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International audience; Biological control of bruchid beetles, Callosobruchus maculatus (Fabricius) (Coleoptera: Bruchidae), infesting cowpea seeds, Vigna unguiculata (L.)Walpers (Fabaceae), can be performed via augmentative releases of Dinarmus basalis Rondani (Hymenoptera: Pteromalidae) parasitoid wasps. Females of the latter species are therefore likely to experience intense intraspecific competition: they should encounter numerous previously parasitized hosts but also conspecific competitors, with which they may fight to secure hosts on which to lay their eggs. Such contests might therefore disrupt biological control programs. Here, we studied aggressive behavior that D. basalis females show toward conspecific competitors and subsequent host exploitation strategies. We further investigated factors that classically affect contest intensity and outcomes in animals, such as the effect of ownership status, by manipulating the residency period before the intruder's arrival. In addition, we tested the effect of the size of female reproductive tissue (measured in terms of egg load) and the quality of the habitat previously experienced by females (either rich or poor in hosts). These two factors are expected to influence the value that females place on the host and therefore the costs they are willing to pay to win it. Finally, we discussed the consequences of agonistic behaviors on females' host exploitation strategies. Our results suggest that contest competition may actually enhance host control by favoring parasitoid dispersion, rather than disrupting it.

The parasitoid species, Dinarmus basalis, attacks bruchid beetle larvae concealed within hard seeds. Female wasps are able to discriminate the quality of their host but the position in the concealed host system where this discrimination takes place remains unclear. Under laboratory conditions, we investigated whether the cues were detectable: (1) at a medium or short distance from the seed, (2) on the seed, or (3) within the seed, at the point of contact with the host. In particular, we investigated whether internal or external markings are applied by the parasitoid and then recognized in a subsequent encounter or by a conspecific encountering the host. Detailed behavioral observations did not provide evidence of markings being applied onto the seed or used, but showed that host discrimination occurred on the basis of internal cues. The lack of evidence of external markings is unexpected and raises the question as to which circumstances in the parasitoid's ecology and life history could result in such a lack not causing a reduction of fitness.

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.

Dinarmus basalis (Rond) and Epelmus vuilleti (Crw) are two Hymenopteran species, which are solitary ectoparasitoids of bruchid larvae. In the presence of seeds of Vigna unguiculata (Walp) containing hosts parasitized by E. vuilleti, a high percentage of D. basalis females avoided multiparasitism whatever the age of the eggs or the larvae present on the host. The least avoidance was observed when the hosts were parasitized by E. vuilleti 30 min beforehand. This avoidance behavior is adaptive and is related to the low survival chances of the D. basalis larvae when they are in interspecific competition with E. vuilleti larvae. The analysis of the behavior of D. basalis demonstrated that the avoidance of multiparasitism could be due to the perception of two signals; an external signal deposited on the surface of the seeds during the E. vuilleti oviposition phase and an internal signal due to the presence of the eggs and larvae at the surface of the hosts. E. vuilleti females did not avoid multiparasitism and multiparasitized the hosts bearing D. basalis eggs or larvae. The behavior of E. vuilleti females was not disturbed by the presence of its competitor. Under these conditions of interspecific competition, the survival chances of E. vuilleti larvae were very high whatever the age of its competitor D. basalis. The two species of parasitoids could move in a column containing healthy seeds of V. unguiculata and patches with seeds containing parasitized or unparasitized larvae. The distribution of D. basalis females introduced into these columns depended on the host quality. They avoided the patches containing the hosts parasitized E. vuilleti and were found in the patches with healthy hosts. The behavior of E. vuilleti females was very different; the distribution of the females and the parasitism and multiparasitism rates were not affected by the quality of the hosts present in the patches. The adaptive significance of the behaviors of these two species was analyzed in relation to the survival chances of their offspring.

In the solitary ectoparasitoid, Dinarmus basalis (Hymenoptera: Pteromalidae), the occurrence of superparasitism according to the unparasitised host density, and the nature of the host(s) provided was investigated in laboratory studies. In this species superparasitism was observed whatever the experimental conditions used, but the degree of superparasitism depended on the density of its host, Bruchidius atrolineatus (Coleoptera: Bruchidae). Superparasitism was due to successive egg-laying phases on the same host. However, females were able to discriminate between unparasitised hosts and hosts parasitised from 8 h to 72 h beforehand by themselves or by conspecifics. There was no conclusive evidence that superparasitism in the presence of a host parasitised 30 min before was linked to an absence of host discrimination. Host discrimination in this species is achieved by host-quality markers. These are individual-specific markers since conspecific superparasitism rates were often higher than self superparasitism rates. One deterrent substance is emitted by the females during oviposition onto the egg or released by the 16 to 24 h-old egg itself. Another host-quality marker is associated with the presence of a larva on its host. On the other hand, host discrimination ability did not always imply avoidance of superparasitism. In D. basalis there exists a positive relationship between the survival probability of the second egg and the tendency to superparasitise, and superparasitism could therefore result in a significant fitness gain. Under our experimental conditions, D. basalis females exhibited a wide range of oviposition behavioural plasticity in relation to the parasitoid developmental stage, the type of superparasitism, and the encounter rate with unparasitised hosts.

Eupelmus vuilleti (Crw) is a solitary ectoparasitoid of several bruchid species which develop inside Leguminosae seeds. In this hymenopteran a preemergence learning process is responsible for a specific response of females toward the host and host-plant volatiles. This learning occurs after the imaginal molt, while the adult remains in the larval chamber of its host. The persistence of the response toward host and host-plant volatiles of females reared onBruchidius atrolineatus (Pic) larvae developing inVigna unguiculata (Walp) seed was investigated in this study. In the absence of any reinforcement, the response towardB. atrolineatus larvae persisted for 6 days after female emergence from the seeds, whereas the response towardV. unguiculata seed volatiles was maintained for 7 days. Our experiments showed that a later learning process also takes place inE. vuilleti. An ovipositional experience on a given plant-host complex enabled females to learn the stimuli from that complex. This learning occurred even if the complex experienced during oviposition differed from the one on which females were reared. After an ovipositional experience on a new plant-host complex, the response acquired through preemergence learning was maintained. These experiments demonstrate that theE. vuilleti females can memorize cues from several species during successive learning processes. The adaptive significance of these learning capacities, which confer an important behavioral plasticity to this generalist parasitoid, is discussed.

The response to different host and plant species odours was investigated in Eupelmus vuilleti (Crw). This hymenopteran is a solitary ectoparasitoid of several species of bruchids developing inside Leguminosae seeds. The locomotor behaviour of females reared on Bruchidius atrolineatus (Pic) larvae developing in Vigna unguiculata (Walp) seeds was analysed using a tubular olfactometer. Females showed a specific sensitivity to the semiochemicals emanating from the host and the seed species on which they had developed. Odours from V. unguiculata seeds were attractive to the parasitoid and stimulated their locomotor activity. Odours from Vigna radiata (Wil) seeds had no effect on the locomotor behaviour. Odours from B. atrolineatus larvae were attractive to the females whereas odours from Callosobruchus maculatus (Fab), another bruchid species, had no effect. By isolating the females from the seed and the host at different developmental stages, we found that the specific sensitivity observed resulted from an early adult learning. This learning which occurs before the emergence from the seed while the imago is in the larval chamber of its host is dependent on contact with the seed and the host larval remains.

Dinarmus basalis Rond. (Hymenoptera : Pteromalidae) et Eupelmus vuilleti Craw. (Hymenoptera : Eupelmidae) sont les deux principales especes d’ectoparasitoides solitaires de larves et de nymphes de Bruchidae en Afrique de l’Ouest. Des etudes anterieures effectuees sur ces parasitoides ont montre que D. basalis et E. vuilleti entraient en competition interspecifique. E. vuilleti est competitivement superieur a D. basalis et l’elimine quand les deux especes se developpent a densite initiale egale. Nous avons montre, dans les conditions du climat subtropical de type guineen que l’issue de la competition pouvait dependre du nombre de femelles de D. basalis exploitant le patch par rapport a celui de E. vuilleti. Quand la proportion initiale des concurrents est en faveur de D. basalis (75 % de D. basalis et 25 % de E. vuilleti), la descendance est majoritairement constituee de D. basalis. Ces resultats permettent d’expliquer en partie l’abondance de D. basalis par rapport a E. vuilleti dans les greniers de la zone subtropicale de type guineen. Dinarmus basalis Rond. (Hymenoptera : Pteromalidae) and Eupelmus vuilleti Craw. (Hymenoptera : Eupelmidae) are the two main solitary ectoparasitoids of larvae and pupae of Bruchidae in West Africa. Former studies carried out on these species showed that D. basalis and E vuilleti entered interspecific competition. E vuilleti is competitively higher than D. basalis and eliminates it when the two species develop with equal initial density. We showed under subtropical climate conditions in guinean zone that the exitence of the competition could depend on the number offemales of D. basalis exploiting the patch compared to that of E vuilleti. When the initial proportion of the competitors is in favour of D. basalis (75 % of D. basalis and 25 % of E vuilleti), the descent mainly consists of D. basalis. These results permitted to explain the abundance of D. basalis compared to E. vuilleti in guinean zone.

Eupelmus vuilleti (Hymenoptera; Eupelmidae) is a host feeding ectoparasitoid of fourth-instar larvae or pupae of Callosobruchus maculatus (Coleoptera; Bruchidae) infecting Vigna unguiculata seed and pods (Fabacae). Parasitoid females are synovigenic, i.e. they are born with immature eggs and need to feed from the host in order to sustain egg production. In this study, the role of sterols obtained through host feeding in parasitoid oogenesis are examined. Quantitative and qualitative analyses of the sterol contents in each partner of the tritrophic interaction show that a parasitoid female's larval sterol contents is sufficient to produce only 30% of the total number of eggs laid throughout a female's life cycle. In a second step, by manipulating the composition of the sterols hemolymph in the host, it is shown that cholesterol obtained through adult nutrition plays a crucial role in the eggs viability but does not affect the egg production quantitatively. This result has important implications for understanding both the nutrient allocation strategy in this species and the impact of cholesterol in parasitoid reproduction.

Eupelmus vuilleti ( Crawford) is an ectoparasitoid of the seed-eating beetle Bruchidius atrolineatus (Pic), which is an important pest of stored cowpea, Vigna unguiculata Walp, seeds in West Africa. Herein, we investigated the dispersal abilities of females within columns of seeds to assess the potential of E. vuilleti as a biological control agent of bruchids in cowpea granaries. The influence of host presence together with the 2 abiotic factors light and gravity on parasitoid movement and parasitization efficiency were analyzed. E. vuilleti females were able to travel through large seed masses and parasitize hosts located at the end of the seed column opposed to their introduction zone. Parasitoid movement was stimulated by light. E. vuilleti females exhibited a negative geotropism. Females introduced at the bottom of the seed column dispersed more and parasitized more hosts than females introduced at the top. Host presence had some influence on the dispersal of the parasitoids within the seed column at a host density of 10 infested seeds for 16,000-18,000 uninfested seeds. This depended on female introduction zone because gravity was the major factor influencing dispersal. The possible applications of these results for biological control of bruchids in cowpea granaries are discussed.

Host selection in parasitic hymenopterans involves the use of a wide array of chemical signals emanating not only from the host itself but also from its food (Lewis & Martin, 1990; Tumlinson et al., 1993). For each tritrophic combination, only some particular cues which have proved to be reliably associated with the presence of hosts, are used (Vet & Dicke, 1992). Numerous studies show that the use of chemical cues can be genetically fixed but that learning often plays a predominant role (van Alphen & Vet, 1986; Vet & Groenewold, 1992; Tunings et al., 1993). By learning profitable cues, females can adapt to spatial and temporal variations in their foraging environment (Turlings et al., 1989; Lewis et al., 1990). However, little is known about how different learning abilities develop in different species and it is still unknown whether these differences could be better explained by differences in ecological niches or by taxonomic status of the species.

Bruchidius atrolineatus (Pic) is a tropical beetle (Coleoptera Bruchidae) that develops during the larval and pupal stages in the seeds of a legume Vigna unguiculata (Walp). Two species of Hymenoptera, Dinarmus basalis (Rond) and Eupelmus vuilleti (Craw), solitary ectoparasitoids of the larvae and pupae of B. atrolineatus, were introduced successively in the presence of their hosts, varying the interval between the two introductions. When D. basalis females were introduced 24 h, 3 days or 7 days after E. vuilleti, multiparasitism was low. The females had low fecundity, and their eggs were not distributed randomly over the different available hosts. When E. vuilleti females were introduced second, they oviposited on the different hosts availabe and did not avoid multiparasitism. The presence of hosts already parasitised by D. basalis increased the reproduction of E. vuilleti, and the fecundity of the females was higher than in control batches with E. vuilleti alone. E. vuilleti seems capable of detecting the ovipositor shafts drilled by the D. basalis females, and by introducing its own ovipositors killing the D. basalis eggs or larvae. When interspecific competition was occurring the number of E. vuilleti adults emerging from the seeds was no different from that observed in control batches with E. vuilleti alone, and there were always fewer D. basalis adults than in control batches (D. basalis alone). This interspecific competition reduces the influence of the two parasitoids in the biological control of bruchid populations.