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.
1_Maternal and grand-maternal photoperiodic responses of Trichogramma buesi, T. embryophagum, T. evanescens, T. piceum, T. principium, and T. telengai were investigated in laboratory conditions. During the experiment, grand-maternal and maternal generations developed at 20°C and one of the 4 photoperiodic regimes: L : D = 12 : 12, 14 : 10, 16 : 8, and 18 : 6 (in total, 16 combinations) while the progeny developed at L : D = 12 : 12 and one of the 3 thermal regimes: 13, 14, and 15°C. The proportion of diapausing individuals in the progeny of all the studied species was significantly dependent on the direct influence of temperature and on the maternal photoperiodic response. The influence of the photoperiodic conditions during development of the grand-maternal generation was statistically significant in 5 of the 6 studied species, being relatively weak in T. embryophagum and T. telengai, whose geographical ranges extend up to north-western regions of Europe (possibly, these wasps enter diapause so early that the grand-mothers of the diapausing generation develop under long day conditions). Comparative analysis showed that the thresholds of the maternal and grand-maternal photoperiodic responses coincided or almost coincided. The grand-maternal effect was stronger in the progeny of maternal females which developed under short day conditions than in those that developed under long day conditions. This pattern of interaction probably synchronizes the life cycle with seasonal changes because diapause is induced under decreasing day length and thus mothers of diapausing individuals develop at shorter daylength than do grand-mothers., 2_We conclude that the grand-maternal and the maternal effects on Trichogramma progeny diapause are based on one and the same photoperiodic response. In nature, the grand-maternal effect increases the proportion of diapausing individuals in the progeny of females which have developed under short day conditions during two generations, thus achieving a "cumulative" photoperiodic effect., Natalia D. Voinovich, Nina P. Vaghina, Sergey Ya. Reznik., and Obsahuje seznam literatury
The effect of photoperiod on parasitization of the eggs of the Angoumois grain moth, Sitotroga cerealella (Olivier, 1789) by Trichogramma principium Sugonyaev & Sorokina, 1976 was investigated under several photoperiodic regimes of L : D = 3 : 21, 6 : 18, 9 : 15, 12 : 12, 15 : 9, 18 : 6 and 21 : 3. In all regimes, certain wasps delayed ovipositing in this non-preferred host. Potential fecundity of T. principium females (the number of mature ovarial eggs at emergence) and subsequent oogenesis (estimated by the number of mature ovarial eggs in non-ovipositing females) was independent of photoperiod. However, the percentage of females that oviposited was higher for females that developed and were kept under 6-12 h long photophase than for those that developed and were kept under ultra short (3L : 21D) and under long (18L : 6D and 21L : 3D) photophases. The average duration of the pre-oviposition (egg retention) period showed the opposite pattern to the photoperiodic response. A possible explanation of this reaction is that the delay in oviposition is adaptive if the probability of finding a better host is high. In autumn, when the last Trichogramma females are still active but their lepidopteran hosts are already much less abundant, then parasitization of any suitable host is the best strategy.
At emergence females of Trichogramma had a lot of mature eggs in their ovaries, but some delayed parasitization or refused to parasitize a laboratory host. The effect of constant and alternating temperatures on the percentage of Trichogramma buesi females parasitizing Sitotroga cerealella eggs and the duration of the pre-parasitization period were investigated. The temperature dependencies of the rate of preimaginal development, pre-emergence survival, number of eggs laid daily, and total lifetime fecundity were also determined. As the temperature was increased from 12 to 35°C, the median pre-oviposition period decreased from 5 days to 3 h, with maximum values of 24 and 1.5 days, respectively. The rate of induction of parasitization (reciprocal of duration of the pre-parasitization period of the females that parasitized) increased with temperature like the rate of preimaginal development and average number of eggs laid daily by a parasitizing female. Total cumulative percentage of parasitizing females reached a maximum (ca 60%) at temperatures of 25-30°C, while at 12 and 35°C, respectively, 25 and 50% of females parasitized the S. cerealella eggs. Average lifetime fecundity and pre-emergence survival showed a similar dependence on temperature. The influence of the thermorhythm (25°C for 4 h and 15°C for 20 h) was strongly dependent on its position within the photoperiod. When thermophase coincided with photophase, the percentage of females that parasitized was close to that recorded at a constant temperature of 25°C. But when the high temperature pulse coincided with the dark period, the percentage of parasitizing females was the same as at 15°C. Thus, the temperature dependence of ethogenesis (supposedly, an increase in motivation to parasitize or search for a host) in Trichogramma females was similar to that of morphogenesis, although the reaction to alternating temperatures may have been complicated by interaction with the light : dark regime.
This is the first record of the harlequin ladybird Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae) occurring in the Caucasus (Sochi region of Krasnodar territory, Russia). All the adults collected in the field there and reared from collected eggs, larvae and pupae, and their progeny were H. axyridis f. succinea, which is the most common morph in natural populations in South-Eastern Asia and the Russian Far East as well as in invasive populations in the Americas and Europe. In contrast in Western Siberia f. axyridis predominates and this indicates that an occasional introduction from the closest native range in Siberia cannot be considered as the source of the Caucasian population. It is known that populations of H. axyridis can also differ in their photoperiodic responses. The results of earlier experiments on H. axyridis, which originated from the Russian Far East, indicate that the threshold day lengths for the acceleration of preimaginal development and deceleration of reproductive maturation were 13–14 h, while for the invasive populations of this species in Europe these two thresholds are approximately 12 h. In the population studied, the thresholds for both of these photoperiodic responses were also approximately 12 h. Thus, it is concluded that the population of H. axyridis studied is a further eastward continuation of the invasion of this species in Europe., Natalia A. Balyakova, Sergey Ya. Reznik., and Seznam literatury
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.
The effects of photoperiod on pre-imaginal development and reproductive maturation of adult females of the multicolored Asian lady beetle, Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae), were investigated at 20°C and at photoperiods from 10L : 14D to 18L : 6D. Experiments were conducted on a laboratory strain that originated from the Russian Far East. Larvae and adults were fed on the green peach aphid Myzus persicae. Under short day conditions the pre-imaginal development was faster than under long day conditions. The acceleration of pre-imaginal development occurred when day length was shorter than 16 h and the threshold day length was ca 14 h. The rate of reproductive maturation of adult females, in contrast, was twice as high under long day conditions, with the threshold between 14L : 10D and 12L : 12D, although even under short day conditions ca 10% of the females show a tendency to mature more quickly. This difference between the thresholds of these two photoperiodic reactions indicates they are relatively independent of one another. and Sergey Ya. Reznik, Nina P. Vaghina.
Giant hogweed, Heracleum mantegazzianum (Apiaceae), was introduced from the Caucasus into Western Europe more than 150 years ago and later became an invasive weed which created major problems for European authorities. Phytophagous insects were collected in the native range of the giant hogweed (Caucasus) and were compared to those found on plants in the invaded parts of Europe. The list of herbivores was compiled from surveys of 27 localities in nine countries during two seasons. In addition, literature records for herbivores were analysed for a total of 16 Heracleum species. We recorded a total of 265 herbivorous insects on Heracleum species and we analysed them to describe the herbivore assemblages, locate vacant niches, and identify the most host-specific herbivores on H. mantegazzianum. When combining our investigations with similar studies of herbivores on other invasive weeds, all studies show a higher proportion of specialist herbivores in the native habitats compared to the invaded areas, supporting the "enemy release hypothesis" (ERH). When analysing the relative size of the niches (measured as plant organ biomass), we found less herbivore species per biomass on the stem and roots, and more on the leaves (Fig. 5). Most herbivores were polyphagous generalists, some were found to be oligophagous (feeding within the same family of host plants) and a few had only Heracleum species as host plants (monophagous). None were known to feed exclusively on H. mantegazzianum. The oligophagous herbivores were restricted to a few taxonomic groups, especially within the Hemiptera, and were particularly abundant on this weed.