Recognition of heterospecific parasitism: Competition between Aphidiid (Aphidius ervi) and Aphelinid (Aphelinus asychis) parasitoids of Aphids (Hymenoptera: Aphidiidae; Aphelinidae)

The solitary parasitoids Aphidius erviHaliday (Hymenoptera: Aphidiidae) and Aphelinus asychisWalker (Hymenoptera: Aphelinidae) attacked but generally did not oviposit in pea aphids parasitized by the other species. Wasps selectively oviposited in unparasitized hosts when given a choice. Host discrimination depended on the recognition of internal cues. Females of A. asychiseither could not recognize or ignored A. ervi'sexternal host marking pheromone. Under most conditions, A. ervisurvived in superparasitized hosts, killing competing A. asychislarvae by physical attack and possibly physiological suppression. The outcome of larval competition was not affected by oviposition sequence or age difference between larvae; A. asychissurvived only when it had substantially completed larval development before the host was superparasitized by A. ervi.It is suggested that competition for host resources incurs a cost, for the winner in terms of reduced size or increased development time and for the loser in terms of lost progeny and searching time. Consequently, heterospecific host discrimination can be functional. Internal, and probably general, cues enable wasps to recognize and avoid oviposition in hosts already parasitized by an unrelated species.     

Effects of mummy consumption on fitness and oviposition site selection on Harmonia axyridis

Intraguild predation (IGP) has been commonly reported between predators and parasitoids used as biological control agents as predators consuming parasitoids within their hosts. However, the effect of parasitoid–mummy consumption on the fitness of the predator and subsequent oviposition site selection have not been well studied. In our study, we conducted two laboratory experiments to examine the influence of Aphidius gifuensis Ashmead (Hymenoptera: Braconidae) mummies as prey on fitness and subsequently oviposition site selection of Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae). Results indicate that when H. axyridis was reared on A. gifuensis mummies only, its larval development was prolonged, and body weight of the 4th instar larvae and newly emerged adults, and fecundity decreased. Moreover, H. axyridis did not exhibit oviposition preference on plants infested with unparasitized aphids or aphids parasitized for shorter than 9 days. However, compared with plants with mummies (parasitized ≥9 days), H. axyridis laid more eggs on plants with unparasitized aphids. In contrast, H. axyridis previously fed with A. gifuensis mummies did not show a significant oviposition preference between plants with unparasitized aphids and those with mummies (parasitized ≥9 days). Overall, our results suggest that mummy consumption reduced the fitness of H. axyridis. Although H. axyridis avoided laying eggs on plants with A. gifuensis mummies, prior feeding experience on A. gifuensis mummies could alter the oviposition site preference. Thus, in biological control practice, prior feeding experience of H. axyridis should be carefully considered for reduction of IGP and increase of fitness of H. axyridis on A. gifuensis.     

Oviposition avoidance of parasitized aphid colonies by the syrphid predator Episyrphus balteatus mediated by different cues

Oviposition decisions made by members of a guild of natural enemies can have evolved to avoid intraguild predation, potentially avoiding the disruption of the extraguild prey control. We have studied the oviposition preference of the aphidophagous predator Episyrphus balteatus De Geer (Diptera: Syrphidae) within colonies of Myzus persicae Sulzer (Hemiptera: Aphididae) in the presence of two developmental stages of the aphid parasitoid Aphidius colemani Viereck (Hymenoptera: Aphidiidae). Results from a greenhouse choice experiment showed that E. balteatus females lay significantly fewer eggs in colonies with mummified aphids than in unparasitized colonies. Colonies of parasitized, but not yet mummified did not contain significantly fewer eggs than colonies with unparasitized aphids. In three no-choice experiments, we assessed stimuli coming from aphid honeydew, from the aphids themselves and also from extracts of the aphid bodies, and all of these stimuli mediate the discrimination of mummified aphids from healthy aphids. To a lesser extent these stimuli also contribute to the discrimination against aphids that are parasitized but not yet mummified. These results suggest that the effects of these two species could be complementary for the control of M. persicae, since the species that acts as an intraguild predator, E. balteatus, avoids ovipositing on aphid colonies parasitized by the intraguild prey, A. colemani.     

Efficiency,intrinsic competition and interspecific host discrimination of Copidosoma desantisi and Trichogramma evanescens,two parasitoids of Phthorimaea operculella

Trichogramma evanescens West. (Hymenoptera: Trichogrammatidae) and Copidosoma desantisi Annecke & Mynhardt (Hymenoptera: Encyrtidae) are potential parasitoids of the potato tuber moth (PTM), Phthorimaea operculella (Zeller) (Lepidoptera: Gelechiidae) in Egypt. Discrimination of a parasitized host from an unparasitized host would prevent wasting of time, eggs and reduce competition with conspecifics or heterospecifics. Therefore, we evaluated interspecific host discrimination, multiparasitism and intrinsic competition between the two wasp species. In a choice test, females of T. evanescens showed high interspecific host discrimination only when they were offered 2-day-old C. desantisi parasitized and unparasitized PTM eggs. In contrast, C. desantisi showed high host discrimination and preferred unparasitized eggs to PTM eggs harboring 2-h- or 2-day-old T. evanescens’ eggs. We also evaluated the effect of different introduction sequences on the efficacy of the two wasps. Dissection data indicated that the two parasitoids had a negative impact on each other. There was a significant reduction in the total number of deposited eggs as well as total number of parasitized hosts by each parasitoid. Regarding the rearing experiment, the total number of T. evanescens-induced black eggs or C. desantisi formed mummies in combined treatments was significantly lower than in single parasitoid treatments (control). Moreover, C. desantisi was inferior and did not develop from any multiparasitized host regardless of oviposition order. It was suggested that combined release of the two wasps would not elevate rate of parasitism over that of single parasitoid treatments and competition between them would reduce their efficacy.     

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.     

Impact of mating status on egg-laying and superparasitism behaviour in a parasitoid wasp

Most parasitoid female wasps can distinguish between unparasitized and parasitized hosts and use this information to optimize their progeny and sex allocation. In this study, we explored the impact of mating on oviposition behaviour (parasitism and self‐ and conspecific superparasitism) on both unparasitized and already parasitized hosts in the solitary parasitoid wasp Eupelmus vuilleti (Crw.) (Hymenoptera: Eupelmidae). Virgin and mated females had the same oviposition behaviour and laid eggs preferentially on unparasitized hosts. The sex ratio (as the proportion of females) of eggs laid by mated females in parasitism and conspecific superparasitism was 0.67 ± 0.04 and 0.57 ± 0.09, respectively. Likewise, females laid more eggs in conspecific superparasitism than self‐superparasitism under our experimental conditions. These experiments demonstrate that E. vuilleti females can (i) discriminate between unparasitized and parasitized hosts and adapt the number of eggs they lay accordingly, and (ii) probably discriminate self from conspecific superparasitized hosts. Finally, mating does not appear to influence the host discrimination capacity, the ovarian function, or the oviposition behaviour.     

Costs of reproduction and geographic variation in the reproductive tactics of the mosquito Aedes triseriatus

Intraspecific host discrimination is widespread in solitary parasitoids whose adult females forage for and evaluate host suitability, whereas interspecific discrimination is less common. In some parasitoid species, mostly Diptera and Coleoptera, the larva performs the last step of host searching. It has been suggested that host discrimination will rarely occur in such host-seeking larvae because their low mobility results in a low host encounter rate. We determined the extent to which the larvae of Aleochara bilineata Gyllenhal (Coleoptera: Staphylinidae), a solitary parasitoid of aggregated Diptera pupae: (1) discriminated between unparasitized hosts and hosts parasitized by conspecifics; (2) used semiochemical cues to discriminate; (3) were influenced by life expectancy, presence of conspecifics and host availability in their host acceptance decision; and the extent to which (4) A. bilineata and A. bipustulata L., a species exploiting the same hosts and occurring sympatrically, showed interspecific host discrimination. A. bilineata larvae were able to discriminate between unparasitized hosts and hosts parasitized by conspecifics in a choice experiment. Such behavior has never previously been described for a coleopteran parasitoid or for a parasitoid species whose larvae perform host searching. Host discrimination in this species was not based on the presence of visual or tactile cues (e.g., entrance holes) but rather on chemical cues. The life expectancy of A. bilineata larvae was significantly shorter in the presence than in absence of hosts, and older larvae had lower parasitism success than young larvae in a 24-h experiment. However, the host acceptance decision of A. bilineata larvae was not influenced by larval age or the presence of conspecifics when the ratio of hosts per larva was greater than or equal to 1. When hosts were scarce, the degree of superparasitism increased significantly with the number of foraging conspecifics and the age of the larvae. Both species of Aleochara showed intra- and interspecific host discrimination in a choice experiment. In contrast to A. bipustulata, A. bilineata larvae more frequently parasitized hosts parasitized by A. bipustulata than those parasitized by conspecifics. We suggest that host discrimination will be frequent in solitary parasitoids with host-seeking larvae when hosts are aggregated. Received: 4 June 1998 / Accepted: 1 September 1998     

Host-range study about four aphid parasitoid species among 16 aphid species for constructing banker–plant systems

To provide fundamental information for the biological control of aphids in vegetable greenhouses, we compared the host ranges of four aphid parasitoid species, Aphidius colemani Viereck, Aphidius gifuensis Ashmead, Diaeretiella rapae (M’Intosh), and Ephedrus nacheri Quilis (Hymenoptera: Braconidae: Aphidiinae). The acceptability as host of 11 vegetable-pest aphids, Acyrthosiphon pisum (Harris), Aphis craccivora Koch, Aphis gossypii Glover, Aulacorthum solani (Kaltenbach), Brevicoryne brassicae (Linnaeus), Chaetosiphon fragaefolii (Cockerell), Lipaphis erysimi (Kaltenbach), Macrosiphoniella sanborni (Gillette), Macrosiphum euphorbiae (Thomas), Myzus persicae (Sulzer), and Uroleucon formosanum (Takahashi), in addition to five aphid species, Melanaphis sacchari (Zehntner), Rhopalosiphum maidis (Fitch), Rhopalosiphum padi (Linnaeus), Schizaphis graminum (Rondani), and Sitobion akebiae (Shinji) (Hemiptera: Aphididae) that serve as alternative hosts in banker–plant systems for the four aphid parasitoid species, were investigated. A newly emerged pair of parasitoid adults were provided to 100 aphids of each species on caged host plants in a 25 °C chamber for 24 h. The numbers of mummified aphids and emerged adults were counted in 10 trials for each aphid species. Aphidius colemani, A. gifuensis, D. rapae and E. nacheri parasitized four, two, three, and eight pest species, respectively, and four, three, three, and five alternative host species, respectively. Ephedrus nacheri had the broadest host range among the four species, and all the four species parasitized M. persicae, R. maidis, and S. graminum. This information will be useful for selecting candidate of biological control agents for aphids and for constructing banker–plant systems.

     

Functional responses of aphid parasitoids,Aphidius colemani (Hymenoptera: Braconidae) and Aphelinus asychis (Hymenoptera: Aphelinidae)

We evaluated the functional responses of two aphid parasitoids: Aphidius colemani on the green peach aphid Myzus persicae (Hemiptera: Aphididae), and Aphelinus asychis on M. persicae and the potato aphid, Macrosiphum euphorbiae (Hemiptera: Aphididae). Parasitoid oviposition occurred at host densities of 5, 10, 20, 30, 50, 80 or 100 aphids for A. colemani and 5, 10, 20, 30 or 50 aphids for A. asychis. More M. persicae were parasitized by A. colemani than by A. asychis at an aphid density of 50. Among the three types of functional response, type III best described the parasitoid response to the host densities both in A. colemani and A. asychis. The estimated handling time was shorter for A. colemani than for A. asychis (0.017 and 0.043 d, respectively). The proportion of aphids that were parasitized exhibited the same characteristic curve among the three host-parasitoid combinations: a wave form that appeared to be a composite of a decelerating (as in type II) response at low host density and an accelerating-and-decelerating (as in type III) response at medium to high host density. We hypothesize that the novel host species (and its host plant), density-dependent superparasitism, and/or density-dependent host-killing may have induced the modified type III response.     

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.     

Interspecific host discrimination and competition in Apoanagyrus (Epidinocarsis) lopezi and A.(E.)diversicornis, parasitoids of the cassava mealybug Phenacoccus manihoti

Abstract.

• 1 Choice experiments on interspecific host discrimination in A.lopezi and A. diversicornis were carried out on discs of cassava leaf containing four hosts (P.manihoti) that had been parasitized by the other species and four unparasitized hosts.
• 2 A.lopezi accepted both host types equally for oviposition, whereas A.diversicornis accepted fewer hosts that had been parasitized by A.lopezi than unparasitized ones. A.diversicornis is therefore capable of interspecific host discrimination, but such a capability was not demonstrated for A.lopezi.
• 3 Survival probability in singly parasitized hosts was 0.85 for both parasitoid species. When the time interval between ovipositions was 2 h or less, survival in multiparasitized hosts was 0.68 for A.lopezi and 0.17 for A.diversicornis, irrespective of priority. Increasing A.lopezi priority to 24±2h did not increase A.lopezi survival. A.diversicornis survival, however, increased to 0.43 when A.diversicornis was given 24 ± 2 h priority. A.diversicornis eggs took 19 h longer than A.lopezi eggs to hatch. This could explain the difference in competitive abilities in multiparasitized hosts.
• 4 The observed difference in host selection behaviour between A.lopezi and A.diversicornis is in accordance with the different benefits of multiparasitism: A.lopezi gains more than A.diversicornis because of its superior within-host competitive abilities.
• 5 Neither species avoided multiparasitism completely. The low survival probability of A.diversicornis in multiparasitized hosts may partly explain its failure to establish when introduced into Africa as part of a biological control programme of P.manihoti.

     

Cues of intraguild predators (Harmonia axyridis) and different temperatures alter foraging and oviposition strategies of the parasitoid Aphidius gifuensis

Organisms live in complex systems where they simultaneously balance responses to multiple factors. Insects are dealing with indirect pressures from predators under the influence of temperature. Currently, little is known about how they balance these factors. Under laboratory conditions, we investigated the effects of the predator Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae) cues and temperature (23, 28, and 33 °C) on the foraging and oviposition behavior of the parasitoid Aphidius gifuensis Ashmead (Hymenoptera: Braconidae) using the aphid Myzus persicae (Sulzer) (Hemiptera: Aphididae) as a host. In all trials, the frequency and duration of stinging, walking, or stationary behavior of A. gifuensis were not influenced by H. axyridis cues. However, in trials with predator cues, A. gifuensis produced less offspring than trials without predator cues. On the other hand, the frequency of A. gifuensis stinging, walking, or stationary behavior, and the number of A. gifuensis larvae increased with rising temperature. Specially, when the predator cues were present, parasitoid responded to rising temperatures by increasing frequency of each behavior and an increase in offspring number. Our findings indicate that A. gifuensis might show flexible foraging and oviposition behavior in response to H. axyridis associated cues and different temperatures. Thus, the effects of both temperature and H. axyridis cues should be considered when assessing the foraging outcomes of A. gifuensis prior to combining these two enemies in aphid management practice.     

Influence of egg load and oviposition time interval on the host discrimination and offspring survival of Anagyrus pseudococci (Hymenoptera: encyrtidae), a solitary endoparasitoid of citrus mealybug, ++Planococcus citri (Hemiptera: pseudococcidae)

Oviposition and host discrimination behaviour of unmated Anagyrus pseudococci (Girault), an endoparasitoid of the citrus mealybug Planococcus citri (Risso), were investigated in the laboratory. Female parasitoids were able to discriminate between parasitized hosts and healthy ones. The mean number of ovipositions was significantly higher in unparasitized than in parasitized hosts. Conspecific-superparasitism occurred more often than self-superparasitism. Changes in consecutive ovipositions over three hours by A. pseudococci suggested that egg load influenced the discrimination behaviour of the parasitoids, with females which had low egg loads mostly avoiding oviposition in already parasitized hosts at time intervals ranging from 0 h to 96 h, and distributing their eggs in the high quality (unparasitized) hosts. The parasitized hosts were rejected more commonly through antennal perception of external markers than during ovipositor probing which could have encountered internal markers but this relationship changed with increasing time after oviposition. The parasitoid's oviposition rate in unparasitized and conspecific-parasitized hosts varied at the different oviposition time intervals when the females had fewer eggs in the ovaries. Percentage emergence of parasitized offspring was not significantly influenced by whether they developed in single or superparasitized mealybugs. The significance of host discrimination by A. pseudococci is discussed.     

Infectivity of entomopathogenic nematode Steinernema carpocapsae Pocheon strain (Nematoda: Steinernematidae) on the green peach aphid Myzus persicae (Hemiptera: Aphididae) and its parasitoids

Infectivity of entomopathogenic nematode (EPN) Steinernema carpocapsae Pocheon strain on the green peach aphid Myzus persicae and its parasitic wasps (e.g., Aphidius colemani, Aphidius gifuensis and Diaeretiella rapae) was evaluated under laboratory conditions. Infective juveniles (IJs) of S. carpocapsae Pocheon strain had low infectivity against nymph and adult stages of M. persicae, showing 2% and 6.7% of mortality, respectively. Application of the EPNs had little effect on mummies caused by the three parasitoid species, allowing them to remain intact. No IJ invaded the host, regardless of EPN application rate. The parasitoid emergence from mummies ranged from 80% to 85% in the presence of EPN while 79–86% was recorded in the absence of EPN. However, the presence of the IJs reduced oviposition by the three parasitoid species, decreasing the rate up to 59% when the nematodes were applied before parasitoid release, while little difference in oviposition was observed when nematodes were applied after parasitoid release.     

Comparison of Foraging Behavior, Interspecific Host Discrimination, and Competition of Encarsia formosa and Amitus fuscipennis

The foraging behavior of Amitus fuscipennis MacGown & Nebeker and Encarsia formosa Gahan was studied on tomato leaflets with 20 Trialeurodes vaporariorum (Westwood) larvae in the first or third stage. Ten of the whitefly larvae were previously parasitized and contained a conspecific or a heterospecific parasitoid egg or larva. The host type (host stage and/or previous parasitization) did not influence the foraging behavior of either parasitoid species. The residence time on these tomato leaflets was about 0.9 h for A. fuscipennis and 1.9 h for E. formosa. Amitus fuscipennis hardly stood still and fed little, while E. formosa showed extensive standing still and feeding. As a result, the time walking while drumming was similar for both parasitoid species. The numbers of host encounters and ovipositions per leaflet were similar for both parasitoid species. However, the residence time of A. fuscipennis was half as long as that of E. formosa so the rate of encounters and ovipositions was higher for A. fuscipennis. Amitus fuscipennis is more efficient in finding and parasitizing hosts under these conditions. The walking activity and host acceptance of the synovigenic E. formosa diminished with the number of ovipositions, but not those of the proovigenic A. fuscipennis. Encarsia formosa is egg limited, while A. fuscipennis is time limited because of its short life span and high egg load. Both parasitoid species discriminated well between unparasitized larvae and self-parasitized larvae, but discriminated poorly those larvae parasitized by a conspecific and did not discriminate larvae parasitized by a heterospecific. Self-superparasitism, conspecific superparasitism, and multiparasitism were observed for both parasitoid species. Superparasitism always resulted in the emergence of one parasitoid and multiparasitism resulted in a higher emergence of one parasitoid of the species that had parasitized first. The data suggest that A. fuscipennis is a good candidate for use in biological control of high-density spots of T. vaporariorum when we consider its high encounter and oviposition rate.     

Dynamic host-feeding and oviposition behavior of an aphid parasitoid <Emphasis Type="Italic">Aphelinus asychis</Emphasis>

In order to maximize the lifetime reproductive success of parasitoids, they should be induced to dynamically accept individual hosts that have different suitability for oviposition. Parasitoids tend to exhibit higher host-selective behavior when their egg load is limited, and are less selective if they are facing time constraints. Here, we evaluated the effects of parasitoid age on egg load, fecundity and host instar preference of a honey-fed aphid parasitoid, Aphelinus asychis Walker (Hymenoptera: Aphelinidae). Host selective experiment was conducted to measure host-preference of honey-fed A. asychis females at different ages, using the second and fourth instars of the green peach aphid Myzus persicae as their hosts. The results showed that the choice of host-instar for oviposition was significantly influenced by the parasitoid age. Honey-fed parasitoids in the age groups of 1, 5, 10 and 20 days tended to parasitize predominantly second-instar aphids, whereas 15-days old parasitoids showed no significant preference of host instars. On the other hand, host-feeding preference was not affected by parasitoid age. Parasitoid females of all ages preferred younger aphids to older aphids. This result could help evaluate the effectiveness of A. asychis for biological control of M. persicae when they encountered mixed-instar aphids in the field. In addition, the results might be helpful in assessing the host killing effects of other host-feeding parasitoids.     

Parasitoid spectrum (Hymenoptera: Braconidae; Aphelinidae) of the soybean aphid Aphis glycines (Homoptera: Aphididae) in Japan and Indonesia (Java and Bali)

Aphidiine and aphelinid parasitoids collected from the soybean aphid, Aphis glycines, on Glycine max in Japan and Indonesia (Java and Bali) were identified to clarify the parasitoid spectrum of the aphid there. Nine parasitoid species from Japan (Aphidiinae: Aphidius gifuensis, Aphidius sp., Binodoxys communis, Diaeretiella rapae, Lipolexis gracilis, Lysiphlebia japonica; Aphelinidae: Aphelinus asychis, A. gossypii, A. varipes) and two parasitoid species from Indonesia (B. communis, A. gossypii) were found to be associated with A. glycines.     

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 (attaction 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.     

Oviposition behaviour ofEphedrus cerasicola [Hym.: Aphidiidae] parasitizing different instars of its aphid host

The parasitoidEphedrus cerasicola Starý oviposited in all 4 nymphal instars and in newly moulted adults ofMyzus persicae (Sulzer). The different host categories were offered with no choice. The duration of an oviposition increased with the age of nymphs, being about 13, 18, 21, 22, and 17 s from 1 st instars to adults, respectively. Observations of number of stabbing attacks prior to oviposition, percent of the encounters not resulting in oviposition, time from first encounter to oviposition, handling time and aphid defensive behaviour also indicated that 1 st instarM. persicae are most easily parasitized. The behaviour ofE. cerasicola in encounters with unparasitized and parasitized hosts, suggested that the parasitoid could discriminate. In encounters with parasitized 1 st to 4th instar aphids,E. cerasicola used only the antennae in 80% of the encounters that resulted in discrimination.     

Behavioural and physiological responses to conflicting oviposition stimuli in a synovigenic parasitoid

The behavioural and physiological responses of the synovigenic parasitoid, Dinarmus basalis (Rond.) (Hymenoptera: Pteromalidae), were investigated whilst the wasps were submitted to conflicting oviposition stimuli: an environment which induced them to restrict egg-laying and a physiological factor, egg-load pressure, which induced them to lay eggs. When individual females depleted patches containing hosts parasitized 24 h or 72 h beforehand, they laid preferentially on unparasitized hosts and the number of eggs laid decreased significantly. Although they remained for several days in such patches, egg-load pressure did not lead them to modify their strategy (i.e. to accept all the available hosts including the already parasitized ones). In this synovigenic species, the oviposition constraint resulted in the retention of oocytes. Dissection and histological observation showed that the unlaid eggs are resorbed, a reproductive regulatory process which has seldom previously been reported in the presence of unparasitized hosts. This physiological response was induced by the mere presence of the parasitized hosts rather than their density. The females’ response was also influenced by the presence of conspecifics. They still avoided laying on parasitized hosts but the number of eggs laid daily became comparable to that observed in patches containing only unparasitized hosts. Thus, D. basalis females adapted their strategy in response to their environment, in which the most significant factor seemed to be the quality of the hosts provided, and their physiology. The originality and the importance of these results are discussed.