Effects of within- and among-patch experiences on the patch-leaving decision rules in an insect parasitoid

The present study aimed to address how an insect parasitoid makes patch-departure decisions from various types of host patches and how previous patch experiences in the environment modify this decision-making process. Experiments were done with the parasitic wasp Aphidius rhopalosiphi attacking the grain aphid Sitobion avenae. In the experiments, wasps were observed in a laboratory environment containing several patches of various host densities, and behavioural records were analysed using a Coxs proportional hazards model. Consideration of the effect of the within-patch experience gave a classic pattern of patch-leaving decision rules in parasitoids: A. rhopalosiphi used local information on host quality (i.e. numbers of ovipositions or rejections) and availability (i.e. patch density) to determine departure decision. However, consideration of previous patch experiences provided evidence that these departure rules are fundamentally dynamic, responding to the physiological state of the animal and the information it has about its environment. Results showed that A. rhopalosiphi decreased its tendency to leave the visited patch after an oviposition. However, when a female has already laid several other eggs in the environment, such an incremental mechanism gradually switched to a decremental one. Hence, A. rhopalosiphi responded to egg-load depletion by leaving the visited patches sooner and by depositing a smaller number of eggs in those patches, which probably led to a decreased level of superparasitism. Results also indicated that previous experiences enabled wasps to estimate spatial host distribution and then to adjust their behavioural decisions accordingly. Thus, A. rhopalosiphi was shown to adjust its patch residence time according to the quality and the number of the patches previously visited. These proximate mechanistic rules adopted by A. rhopalosiphi females are discussed in the context of general predictions from optimality models.Communicated by D. Gwynne     

Unattractiveness of mated females to males in the parasitoid wasp Spalangia endius

Despite common stereotypes, males are not always indiscriminate and eager when it comes to mating. In the parasitoid wasp Spalangia endius, the initial response of males to females was almost always one of apparent excitement; however, this was followed by a clear preference for virgin females over mated females in both no-choice and choice situations. The no-choice data were collected from videotapes of male-female pairs of all possible combinations of mated and virgin individuals. Neither female nor male mating status had a significant effect on likelihood of, or time until, contact or male courtship fanning. However, a males first retreat was sooner when the female was mated than when she was virgin; mated males exhibited their first retreat sooner than did virgin males; and mated females were less likely to be mounted than were virgin females. In addition to the videotapes, male-choice experiments were performed. When given a choice of a virgin and a mated female, both virgin and mated males were more likely to mount and copulate with the virgin. The difference in response to virgin versus mated females seemed to be less in virgin males than in mated males, perhaps due to virgin males greater eagerness to mate: when a virgin male and a mated male were presented with a dead virgin female, the virgin male was usually the first to respond to the female. That males preferentially retreated from and avoided mounting mated females appears to be adaptive given that mated females rarely copulated.Communicated by N. Wedell     

Partial compensation of the sublethal effect of deltamethrin on the sex pheromonal communication of Trichogramma brassicae

Pyrethroid insecticides are widely used and lead to a sizable environmental pollution that could interfere with the population biology of insects. Trichogramma is a beneficial insect used in biological control and which natural populations contribute to the control of Lepidopterus pests. In this work, we determined the effect of a sublethal dose of deltamethrin on the sex pheromonal communication of Trichogramma. The dose used (LD 0.1) induces no detectable mortality (the theoretical mortality is only one insect over 1000) and can be a good representation of contamination by this insecticide from environmental pollution. The insecticide was shown to have opposite effects on the sex pheromonal communication of Trichogramma, depending on which sex was exposed (Delpuech, J.M., Legallet, B., Terrier, O., Fouillet, P., 1999. Chemosphere 38, 729–739). We show that, when both sexes are simultaneously exposed to the insecticide, this effect is only partially neutralized. The mean response of treated males responding to the sex pheromone from treated females is not significantly different from that of controls, but the kinetics of their response is not the same. When both sexes are treated, the response of males to the sex pheromone is lower at the beginning but their response does not decrease during time contrary to controls and becomes finally higher than that of controls. Therefore, the sublethal effect of deltamethrin in the field can be either advantageous or disadvantageous depending on the difficulty in finding females and their scarcity.     

Latency to resume calling after disturbance in the field cricket,<Emphasis Type="Italic"> Teleogryllus oceanicus</Emphasis>, corresponds to population-level differences in parasitism risk

A possible parasitoid-evasion behavioral adaptation is examined in male field crickets, Teleogryllus oceanicus, from three Hawaiian islands where parasitoid prevalence varies naturally among islands. Ormia ochracea, the parasitoid fly that parasitizes T. oceanicus on these islands, uses male calling song to locate its hosts. We used laboratory-reared males from three Hawaiian islands to determine if there are population differences in the time it takes for calling males to resume calling after a standardized disturbance. Males follow the expected pattern; males from the island with the greatest risk of parasitism have the longest latency to resume calling, and males from the island with the least risk of parasitism have the shortest latency to resume calling. Results are discussed in the context of behavioral adaptations to differing parasitism levels, and trade-offs between natural and sexual selection.Communicated by D. Gwynne     

Pre-pupation behaviour of the aphid parasitoid <Emphasis Type="Italic">Aphidius ervi</Emphasis> (Haliday) and its consequences for pre-imaginal learning

Olfactory learning may occur at different stages of insect ontogeny. In parasitoid wasps, it has been mostly shown at adult emergence, whilst it remains controversial at pre-imaginal stages. We followed larval growth of the parasitoid wasp, Aphidius ervi Haliday, inside the host aphid, Acyrthosiphom pisum Harris, and characterised in detail the behaviour of third instar larvae. We found that just before cocoon spinning begins, the third instar larva bites a hole through the ventral side of the mummified aphid exoskeleton. We then evaluated whether this period of exposure to the external environment represented a sensitive stage for olfactory learning. In our first experiment, the third instar larvae were allowed to spin their cocoon on the host plant (Vicia faba L.) surface or on a plastic plate covering the portion of the host plant exposed to the ventral opening. Recently emerged adults of the first group showed a preference for plant volatiles in a glass Y-olfactometer, whereas no preference was found in adults of the second group. In a second experiment, during the period in which the aphid carcass remains open or is being sealed by cocoon spinning, third instar larvae were exposed for 24 h to either vanilla odours or water vapours as control. In this experiment, half of the parasitoid larvae were later excised from the mummy to avoid further exposure to vanilla. Adult parasitoids exposed to vanilla during the larval ventral opening of the mummy showed a significant preference for vanilla odours in the olfactometer, regardless of excision from the mummy. The larval behaviour described and the results of the manipulations performed are discussed as evidences for the acquisition of olfactory memory during the larval stage and its persistence through metamorphosis.     

Ambient ultraviolet radiation induces protective responses in soybean but does not attenuate indirect defense

We investigated the effects of ambient ultraviolet (UV) radiation on (i) the performance and chemistry of soybean plants, (ii) the performance of Spodoptera frugiperda and (iii) the foraging behavior of the herbivore's natural enemy Cotesia marginiventris which exploits herbivore-induced plant volatiles (VOC) for host location. The accumulation of protective phenolics was faster in plants receiving ambient UV than in controls exposed to sun light lacking UV. Accordingly, isorhamnetin- and quercetin-based flavonoids were increased in UV exposed plants. No UV effects were found on the performance and feeding behavior of S. frugiperda. Herbivore-damaged plants emitted the same VOC when grown under ambient or attenuated UV for 5, 10 or 30 days. Consequently, C. marginiventris was attracted but did not discriminate between exposed and unexposed soybeans. In summary, ambient UV radiation affected soybean morphology and physiology but did not destabilize interactions between trophic levels.     

Sex ratio and oviposition responses to host age and the fitness consequences to mother and offspring in the parasitoid wasp Spalangia endius

In the parasitoid wasp Spalangia endius more offspring and a greater proportion of daughters were oviposited in, and emerged from 0-day-old versus 3-day-old hosts. Offspring that developed on the younger hosts (1) were larger at adulthood, (2) developed more quickly, (3) had higher survivorship to adulthood, and (4) were more often able to chew their way out of the host. Sons and daughters did not differ in how host age affected their size, development rate, or survivorship. The greater proportion of daughters from the younger hosts may be adaptive, as described by the host quality model (a variant of the Trivers and Willard hypothesis). It is adaptive if greater size or more rapid development has a more positive effect on a daughter’s than a son’s fitness and the positive effect is large enough to compensate for sons being trapped disproportionately to daughters in the older hosts. Despite greater success at drilling the younger hosts, mothers did not try to drill them sooner or more often. Having previously oviposited on the older rather than the younger hosts had no detrimental effect on the mother’s subsequent longevity or offspring production. Received: 8 March 2000 / Revised: 9 June 2000 / Accepted: 24 June 2000     

The larval parasitoid <Emphasis Type="Italic">Microplitis croceipes</Emphasis> oviposits in conspecific adults

Microplitis croceipes (Hymenoptera: Braconidae) is a larval parasitoid of Helicoverpa/Heliothis spp. In the course of mass rearing of M. croceipes, we found that females oviposited in the conspecific adults in rearing cages. When 20 pairs of inexperienced females and males or of experienced females and males were reared in a cage, the males lived for 14–15 days and the females for 18–20 days on average. At their death, 37–42% of the males and 50–57% of the females contained conspecific eggs or first instar larvae in their abdominal cavity. When two of inexperienced females met on a host-infested leaf of soybean, they attempted to sting each other. Of the attacked females, 30% contained a conspecific egg laid in their abdomen. In abdominal cavity of the adults parasitized by a conspecific female, the majority of the parasitoid eggs laid disappeared within 1 day after oviposition. Only 10–30% of the parasitoid eggs laid in conspecific adults hatched 3–4 days after oviposition, but those larvae never molted to second instar. When the adults were stung by one or two conspecific females, their subsequent longevity was significantly shorter than that for the control adults. Oviposition in conspecific adults may be prevalent in other parasitic wasps that quickly oviposit without intensive host examination, and have cuticle and size of abdomen to be stung by conspeicifcs.     

Learning affects host discrimination behavior in a parasitoid wasp

Learning is generally predicted not to be important in host discrimination by parasitoids, because the stimuli involved are less variable than those used in habitat location. However, Anaphes victus (Hymenoptera: Mymaridae), an egg parasitoid of Listronotus oregonensis (Coleoptera: Curculionidae) apparently learns to associate external pheromones with the presence of a conspecific in a host. In this species, females can reject a parasitized host either after antennal drumming (antennal rejection) or after the insertion of their ovipositor (sting rejection). When they encountered a series of parasitized hosts, females A. victus learned to associate the presence of the external pheromone with the presence of the internal one. Learning lasted less than 4 h and occurred earlier in a series when the female marking the egg and the one detecting that mark were close relatives. This behavior could be adaptive because antennal rejection is faster than sting rejection. Received: 11 March 1997 / Accepted after revision: 30 August 1997     

Test of the adaptiveness of sex ratio manipulation in a parasitoid wasp

In behavioral ecology it is generally assumed that behavior is adaptive. This assumption is tested here for sex ratio manipulation in response to host size in the parasitoid wasp Spalangia cameroni. Females produce a greater proportion of daughters on larger hosts. If this behavior is adaptive, it is not through a positive effect of host size on the fitness of daughters, as theory suggests and as found for other species. Females that developed on larger hosts were not more successful at drilling into hosts, were not more successful at interspecific competition for hosts, and did not have greater dispersal ability as measured by wing loading (weight/area of wing and thorax). The possibility that S. cameroni's sex ratio manipulation may be adaptive through a negative effect of host size on the fitness of sons cannot be ruled out. Relative to males from larger hosts, males from smaller hosts had lower wing loading and thus potentially greater dispersal ability. The actual effect of wing loading on fitness remains to be tested.