasps that successfully parasitized host eggs, the number of host eggs parasitized, and the rate of parasitization all decreased by >50%. The number of O. furnacalis eggs parasitized per female T. dendrolimi increased with the number of host eggs available, and reached 22.9 in a 24 h period. However, the parasitic capacity of female T. dendrolimi on eggs of O. furnacalis laid on plant leaves was similar to that of O. furnacalis eggs laid on wax paper. Levels of parasitism of O. furnacalis eggs from two widely separated localities, i.e. Changchun (43.50° N, 125.20° E) and Hangzhou (30.18° N, 120.07° E), were similar. Cold storage of O. furnacalis eggs at 4 °C for 5 days did not affect parasitization. Results obtained in this study indicate that although O. furnacalis is a less preferred and less suitable host than many other hosts, such as Dendrolimus punctatus, Actias selene ningpoane, Philosamia cynthia, A. pernyi, C. cephalonica, within the host-species range of T. dendrolimi, the parasitoid has the potential to achieve 50–60% or even higher rates of parasitization of O. furnacalis eggs in corn fields under suitable conditions, and could be used in the biological control of the pest.     

Host examination walk and oviposition site selection ofTrichogramma minutum: Studies on spherical hosts

The host examination behavior of the parasitoid wasp Trichogramma minutumon spherical Manduca sextahosts is described. The mean walking speed during host examination was 0.64 ± 0.03 mm s^–1and is independent of wasp size. The paths taken by the wasps are not evenly distributed over the host surface; the majority of time is spent below 45 ° latitude. The distribution of oviposition sites is also nonrandom with respect to latitude and is not influenced by phototaxic and geotaxic responses. The initial transit made by the wasps over the host surface is a straight path which frequently passes across the highest point on the host. The length and duration of the initial transit are independent of wasp body length. However, stride length and stepping rates are proportional to wasp body length, and small wasps take more steps to complete their initial transit. The roles of the examination walk in host recognition and host volume measurement by Trichogrammaare also discussed.     

Oviposition‐induced plant cues: do they arrest Trichogramma wasps during host location?

Plants can defend themselves against herbivorous insects before the larvae hatch from eggs and start feeding. One of these preventive defence strategies is to produce plant volatiles, in response to egg deposition, which attract egg parasitoids that subsequently kill the herbivore eggs. Here, we studied whether egg deposition by Pieris brassicae L. (Lepidoptera: Pieridae) induces Brussels sprouts plants to produce cues that attract or arrest Trichogramma brassicae Bezdeko (Hymenoptera: Trichogrammatidae). Olfactometer bioassays revealed that odours from plants with eggs did not attract or arrest parasitoids. However, contact bioassays showed that T. brassicae females were arrested on egg‐free leaf squares excised from leaves with 72 h‐old egg masses, which are highly suitable for parasitisation. We tested the hypothesis that this arresting activity is due to scales and chemicals deposited by the butterflies during oviposition and which are thus present on the leaf surface in the vicinity of the eggs. Indeed, leaf squares excised from egg‐free leaves, but contaminated with butterfly deposits, arrested the wasps when the squares were tested 1 day after contamination. However, squares from egg‐free leaves with 72 h‐old butterfly deposits had no arresting activity. Thus, we exclude that the arresting activity of the leaf area near 72 h‐old egg masses was elicited by cues from scales and other butterfly deposits. We suggest that egg deposition of P. brassicae induces a change in the leaf surface chemicals in leaves with egg masses. A systemic induction extending to an egg‐free leaf neighbouring an egg‐carrying leaf could not be detected. Our data suggest that a local, oviposition‐induced change of leaf surface chemicals arrests T. brassicae in the vicinity of host eggs.     

Effect of insectary rearing on host preference and oviposition behavior of the fruit fly parasitoid Diachasmimorpha longicaudata

Diachasmimorpha longicaudata (Ashmead) has been produced in the laboratory for >160 generations on the larvae of oriental fruit fly, Bactrocera dorsalis (Hendel), the propagation hosts raised routinely on a semi-synthetic wheat diet formulation. Choice tests using modified stinging units were conducted in the laboratory to investigate whether insectary rearing had altered the host seeking and oviposition behavior of female parasitoids. Results showed that fruit fly larvae that developed in papaya, Carica papaya L. var. solo, were less preferred for oviposition than fruit fly larvae that developed on wheat diet when both were exposed concurrently to naive D. longicaudata females (= females without prior oviposition experience). The substrates (pureed papaya or wheat diet) in which treatment larvae were exposed to parasitoids did not affect oviposition preference of gravid D. longicaudata for wheat diet-reared fruit fly larvae. Our study demonstrated the possibility that rearing in an insectary system may have modified the parasitization behavior of female D. longicaudata.     

Is the parasitization capacity of Trichogramma cordubensis influenced by the age of the females?

We investigated the parasitization capacity of Trichogramma cordubensis Vargas & Cabello (Hymenoptera: Trichogrammatidae) females aged 24 h, 48 h, 72 h, 96 h, 120 h, and 144 h, using Ephestia kuehniella Zeller (Lepidoptera: Pyralidae) eggs as hosts. Wasps were held without hosts during the period of ageing, therefore being increasingly time-limited with respect to parasitization as they got older. The total number of parasitized hosts decreased as the age of the parasitoid increased. However, the proportion of lifetime parasitism carried out on the first day increased with wasp age, up to 120-h old females. These results show that the parasitization capacity of ageing T. cordubensis females changes as they become time-limited. The consequences of such changes for biological control programs are discussed.     

Endogenous and exogenous factors affecting parasitism of gypsy moth egg masses by shape Ooencyrtus kuvanae

Factors affecting the orientation, reproduction, and sex ratio of the egg parasitoid Ooencyrtus kuvanae Howard were examined. Adult females were attracted to airborne volatiles from the egg mass and accessory gland of the primary host, the gypsy moth Lymantria dispar L. Visual cues also affected host selection. Background colors against which egg masses were placed affected oviposition preference. In the absence of egg masses, color variation did not affect wasp behavior. Light is required for parasitism by O. kuvanae. The age and density of both the host and parasitoid affected wasp reproduction and sex ratios. Older egg masses issued relatively fewer wasps and higher proportions of males than did young egg masses. Likewise, wasp reproduction and the proportion of females declined with wasp age. Larger egg masses produced more wasps and lower proportions of males than did smaller egg masses. The number of offspring per female, and the proportion of female offspring, were inversely related to wasp density. Implications to biological control of the gypsy moth and parasitoid ecology are discussed.     

Effect of host diet on acceptance of eastern spruce budworm eggs byTrichogramma minutum

The acceptance behaviour of the parasitic wasp, Trichogramma minutum Riley (Hymenoptera: Trichogrammatidae), was studied on two types of egg masses of eastern spruce budworm (SBW), Choristoneura fumiferana Clemens. Host eggs were obtained from moths that had been fed balsam fir or artificial diet during their larval stages. Parasitoids were reared from both types of hosts. Host acceptance assays were conducted where females were provided with a single egg mass and with a choice between egg masses. Regardless of the parasitoid's rearing host, females parasitised hosts reared from artificial diet more often than those from balsam fir. Response was consistent in both no-choice and choice assays. Rejection of hosts tended to occur during the examining stage of behaviour, suggesting that acceptability is judged externally. If a host was accepted, the same number of eggs were laid per egg mass regardless of treatment.     

Refrigeration of Riptortus clavatus (Hemiptera: Alydidae) eggs for the parasitization by Gryon japonicum (Hymenoptera: Scelionidae)

Gryon japonicum (Ashmead) (Hymenoptera: Scelionidae) is a solitary egg parasitoid of Riptortus clavatus (Thunberg) (Hemiptera: Alydidae) one of the most important pests in soybean in Korean and Japan. Refrigeration of R. clavatus eggs for up to 120 days was evaluated as a method of mass production of G. japonicum. The refrigeration reduced the eclosion of eggs, thus no nymph emerged after 30 days of egg refrigeration. Increased duration of host acceptance behaviors by G. japonicum and some detrimental effects on parasitism rate, developmental time, longevity, and adult size of the parasitoid were found in refrigerated host eggs. However, all the biological parameters of G. japonicum were unaffected by the refrigeration of up to 30 days. Gryon japonicum parasitized 16 and 14 host eggs daily that were refrigerated for 15 and 30 days, respectively, which did not differ from parasitization of fresh host eggs. Furthermore, refrigeration of host eggs did not reduce the reproduction of the emerged adult parasitoids and emergence and sex ratio of their progeny. Gryon japonicum also parasitized 14 and 13 refrigerated host eggs per day kept at 26.3°C and 78.7% RH for 2 and 4 days of post-refrigeration without significant reduction, respectively. These results show that refrigeration of R. clavatus eggs can be a good method for mass rearing of the parasitoid, and the host eggs killed by cold storage can be supplemented in the field to boost field parasitism.     

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.