Interspecific Relationships Between the Solitary Ectoparasitoid, Eupelmus vuilleti (Crw.) (Eupelmidae), and Its Sympatric Species, Dinarmus basalis (Rond.) (Pteromalidae), in the Presence of Their Host, Callosobruchus maculatus Pic (Coleoptera Bruchidae)

The eupelmid Eupelmus vuilleti CRW. and the pteromalid Dinarmus basalis Rond. are sympatric in West Africa. The reproduction of E. vuilleti and D. basalis females was analyzed in 10, 20, and 40 host patches when present alone or together. In 10- and 20-hosts patches, the presence of E. vuilleti affected the reproduction and offspring survival chances of D. basalis. In 40-hosts patch, only the offspring survival of D. basalis was affected by the presence of E. vuilleti. In contrast, the presence of D. basalis did not influence the reproduction of E. vuilleti and the survival chances of its offspring at all host densities tested. Multiparasitism did not occur at random. In a choice test, E. vuilleti showed a preference for hosts previously parasitized by D. basalis on healthy hosts. This attraction is mediated by chemical markers deposited by D. basalis females on the surface of the seed. At high host density, the likelihood of encountering a seed containing a host previously parasitized by D. basalis was lower for E. vuilleti, affecting moderately D. basalis reproductive success.     

Marking Behavior and Discrimination of Concealed Hosts by the Ectoparasitoid, Dinarmus basalis Rond. (Hym. Pteromalidae)

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.     

Analysis of multiparasitism by Eupelmus vuilleti (Craw) (Eupelmidae) and Dinarmus basalis (Rond) (Pteromalidae) in the presence of one of their common hosts,Bruchidius atrolineatus (Pic) (Coleoptera Bruchidae)

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.     

Influence of two successive learning processes on the response ofEupelmus vuilleti Crw (Hymenoptera: Eupelmidae) to volatile stimuli from hosts and host plants

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.     

Self-, Intra-, and Interspecific Host Discrimination in Telenomus busseolae Gahan and T. isis Polaszek (Hymenoptera: Scelionidae), Sympatric Egg Parasitoids of the African Cereal Stem Borer Sesamia calamistis Hampson (Lepidoptera: Noctuidae)

Oviposition behavior, intra- and interspecific host discrimination, and super- and multiparasitism by the scelionids Telenomus busseolae and T. isis were studied using batches of eggs of the noctuid stalk borer Sesamia calamistis as the host. Both Telenomus species were able to discern eggs already parasitized. As a result, self-superparasitism was only 4.0% for T. busseolae and 5.8% for T. isis. Likewise, intraspecific superparasitism was avoided by both species and was significantly higher for T. busseolae than T. isis, and higher when parasitized eggs were offered immediately (0 h) and after 48 h than after 24 h; apparently, the recognition of parasitized eggs at 24 and 48 h was based on the presence of parasitoid larvae rather than a specific marking substance. Multiparasitism was avoided if the female had a choice between unparasitized and parasitized eggs. In a choice experiment, it was 10.2 and 2.5% for T. busseolae and T. isis, respectively. In a nonchoice experiment, multiparasitism did not vary between species and time treatment; it was low, varying between 9.6 and 24.1%. In the 0-h treatment, T. busseolae always outcompeted T. isis, accounting for 63.4 or 91.7% of the offspring, when T. isis or T. busseolae, respectively, was the first species. In the 24-h treatment, the first species emerged more often than the second did but the host egg mortality, i.e., eggs from which neither borer larvae nor parasitoid emerged, was >40%. Whereas T. busseolae is ubiquitous in Africa, T. isis has never been reported from eastern Africa, and it has been proposed for introduction against the prevailing noctuid pest Bussoela fusca. Based on earlier life table and host suitability studies and on the present findings, it is concluded that T. isis would establish in the midaltitudes but not the highlands of eastern Africa.     

Host selection by virgin and inseminated females of the parasitic wasp, Dinarmus basalis (Pteromalidae, Hymenoptera)

1. Fitness is related to reproduction and survival. There apparently exists a negative correlation between the numbers of male and female offspring. There also exists a trade-off between survival and reproduction. This paper investigates optimal decisions with the reproduction and survival trade-off in host selection by wasps.
2. Whereas inseminated female wasps could manipulate the sex of their offspring, virgin females produced only male offspring. I surveyed behavioural differences and the consequences of oviposition by inseminated and virgin females of a solitary parasitic wasp in host choice situations.
3. Two host types were available at the same time to both inseminated and virgin female wasps: one (a 17-day-old host in one bean) presenting difficulties for the laying of eggs, but more benefits for the offspring and the other (five 12- or 13-day-old hosts in one bean) easier for the female wasp for laying of eggs but less beneficial for the offspring.
4. Inseminated female wasps chose more 17-day-old hosts than 12-day-old hosts, but more 13-day-old hosts than 17-day-old hosts in each pair-wise choice. Virgin females chose the smaller hosts in both situations.
5. Virgin females, having greater longevity than inseminated females, laid larger numbers of eggs than the inseminated females during their lifetime by adopting an energy-saving host choice that had little effect on male offspring fitness.     

Host plant conspicuousness and the distribution of eggs and larvae in the butterfly,Anthocharis scolymus (Lepidoptera: Pieridae)

We investigated how the distribution pattern of eggs and larval on the host plant, Turritus glabra, was influenced by the oviposition behavior of the pierid butterfly Anthocharis scolymus. Females searched for the host plants visually and they frequently approached taller host plants with sparse surrounding vegetation. After encountering host plants, oviposition behavior of females was independent of host plant characteristics such as height, density, and type of surrounding vegetation. A female laid eggs singly on a host plants. Most females appeared to lay their eggs regardless of the presense of eggs on the host plant. Consequently egg and larva tended to be abundant on conspicuous host plants as measured by height or relative isolation from other plants. However, overcrowding of eggs on an individual host decreased the survival rate of larvae.     

Interactions of the Males and Females of Three Sympatric Drosophila virilis-Group Species, D. montana, D. littoralis, and D. lummei, (Diptera: Drosophilidae) in Intra- and Interspecific Courtships in the Wild and in the Laboratory

Four species of the Drosophila virilis group, D. montana, D. littoralis, D. lummei, and D. ezoana, occur sympatrically in several locations in northern Europe. Courtship interactions between the flies of the three first-mentioned species were observed at malt baits in Kemi, northern Finland, to find out how the flies of different species recognize conspecific individuals and how interspecific courtships differ from intraspecific ones in the wild. Intraspecific courtships (including females of different reproductive stages) and interspecific courtships were also videotaped and analyzed in laboratory. In the wild the males courted both conspecific and allospecific females, even though the species varied in how much the males were attracted to females of different species. Interspecific courtships usually broke off when the male touched the female or when the male and/or the female vibrated his/her wings, producing acoustic cues. In the laboratory males courted conspecific females irrespective of the reproductive stage of the female, even though the courtships directed toward immature and fertilized females usually included only orienting and touching (no licking and singing). D. littoralis, and very rarely D. montana and D. lummei, males courted also allospecific females. In the few interspecific courtships between these three species, where the male proceeded to singing, females responded to male singing by vibrating their wings. This ended the courtship. It is suggested that both the chemical cues affecting female attractivity and the acoustic signals of males and females, which are produced by wing vibration, function in maintaining sexual isolation between these three species.     

Instar-specific defense of the pea aphid,Acyrthosiphon pisum: Influence on oviposition success of the parasiteAphelinus asychis (Hymenoptera: Aphelmidae)

The parasite Aphelinus asychisWalker (Hymenoptera: Aphelinidae) oviposits in all four instars of the pea aphid, Acyrthosiphon pisum(Harris) (Homoptera: Aphididae). Searching females display a highly stereotyped sequence of behaviors when encountering a host. Once recognized, an aphid is examined and probed by the wasp with the everted ovipositor prior to oviposition. Oviposition success is influenced by aphid behavior that is related to aphid size and expressed through instarspecific escape and defense reactions. Being smaller and less able to defend themselves, first and early-second instars of pea aphid are more susceptible to successful parasitism than third and fourth instars, in that order. Observed patterns of preference by Aphelinus females for particular aphid species and instars reflect the outcome of behavioral interactions between the hosts and the parasites, rather than preference in the strict sense.     

Exploitation of the Fecal Shield of the Lily Leaf Beetle, Lilioceris lilii (Coleoptera: Chrysomelidae), by the Specialist Parasitoid Lemophagus pulcher (Hymenoptera: Ichneumonidae)

We investigated signal sources used by the parasitoid Lemophagus pulcher in locating and accepting larvae of its host, the lily leaf beetle, Lilioceris lilii. Olfactometer bioassays revealed that larvae with fecal shields, larvae without shields, the shield alone, and lily leaves damaged by L. lilii were all attractive to female parasitoids. In contact bioassays, L. pulcher females were attracted to shields and showed ovipositor probing independently of whether the larva underneath was L. lilii or a nonhost, suggesting that the shield plays a primary role in short-range host location and host acceptance by L. pulcher. The attractiveness of the shield is at least partly of a chemical nature, since shield extracts applied to dummies increased contact duration and induced ovipositor probing by L. pulcher. Another putative defense system of L. lilii, i.e., oral discharge which is emitted by disturbed larvae, was also attractive to experienced, but not to naive, female parasitoids. In all other tests, naive and experienced female L. pulcher responded to the same signal sources tested, suggesting that the host-selection behavior of this biological control candidate is governed largely by innate responses to host-associated cues.     

Control of stored grain pest,Callosobruchus maculatus (F.) (Coleoptera: Bruchidae), using the essential oils isolated from Zingiber officinale (L.) and Mentha pulegium (L.) in laboratory condition

Fumigant activity of essential oil vapours distilled from Zingiber officinale (L.) and Mentha pulegium (L.) was tested against eggs, larvae and adults of Callosobruchus maculatus (F.). Fumigant toxicity was assessed at 27?±?1?°C and 60?±?5% RH, in dark condition. The influence of different concentrations of the essential oil vapours on egg hatchability, larval and adult mortality was significant. Data probit analysis showed that lethal concentration of the essential oil to kill 50% of the population (LC₅₀) for egg, larvae and adult was found to be 1.151, 2.336 and 2.183?μl/l air of Z. officinale, followed by 0.072, 0.113 and 0.093?μl/l air essential oil of M. pulegium, respectively. Between these essential oils, Z. officinale is almost more toxic than M. pulegium on all growth stages of C. maculatus. The present study suggests that essential oils from these medicinal plants may be potential grain protectants as botanical alternative fumigants and could be used in the management of various life stages of C. maculatus.     

Flight Toward a Learned Odor and Factors Inducing Landing of Female Brachymeria intermedia (Hymenoptera: Chalcididae), a Parasitoid of the Gypsy Moth, Lymantria dispar (Lepidoptera: Lymantriidae)

Through a single oviposition experience in the presence of vanilla odor, female Brachymeria intermedia were conditioned to fly toward a source of vanilla odor in a wind tunnel. Conditioning was an associative process, and wasps were able to fly toward odor sources at much lower doses than the dose used in conditioning. Furthermore, upon approaching vanilla-scented paper rolls hung on a vertical cylinder, conditioned wasps landed on them readily, whereas few wasps landed on real pupae. However, more wasps reached pupae or pupal cases than white paper rolls. Visual and olfactory cues appeared to mediate the foraging behavior of wasps in conflicting ways. Therefore, the extent to which olfactory learning mediates foraging success in this species remains unresolved.     

Oviposition behavior of an ant-parasitizing fly,Neodohrniphora curvinervis (Diptera: Phoridae), and defense behavior by its leaf-cutting ant hostAtta cephalotes (Hymenoptera: Formicidae)

This study examines the oviposition behavior of the phorid parasitoid Neodohrniphora curvinervisand the antiparasitoid defense behavior of its leafcutting ant host Atta cephalotes. N. curvinervisfemales are diurnal sit- and- wait parasitoids that attack only outbound foragers of head width 1.6 mm or greater. Females deposit a single egg through the foramen magnum of each host successfully parasitized. Pursuit of hosts is usually initiated when an outbound forager of acceptable size passes by a parasitoid perch site. Individual foragers defend themselves against pursuing parasitoids by outrunning them along the foraging trail or by standing their ground and fending them off with their legs,antennae, and mandibles. At the colony level, susceptible foragers are protected against parasitism by a shift in the forager size distribution toward smaller unsusceptible sizes during the day when parasitoids are active and toward larger sizes at night when parasitoids are inactive. The frequency of parasitism of susceptible foragers was 15%, which is more than five times the frequency found in another system involving the phorid parasitoid Apocephalus attophilusand the leafcutting ant host Atta colombica.We offer several possible explanations for such differences in the frequency of parasitism and also examine reasons for the high incidence of superparasitism (19%) observed in the system studied.     

Responses of Two Potential Host Species (Formica gnava and Formica occulta) to Pupae of the Obligatory Slave-Making Ant, Polyergus breviceps (Hymenoptera: Formicidae)

Queens of the slave-making ant, Polyergus breviceps, take over nests of adult Formica workers when establishing new colonies. Although naïve to slave-maker brood, the usurped Formica rear Polyergus offspring and nests containing both host and parasite species forms. Host worker acceptance of parasite brood has been attributed to the similarity of brood tending signals between these closely related taxa and/or the presence of an attractive pheromone in the slave-maker brood. By presenting single-species groups of Formica occulta and Formica gnava (two host species of P. breviceps) with a choice of Formica pupae of both species or with a choice of P. breviceps pupae from both types of mixed-species nests, it seems that neither close phylogenetic relatedness nor an attractive brood odor alone can account for the propensity of host workers to adopt slave-maker pupae. Significantly greater numbers of P. breviceps pupae were adopted by enslaved workers than by free-living workers, and within the enslaved groups and the free-living F. gnava group, greater numbers of P. breviceps pupae were adopted if they were from nests where the host species was conspecific to workers used in tests. When presented with F. gnava and F. occulta pupae, Formica workers adopted conspecific pupae almost exclusively and ignored or consumed pupae of the other host species. Taken together, these results imply that P. breviceps pupae have both a species-specific odor and a general brood-tending pheromone, upon which a host odor may be imposed. The disparate requirements of immatures at different stages of development for cue specificity or generality in maintaining nest exclusivity and maximizing inclusive fitness are discussed.     

Effects of postemergence experience on searching and landing responses of the insect parasitoid,Cotesia congregata (Say) (Hymenoptera: Braconidae), to plants

Postemergence experience with one of six plant species, in the presence of the host larva, modified the searching response of reproductively mature females of Cotesia congregata(Say) to these plants in at least one of three ways: (1) an increased response to the plant experienced at emergence, (2) an increased response to other plants, or (3) an inhibited response to other plants. Landing and searching responses were differentially affected by postemergence experience. For example, postemergence experience with tobacco (a common plant) in the presence of the host larva induced a landing preference for this plant over parsley (a novel plant) but did not affect searching responses to either plant, whereas experience with parsley and the host larva induced an increased searching response to parsley but a landing preference for tobacco. Differential effects of postemergence experience may reflect the type of stimuli involved in searching or landing and may have adaptive significance.     

Searching and Oviposition Behavior of Anagrus atomus L. (Hymenoptera: Mymaridae) on Four Host Plants of Its Host, the Green Leafhopper Empoasca decipiens Paoli (Homoptera: Cicadellidae)

Anagrus atomus L. is an important egg parasitoid of the green leafhopper Empoasca decipiens Paoli. In this study the ability of the parasitoid to locate and parasitize its host was investigated on four host plants, i.e., broad beans (Vicia faba L.), sweet pepper (Capsicum annuum L.), cucumber (Cucumis sativus L.), and French beans (Phaseolus vulgaris L.). For each plant species, the behavior of the parasitoid was observed on E. decipiens infested and noninfested plants. Searching and oviposition behavior were characterized by drumming, probing, and resting. Parasitoids spent significantly less time on non-infested than infested plants, 274.5 and 875.7 s, respectively, and no probing behavior was observed on non-infested plants. Frequency of resting behavior was significantly greater on non-infested than on infested plants. Total foraging time was significantly longer on infested than on non-infested plants, indicating that A. atomus females can efficiently discriminate between leaves with and without infestation. Parasitism of A. atomus was influenced by parasitoid density, with the highest parasitism rate (64.0%) obtained at a density of 10 A. atomus females/0.1356 m² but the number of parasitized eggs per female and the searching efficiency decreased with increasing parasitoid density.