Jean-Pierre Nénon

Systemic Release of Herbivore-Induced Plant Volatiles by Turnips Infested by Concealed Root-Feeding Larvae Delia radicum L.

When attacked by herbivorous insects, many plants emit volatile compounds that are used as cues by predators and parasitoids foraging for prey or hosts. While such interactions have been demonstrated in several host–plant complexes, in most studies, the herbivores involved are leaf-feeding arthropods. We studied the long-range plant volatiles involved in host location in a system based on a very different interaction since the herbivore is a fly whose larvae feed on the roots of cole plants in the cabbage root fly, Delia radicum L. (Diptera: Anthomyiidae). The parasitoid studied is Trybliographa rapae Westwood (Hymenoptera: Figitidae), a specialist larval endoparasitoid of D. radicum. Using a four-arm olfactometer, the attraction of naive T. rapae females toward uninfested and infested turnip plants was investigated. T. rapae females were not attracted to volatiles emanating from uninfested plants, whether presented as whole plants, roots, or leaves. In contrast, they were highly attracted to volatiles emitted by roots infested with D. radicum larvae, by undamaged parts of infested roots, and by undamaged leaves of infested plants. The production of parasitoid-attracting volatiles appeared to be systemic in this particular tritrophic system. The possible factors triggering this volatile emission were also investigated. Volatiles from leaves of water-stressed plants and artificially damaged plants were not attractive to T. rapae females, while volatiles emitted by leaves of artificially damaged plants treated with crushed D. radicum larvae were highly attractive. However, T. rapae females were not attracted to volatiles emitted by artificially damaged plants treated only with crushed salivary glands from D. radicum larvae. These results demonstrate the systemic production of herbivore-induced volatiles in this host-plant complex. Although the emission of parasitoid attracting volatiles is induced by factors present in the herbivorous host, their exact origin remains unclear. The probable nature of the volatiles involved and the possible origin of the elicitor of volatiles release are discussed.

Intra- and interspecific host discrimination in two closely related egg parasitoids

Intraspecific host discrimination is frequently found in solitary parasitoids, but interspecific host discrimination, where female parasitoids recognize hosts already parasitized by females of other species, is rare. This particular behaviour appears to be adaptive only under specific circumstances. In this paper, we quantified intraspecific host discrimination in Anaphes n. sp. (Hymenoptera: Mymaridae), an endoparasitoid of the eggs of Listronotus oregonensis (LeConte) (Coleoptera: Curculionidae) and interspecific host discrimination toward eggs parasitized by Anaphes sordidatus (Girault), a sympatric species competing for the same resource in similar habitats. To examine host discrimination, choice experiments were used where the females had to choose between different categories of eggs (unparasitized, parasitized by Anaphes n. sp. or A. sordidatus). Superparasitism and multiparasitism were avoided in experiments where the female had a choice between unparasitized hosts and hosts parasitized by the same female, by a conspecific or by a female A. sordidatus. When all hosts available were parasitized, conspecific superparasitism occurred more often than self-superparasitism or multiparasitism. These results indicated that females Anaphes n. sp. were capable of self-, conspecific and interspecific discrimination. Self-discrimination followed recognition of an external marking while interspecific discrimination occurred mostly after insertion of the ovipositor. Interspecific discrimination could result from the recent speciation of these species and could be associated with a genotypic discrimination. This behavior appears to be adaptive because of the competition for common hosts between the two parasitoid species.

Life history comparison between two competitive Aleochara species in the cabbage root fly, Delia radicum: implications for their use in biological control

This study evaluates the efficacy of Aleochara bilineata Gyll and Aleochara bipustulata L. (Coleoptera: Staphylinidae) as biological control agents against the cabbage root fly, Delia radicum L. (Diptera: Anthomyiidae). Biological and demographic characters are documented and comparisons made between the two species. The following results were obtained:

Host-stage selection by Trybliographa rapae, a parasitoid of the cabbage root fly Delia radicum.

Host‐stage selection by Trybliographa rapae Westwood (Hymenoptera: Figitidae) was studied in choice and no‐choice experiments in the laboratory. The parasitoid was able to reproduce in first, second, and third instars of the cabbage root fly, Delia radicum L. (Diptera: Anthomyiidae), but oviposition occurred more frequently in third instars when all three developmental stages were offered simultaneously. Oviposition in third instars increased the rate of development of offspring and their body size, but did not alter sex ratio. Results are discussed in the light of predictions made by the theory of optimal host acceptance.

Intraspecific hyperparasitism in a primary hymenopteran parasitoid

In solitary parasitoids, in which only one individual can emerge per host, the adaptive value of conspecific superparasitism is a function of the survival probability of the egg laid by the superparasitizing female. In the few cases which these probabilities are compared, the oldest immature has an advantage over the other individuals. We measured the acceptance rate of parasitized hosts and survival rate of supernumerary larvae in Anaphes victus (Hymenoptera: Mymaridae) in relation to the interval between ovipositions. When this interval was 5–7 days, the first immature was at the prepupa and pupa stage respectively, and female Anaphes victus changed their oviposition behavior markedly. They killed the developing parasitoid of their own species before ovipositing in it. The progeny of these females, which are normally primary parasitoids, developed thereafter as hyperparasitoids. Indeed, in contrast with other species, the survival of the second females progeny increased with the time interval between ovipositions. This type of facultative intraspecific hyperparasitism is different from autoparasitism in Aphelinidae and has never been mentioned in other parasitoids; it would be adaptive if females of this short-lived species encounter low-quality patches.

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.

Intraspecific Variations in Host Discrimination Behavior in the Pupal Parasitoid Pachycrepoideus vindemmiae Rondani (Hymenoptera: Pteromalidae)

Abstract In solitary parasitoid species, superparasitism incurs a high cost because only one individual can emerge per host. While avoiding already-parasitized hosts seems advantageous, it requires an ability to discriminate between parasitized and unparasitized hosts. The ability to discriminate can be based on physical or chemical cues or signal associated either internally or externally with a given host. The type of stimuli used to recognize parasitized hosts generally depends on the features of these hosts but also on costs and benefits associated with reusing them. Some local factors such as mortality rate of females, host availability, and competition level can influence this trade-off. In species that occupy a large geographic range, local conditions may favor either external or internal mechanisms of host discrimination. We describe the behaviors associated with host discrimination in Pachycrepoideus vindemmiae Rondani (Hymenoptera: Pteromalidae), a solitary pupal parasitoid of cyclorraphous dipteran species. To detect potential intraspecific variability in their host discrimination behavior, we compared two P. vindemmiae populations originating from different geographical areas in France. Our results revealed different host discrimination strategies in both populations and indicated a potential trade-off between the speed and the accuracy of host discrimination. One population discriminated hosts externally (faster but less accurately), whereas the other discriminated internally (more slowly but more accurately). In our experimental conditions, these two strategies resulted in differences in the fitness gains of both populations. The ecological conditions that could have selected for such differences in the host discrimination and oviposition strategies of the two populations remain to be investigated.

Host location and discrimination mediated through olfactory stimuli in two species of Encyrtidae

The role of olfactory stimuli in host detection and evaluation was studied in two encyrtid Hymenoptera. The first, Epidinocarsis lopezi De Santis, is a monophagous parasitoid of the cassava mealybug Phenacoccus manihoti Matile‐Ferrero, itself feeding exclusively on cassava, Manihot esculenta Crantz. The second, Leptomastix dactylopii Howard, is a monophagous parasitoid of the Citrus mealybug, Planococcus citri Risso, but this latter is highly polyphagous. The behaviour of females of both parasitoids (attraction and locomotion) was compared in a tubular olfactometer for the odours of their respective hosts on cassava and poinsettia. Tests were made using: 1) healthy host‐plant alone; 2) host‐plant infested with unparasitized mealybugs; 3) unparasitized mealybugs only; 4) host‐plant infested with parasitized mealybugs and 5) parasitized mealybugs only. Only E. lopezi was attracted by the odour of the host‐plant alone, but both species were attracted by the odour of an infested host‐plant and that of unparasitized mealybugs. The odour of parasitized mealybugs, alone or on host‐plant, induced an undirected activity. The attraction of E. lopezi to the odour of the host‐plant alone could be linked to the monophagous diet of its host, whereas the attraction of the two species of parasitoids to the odours of infested host‐plants and unparasitized mealybugs could be due to the fact that both parasitoids are specialists. The behavioural response of both species to the odour of parasitized mealybugs revealed a new aspect in host discrimination: the identification of parasitized hosts could be partly mediated through olfactory stimuli, and not only through gustatory stimuli.

Intraspecific larval competition in two solitary parasitoids, Apoanagyrus (Epidinocarsis) lopezi and Leptomastix dactylopii.

Encyrtid Hymenoptera, Apoanagyrus (Epidinocarsis) lopezi De Santis and Leptomastix dactylopii Howard are strictly solitary endoparasitoids of mealybug larvae and adults, Phenacoccus manihoti Matile‐Ferrero and Planococcus citri Risso (Pseudococcidae), respectively. For both species, the survival rate of the supernumerary parasitoid according to the time interval between the two ovipositions has been determined by dissecting superparasitized hosts. For A. lopezi, the survival rate of the second parasitoid is at least equal to 30% when the time interval between the two ovipositions is less than 2 days, then it drops rapidly down to 10%. For L. dactylopii, the survival rate of the second parasitoid is 23% for a one hour interval between the two ovipositions and thereafter decreases very rapidly to reach a stable value of 7%. Differences between the two species might be partly due to the immunological defense reactions of their respective hosts. In the two Encyrtidae, supernumerary individuals are eliminated partly by physical attack between larvae at the L2 stage and partly by a physiological suppression of the second parasitoid by the first at the egg or larval stage. Earlier findings had shown that females of each species oviposit preferentially in hosts in which their progeny has the highest probability of survival: this would indicate that the adaptive value of conspecific superparasitism depends essentially on the survival rate of the second parasitoid in superparasitized hosts.

Costs of secondary parasitism in the facultative hyperparasitoid Pachycrepoideus dubius: does host size matter?

Although hyperparasitism frequently occur in parasitic insects, many aspects of this strategy remain unknown. We investigated possible fitness costs of hyperparasitism as influenced by host size. Our study was conducted with the facultative hyperparasitoid Pachycrepoideus dubius Ashmead (Hymenoptera: Pteromalidae), which parasitizes host species differing greatly in size. We compared some fitness traits (level of successful parasitism, development time, sex ratio and offspring size) of P. dubius developing on large secondary/primary (Delia radicum L. (Diptera: Anthomyiidae)/Trybliographa rapae Westwood (Hymenoptera: Figitidae)) or small secondary/primary host species (Drosophila melanogaster L./Asobara tabida Nees (Hymenoptera: Braconidae)). In no‐choice and choice experiments, P. dubius was able to develop on different stages of T. rapae (L2 (endophagous), L4 (ectophagous), and pupae) but that it preferred to parasitize unparasitized D. radicum pupae over pupae parasitized by T. rapae. Furthermore, in P. dubius, hyperparasitism was associated with fitness costs (lower level of successful parasitism, smaller adult size) and these costs were greater on the smallest host complex. We hypothesize that the size of D. melanogaster pupae parasitized by A. tabida may be close to the suboptimal host size for P. dubius beneath which the costs of hyperparasitism make this strategy nonadaptive. Hyperparasitism in terms of trade‐offs between host quality and abundance of competitors is discussed.