In many insect species with a pupa covered by various "shells" (puparium, host remains, etc.) pupal-adult ecdysis and emergence to the open air represent two discrete steps. However, in Trichogramma, as well as in other insect parasitoids, these two processes have never been studied separately. We investigated the temporal pattern of pupal-adult ecdysis and of adult emergence from the host chorion in Trichogramma embryophagum Hartig (Hymenoptera: Trichogrammatidae) in laboratory conditions (12L : 12D, 20°C). Adult ecdysis was arrhythmic, while adult emergence showed a strong rhythmicity. The time lag between ecdysis and emergence varied from one to almost two days, depending on the circadian time of the ecdysis. The proportion of ecdysed adults that stayed in the host chorion ranged up to 60% (just before the highest peak of emergence). The cumulative percentage of ecdysed adults gradually increased with time, independently of whether the light was turned on in accordance with the entrained circadian rhythm or 4 h earlier. This arrhythmic ecdysis could be explained by the fact that the ecdysed adults get into a well protected space inside the host chorion and the timing of this event is adaptively neutral.
A comparative study of the direct influence of single, non-repetitive changes of light and/or temperature on the daily pattern of adult eclosion was conducted in laboratory conditions with the egg parasitoid, Trichogramma embryophagum (Hartig). In all experiments, the entire development occurred under the same regimen (12L : 12D, 20°C). However, just in the day when records of eclosion were made, the time of light-on was changed (from the total absence of a dark period to the 12-h-long scotophase). In addition, in different treatments of each experiment a thermophase (30°C during 2 or 4 hours) was applied in various positions relative to light-on. Results showed that light-on or a temperature step-up could induce eclosion in a certain percentage of individuals practically at any time of the 24-h-long cycle (masking effects). The fraction of adults emerging during 2 h after the beginning of the action of an exogenous factor depended not only on the preceding scotophase duration, but also on the stimulus itself (light, temperature or both). The temperature stimulus was found to modify the responsiveness to light considerably, depending on the order of application of the stimuli. The simultaneous action of light-on and a temperature step-up exerted a more considerable influence than could be expected based on their separate impacts. On the contrary, a high temperature pulse preceding the onset of the photophase inhibited the response of individuals to light-on, evidently acting by phase delay of a circadian rhythm of light responsiveness. These effects were more pronounced in the first half of the 12-h-long scotophase. The data suggest a complex interaction between the exogenous factors in their influence on the daily pattern of adult eclosion.
A laboratory study was carried out on photoperiodic control of prepupal diapause in the egg parasitoid Trichogramma embryophagum (Hartig). All experiments were conducted with an isofemale parthenogenetic strain. The maternal generation was reared at 20°C and photoperiods of L:D = 3:21, 6:18, 9:15, 12:12, 15:9, 18:6, 21:3 or 24:0. The tendency to diapause in the progeny was estimated by rearing the daughter generation at 15°C in the dark. Experiments revealed a long-day type response based on maternal influence on the progeny prepupal diapause. However, significant endogenous fluctuations in the pattern of the photoperiodic curve were revealed in successive laboratory generations reared under constant conditions. The left threshold day-length was very variable, while the right threshold kept relative constancy. Experiments with individual females sequentially offered new host eggs demonstrated that the probability of the progeny entering diapause depends significantly on maternal age. At 20°C and 18L : 6D, the percentage of diapause was maximal (ca 15%) in the progeny eclosed from the eggs laid during 1st - 2nd days of maternal life. Then the proportion of diapausing progeny decreased to 0-5% at days 9-11 of female life and later slightly increased in 15-17 days old females. Thus, endogenous factors play an important role in maternal influence on progeny diapause, particularly in environments close to threshold temperature and photoperiod.
Despite the impact of parasitoids on insect populations being extensively studied, indirect parasitoid-mediated effects remain rarely documented in natural communities. We examined the influence of shared parasitoids on the interactions between two functionally monophagous moths, Nonagria typhae and Archanara sparganii. The moths showed a considerable variation in terms of relative abundance and the degree of phenological synchrony between the species. On average, parasitism levels caused by shared parasitoids did not differ between the two host species. Relative parasitism levels of the two hosts, however, varied considerably among different samples. Percentage parasitism of the scarcer species, A. sparganii, thus could not be fully explained by that of the dominant species, N. typhae. The results indicated that A. sparganii may benefit from the presence of N. typhae. In particular, both low relative density as well as high phenological synchrony with N. typhae reduced parasitism levels in A. sparganii. The case thus indicates the presence of parasitoid-mediated indirect effects between the coexisting herbivores. The patterns of host use observed in this study are consistent with the scenario of frequency-dependent host use caused by changes in parasitoid behavior. Such a host use by parasitoids is suggested to promote numerical stability and coexistence of the moth species in the system studied.