Site-specific mate-finding strategies and oviposition behavior inCrocothemis erythraea (Brullé) (Odonata: Libellulidae)

Mate-finding strategies and adaptations in pre- and postcopulatory behaviors to avoid male disturbance were studied in Crocothemis erythraea at two different sites. At ponds without perch sites males patrolled with temporal partitioning of the limited oviposition sites and male-male disputes were rare. The arrival rate of females was high. At temporary marshes with dense emergent vegetation the oviposition sites were widely distributed. Males mainly perched and interacted with longer disputes. At both types of habitats with high male densities females showed a similar number of copulations per visit and oviposition durations. Postcopulatory behavior to avoid male disturbance and to decrease remating of the female differed in both sexes. At the ponds with patrolling males the probability of remating in guarded and unguarded ovipositions was similar and higher than at the marshes. At the marshes 78% of rematings occurred when the guarding male was still involved in disputes with the previously disturbing male. At the ponds females hovered before escaping successfully from approaching males and they changed to another oviposition site where they continued oviposition. Females at the marshes remated after surprise attacks by neighboring males.     

Oviposition and mining by Phytobia betulae (Diptera: Agromyzidae) in genotypes of European white birch (Betula pendula)

Abstract 1 The colonization success of herbivorous insects depends partly on the ability of females to choose suitable host plants. Phytobia betulae Kangas (Diptera: Agromyzidae), a miner of differentiating xylem, infests birch (Betula pendula Roth and Betula pubescens Ehrh.) trees. 2 In a field experiment we measured successful ovipositions in the long shoots of eight 5‐year‐old‐birch genotypes. Later in the growing season, in the same experiment, we counted the number of larvae from the same birch genotypes (different replicates). 3 We attempted to determine where Phytobia females lay their eggs in the birch canopy, what kind of shoots Phytobia utilize, whether birch genotypes differ in their susceptibility to Phytobia, and which tree characteristics correlate with successful utilization of birch for oviposition and larval development. 4 The ovipositions were concentrated in fast growing shoots and only 5% of the available shoots were utilized. Usually a shoot contained only one oviposition (80% of the cases). However, oviposition pattern resulted in significant aggregations (two to four ovipositions per shoot): 11% of shoots selected for oviposition contained more than one Phytobia and 22% of larvae shared their shoot with at least one other larva. Although birch shoots were much more abundant than Phytobia, there may be intraspecific competition for host resources among Phytobia. 5 Birch genotypes differed in the number of ovipositions but not in the number of larvae, although in general the number of larvae was correlated positively with the number of ovipositions. There were more ovipositions in the faster growing genotypes.     

Comparison of oviposition behavior of a solitary and a gregarious parasitoid (Hymenoptera: Mymaridae)

Anaphes victus Huber andAnaphes listronoti Huber (Hymenoptera: Mymaridae) are respectively solitary and gregarious egg parasitoids of the carrot weevil,Listronotus oregonensis (LeConte) (Coleoptera: Curculionidae). We made detailed ethograms of the oviposition behavior on unparasitized and parasitized hosts for the two species. We then compared the behavior of virgin and mated females for the oviposition of male and female progenies. The two species did not always oviposit after insertion of the ovipositor, but these punctures without oviposition could be readily differentiated from oviposition.A. victus oviposited only once by puncture, whileA. listronoti deposited one to three eggs during the same sequence. The variability of the duration of the various components was generally lower for a given female than between females. Two components, the abdominal vibrations and the pause, were significantly shorter in ovipositions that resulted in male progency for the two species. However, an important overlap in duration prevents using these differences to sex the progeny at oviposition. Virgin females of both species, although capable of producing only males, exhibited both behaviors. Parasitized hosts were recognized through internal and external markings that were used in host discrimination.     

The developmental viability and sizes of sequentially oviposited eggs ofBoophilus decoloratus andBoophilus geigyi

The viability and sizes of eggs sequentially laid up to the 10th day of oviposition by engorgedBoophilus decoloratus andBoophilus geigyi were studied. The average length and breadth of eggs laid on the 1st and 2nd days of oviposition were statistically greater than at subsequent ovipositions. The eclosion periods of eggs laid from 7th to 10th day of oviposition were shorter than those of eggs laid earlier by bothBoophilus species. The percentage mortality of eggs of 9th and 10th day of oviposition was significantly higher than that of the earlier ovipositions. The numbers of larvae which engorged on rabbits after hatching from eggs of 7th–10th day ovipositions were significantly fewer than those which hatched from eggs of earlier ovipositions. It is concluded that eggs of early ovipositions ofB. decoloratus andB. geigyi are more viable and survive longer than those of later ovipositions. It is also suggested that the shorter eclosion period of the eggs of later ovipositions is due to the process of embryogenesis which is gradual in the eggs of early ovipositions and faster in those of later ovipositions.     

Reproductive biology and function of multiple mating in the mating system of a tree cricket,Truljalia hibinonis (Orthoptera: Podoscritinae)

A tree cricket,Truljalia hibinonis, is known to show a novel sperm removal during copulation. The pattern of copulations and ovipositions showed that the sperm removal functioned to increase reproductive success for sperm removing males. The sperm removal by males evolves under the system in which female accept multiple mating. The possible benefits of multiple mating for females are examined. Multiple mating did not seem to be necessary for avoiding sperm depletion, because females stored huge number of sperm in their sperm storage organ after finishing oviposition. The ingestion of metanotal secretion during copulation also had no effect on increasing fecundity and egg size. However, mating experience may have a positive effect on increasing fecundity slightly, though there were no differences between once- and twice-mated females. The other possible benefits for each male and female are discussed.     

Host Stage Selection and Sex Allocation by Gyranusoidea tebygi (Hymenoptera: Encyrtidae), a Parasitoid of the Mango Mealybug, Rastrococcus invadens (Homoptera: Pseudococcidae)

Host stage selection and sex allocation by Gyranusoidea tebygi Noyes (Hym,, Encyrtidae) were studied in choice and no-choice experiments in the laboratory. The parasitoid reproduced on first, second, and third instars of the mango mealybug, Rastrococcus invadens Williams (Hem., Pseudococcidae), and it avoided hosts that were already parasitized. Host feeding was occasionally observed. Sex ratios of the offspring produced by individual wasps were highly biased in favor of females, whereas the sex ratio of groups of wasps foraging under crowded conditions varied from male biased in smaller hosts to female biased in larger hosts. Females had longer developmental times than males, developed faster in larger mealybugs than in smaller ones, and were always larger than males emerging from the same host instar. Their size increased with the instar of the host at oviposition. About 90% of all ovipositions in second and third instar nymphs resulted from an attack with multiple stings, starting with a sting in the head of the host for the most part. The function of these head stings is either to assess quality of the host or to subdue hosts prior to oviposition. Encounter rates, number of attacks, and number of stings during one attack increased, while ovipositions decreased with host instar. Time investment per oviposition and time spent preening increased with increasing host age because older hosts defended themselves more vigorously than younger ones. Thus, while fitness of the parasitoid increased with host size, fitness returns per time decreased. The implications of this host selection behavior for the biological control of the mango mealybug are discussed.     

Host size,superparasitism and sex ratio in mass-reared <Emphasis Type="Italic">Diachasmimorpha longicaudata</Emphasis>, a fruit fly parasitoid

We analyzed the relationship among host size, superparasitism and sex-ratio in mass reared Diachasmimorpha longicaudata (Hymenoptera: Braconidae). Individual host pupae of Anastrepha ludens (Diptera: Tephritidae) were measured (length and width), and the number of oviposition scars per pupa was used as a reliable indicator of superparasitism. The probability of an emerging parasitoid being a female was positively associated with the number of oviposition scars on the host cuticle, but not with the host size. The number of scars per host pupae from which females emerged was slightly but significantly higher than in those pupae giving raise to males. In D. longicaudata, the influence of host size on sex allocation decisions of individual females seems to be overridden by the level of superparasitism, which itself was positively correlated with pupa length. This suggests that larger pupae could experience a higher number of ovipositions than their smaller counterparts, and that a high level of superparasitism may conduct to a female biased sex ratio. We discuss the relevance of these findings which could provide new elements (e.g., the manipulation of superparasitism) for optimizing the mass rearing of this parasitoid.     

The adaptive significance of mate guarding in the soapberry bug,Jadera Haematoloma (Hemiptera: Rhopalidae)

Male soapberry bugs (Jadera haematoloma)face severe mating competition at the northern edge of their range due to male-biased adult sex ratios. Copulations lasting up to 11 days may serve a mate guarding function (encompassing four or more ovipositions), but copulation duration is highly variable, with some pairings lasting as little as 10 min. Data were gathered to describe factors that influence the reproductive costs and benefits of prolonged copulation. Estimated copulation durations (mean ± SD) were 20 ± 23 h in the lab and 50 ± 8 h in the field and were only weakly affected by sex ratio. Females mated for 5 min produced as many fertile eggs as those mated for 600 min laid; they became depleted of fertile sperm after about 25 days. In twicemated females, the first male's paternity was reduced by about 60%, and all females (N = 13) whose mates were removed experimentally mated again within an average of 6 min. The outcome of sperm competition on a perclutch basis was not highly predictable. The possibility of increased sperm displacement in longer copulations was not tested. Males often guarded females during oviposition and successfully defended them from intruding single males by recopulating. Such intrusions occurred in the majority of oviposition attempts observed in nature. Even though most females mated promiscuously, in a focal aggregation with a mean sex ratio of 2.2 ± 0.4 males/female, the interval between matings by males was commonly several days. Males appeared to respond facultatively to several aspects of the distribution and availability of females. The intensities of mating competition and sperm competition indicate that monogamous mate guarding should be favored over nonguarding in nature. Unpredicted brief. pairings may result from assessment by males of female reproductive value or of their own physical condition, or from female resistance.     

Mating tactics and male wing dimorphism in the damselfly,Mnais pruinosa costalis selys (Odonata: Calopterygidae)

Males of the damselfly,Mnais pruinosa costalis, exhibit wing color dimorphism: one form has orange wings, and the other hyaline wings which resemble female wings. The former is usually territorial and the latter uses sneaky mate securing tactics. When orange-winged males failed to establish territory, they became floaters that day. Hyaline-winged males perched around their territories and often, formed in tandem without any apparent courtship behavior when they found females. Their copulation frequency was higher and copulation duration longer than those of territorial males. A few females oviposited without remating. Total oviposition duration of females with which a hyaline-winged male mated was more than 32 min per male on average in a day Females that copulated with hyaline-winged males often remated with orange-winged residents before oviposition. Total duration of oviposition bouts of females after mating with floaters was short (15 min), while that with territorial residents was long (66 min). As a result, total oviposition duration of females with which an orange-winged male mated was about 40 min in a day. The reproductive success of the hyaline-winged males may be similar to that of the orange-winged males.     

The avoidance of superparasitism in four species of parasitic wasp—Mathematical models and experimental results

Experimental results, and interpretation, are presented which relate to oviposition behaviour in four species of parasitic wasp (Hymenoptera: Chalcidoidea). All lay their eggs in pupae of the common house fly (Musca domestica) and of other flies associated with man. Probability models suggested byD. J. Daley for describing the avoidance of superparasitism are used to analyse the data. Certain of the models considered by previous authors appear as specieal cases. A satisfactory fit was obtained either by assuming a fixed probability of oviposition in an already parasitised pupa, or by using a model motivated by supposing that a female will tolerate without oviposition some fixed number of visits to already parasitised pupae. A third model was considered which makes the probability depend on the number of previous ovipositions in the pupae now visited. In all four species the presence of other females (with parasite: host ratio unchanged) increased the probability of oviposition upon encountering an already parasitised pupa. In the presence of other females Spalangia endius laid an increased number of eggs, whereas the other three species laid fewer eggs.     

Nymphal cannibalism in relation to oviposition behavior of adults in the assassin bug,Agriosphodrus dohrni signoret

Summary Nymphal cannibalism was studied over a five year period in a natural population of the assassin bug,Agriosphodrus dohrni Signoret. Cannibalism among instar I nymphs was potentially an effective agent to cause rapid decline of the newborn population in contrast to the rarity of cannibalism in other instars. The high incidence of cannibalism among newborn was associated with concentrated oviposition on the same site by two or more females. The rate of the duplicate oviposition was high at hollows on tree trunk where early mortality of nymphs in the absence of cannibalism was significantly low in comparison with other types of sites. However, analyses of oviposition showed that more than half of the overlapping ovipositions took place on the same day and that insuch a situation 95% of the lately deposited egg masses escaped cannibalism suggesting existence of avoiding mechanism of cannibalism in adult females under circumtances of ovipositional site limitation.     

Self-/conspecific discrimination and superparasitism strategy in the ovicidal parasitoid Echthrodelphax fairchildii （Hymenoptera： Dryinidae）

Superparasitism in solitary parasitoids results in fatal competition between the immature parasitoids, and consequently only one individual can emerge. In the semisoli- tary ovicidal parasitoid Echthrodelphaxfairchildii （Hymenoptera： Dryinidae）, 2 adults can emerge under superparasitism with a short interval （〈24 h） between the first and second ovipositions. We determined the female parasitoid＇s behavioral responses under self- and conspecific superparasitism bouts with first-to-second oviposition intervals of 〈2 h. The self- and conspecific superparasitizing frequencies increased up to an oviposition interval of 0.75 h, with the former remaining lower than the latter, particularly for oviposition intervals of _〈0.25 h, suggesting the existence of self-/conspecific discrimination. The superparasitizing frequency plateaued for oviposition intervals of _〉0.75 h, with no dif- ference between self- and conspecific superparasitism. The ovicidal-probing frequency did not differ under self- and conspecific superparasitism, and was usually 〈20%. The females exhibited no preference for the oviposition side （i.e., ovipositing on the side with or without the first progeny） and almost always laid female eggs for any oviposition in- terval under self- and conspecific superparasitism. The sex ratio was not affected by the type of superparasitism, oviposition sides, or the occurrence of ovicidal probing. These observed results about the oviposition side, ovicidal probing, and sex ratios differed from the predictions obtained assuming that the females behave optimally. Possible reasons for the discrepancies are discussed： likely candidates include the high cost of selecting oviposition sides and ovicidal probing, and, for the sex ratio, the low frequency of encountering suitable hosts before superparasitism bouts.     

Plant wound compounds from oviposition scars used in host discrimination by a stem-galling sawfly

The stem-galling sawfly Euura lasiolepisuses one or more plant wound compounds resulting from oviposition scars as cues in host discrimination (avoiding sites occupied by conspecifics). Four experiments were conducted to test hypotheses about how Euurapartitions resources. Experiment 1 demonstrated that Euuraavoids ovipositing on nodes with scars from previous ovipositions. Experiment 2 showed no evidence that the sawfly uses oviposition-deterring pheromones and indicated there is a time lag following oviposition before the oviposition scar becomes a deterrent. Experiment 3 showed that sawflies avoid artificially formed scars, demonstrating that a plant cue alone can lead to host discrimination. Experiment 4 showed that visual or tactile cues are not necessary for host discrimination and indicated that a plant wound compound functions as an oviposition deterrent. Both experimental results and field surveys showed that Euuraoviposition scars were more uniformly distributed than expected if sawflies were ignoring previous ovipositions.     

Reproductive behavior of the Japanese treefrog,Rhacophorus arboreus (Anura: Rhacophoridae)

Reproductive behavior of the foam-nesting treefrog,Rhacophorus arboreus, is described. Oviposition was made either by an amplectant pair (a female, and an amplectant male) alone or by an amplectant pair and other males (joiners). The snout-vent length of males and females was negatively correlated with the date of the first appearance in the pond. The period of staying in the pond in males was longer than that in females. Physical body contacts of other males, and encounter calls by ampletant males were frequently observed before oviposition with joiners. Snout-vent length of the female and an amplectant male were positively correlated. Encounter calls by amplectant males were likely to precede movement of females. There was no difference of snout-vent length between amplectant males and joiners. Females moved the hindlimbs more frequently than males during oviposition. Amplectant males made encounter cells more frequently in oviposition with joiners, than oviposition without joiners. The patterns of transitions between acts of females and amplectant males were similar between oviposition with and without joiners. Females more frequently responded to amplectant males than joiners. Both the duration of a bout of the movements of hindlimbs of females and the interval between the bouts were longer in oviposition without joiners than that with joiners. The number of the movements per bout in oviposition without joiners was smaller than that with joiners. Encounter calls by amplectant males during oviposition tended to precede the hindlimb movements of females.     

Influence of Conspecific Presence, Experience, and Host Quality on Oviposition Behavior and Clutch Size Determination in Anastrepha ludens (Diptera: Tephritidae)

Females of some insect species adjust the number of ovipositions and clutch size adaptively depending on conspecific density and probably experience. In a series of three experiments, we examined the effect of the presence of conspecifics, host quality, and oviposition experience on oviposition behavior and clutch size determination by females of the polyphagous fruit fly Anastrepha ludens (Diptera: Tephritidae). In the first experiment, we determined that grouped (eight females per cage) A. ludens females tended to visit and oviposit in more hosts than did solitary females probably as a result of stimulation by the presence of conspecifics. We also determined that females with previous oviposition experience visited and oviposited in more hosts than inexperienced ones. Importantly, when females were grouped, we observed significantly more landings on unoccupied hosts (i.e., devoid of flies) than on occupied ones (i.e., with at least one fly on it). However, oviposition experience, and not female density, was the most important factor affecting clutch size. Naive females deposited larger egg clutches than experienced ones. In the second experiment, we found that oviposition experience and host quality (i.e., clean fruit or fruit covered with a host marking pheromone [HMP] extract), influenced clutch size and the decision of females to defend or not defend the host. Clutch size and number of fights were greater on clean than on HMP-marked hosts. In the third experiment, we observed that host quality (i.e., size) played a significant role with regard to the number of female fights, host marking behavior, and clutch size. Specifically, females fought and dragged their aculeus longer on small- and medium-sized hosts than on large ones. But this behavior varied according to whether females were kept alone or grouped. Clutch size was greatest in the largest hosts. Considering all the above, we believe that the observed increase in ovipositional activity by grouped A. ludens females can be attributed to competition through mutual interference and not social facilitation as has been reported in other tephritid species.     

Effects of Bacillus thuringiensis on Adults of Cotesia plutellae (Hymenoptera: Braconidae), a Parasitoid of the Diamondback Moth, Plutella xylostella (Lepidoptera: Plutellidae)

To complement existing information on the mortality of larvae of the wasp Cotesia plutellae attacking moth caterpillars infected with Bacillus thuringiensis (Bt) we tested the direct and indirect effects of the bacterium on adult wasp longevity and oviposition behaviour. In one experiment with female parasitoids, mean longevity (SEM) was not significantly different between females exposed to Bt (1.98 +/- 0.08 days) and those not exposed (2.18 +/- 0.13 days). In a second experiment with both males and females, Bt treatment did not significantly effect either male or female parasitoids exposed to Bt. To observe the possible effects of Bt on oviposition behaviour of C. plutellae each of 10 females were given five larvae that had been treated with Bt and five untreated larvae at the same time. All parasitoids were observed to make oviposition attempts in both untreated and treated larvae. Upon dissection of the host larvae, one or more C. plutellae eggs were found in each of the larvae in which a parasitoid attempted oviposition. There was no effect of Bt treatment on parasitoid oviposition. The mean number of ovipositions in treated larvae (4.3 +/- 0.3) was not significantly different from untreated larvae (4.7 +/- 0.2).     

Rhythmicity of mating and oviposition inCallosobruchus subinnotatus (Pic) (Coleoptera: Bruchidae)

Mating and oviposition behaviors were studied inCallosobruchus subinnotatus. Copulation was most frequent during the late scotophase, 2–3 h before onset of photophase. The females were less willing to mate during photophase, which increased the time to initiate mating while decreasing the duration of mating. Females exhibited increased movement prior to mating, resting immediately after mating, and remained stationary for 6 h when oviposition commenced. Multiple mating by both males and females affected the number of eggs laid, duration of mating, and uncoupling time at the end of mating. Females that mated two or three times laid more eggs than females that mated once or more than three times. Females that remainedin copula for less than 18 min showed greater readiness to remate than those that remainedin copula longer. There was a gradual decrease in the number of eggs females could lay with an increase in the number of previous matings by males.     

Olfactory Sex Attraction and Mating Behaviour in the Pine Sawyer Monochamus galloprovincialis (Coleoptera: Cerambycidae)

The pine sawyer Monochamus galloprovincialis Olivier (Coleoptera: Cerambycidae) is a vector of the pine wood nematode Bursaphelenchus xylophilus (Steiner and Buhrer) Nickle. Male and females of this species have a kairomonal attraction to host volatiles and Ips semiochemicals. Once on the host tree, males and females copulate and oviposition occurs. Bioassays using Y-tube olfactometry revealed that females were attracted to volatile compounds produced by males, but not to volatiles produced by females. However, immature males did not seem to release attractants for mature females and immature females did not show any attraction for mature males. A experiment designed to know about mating behaviour in this specie revealed that most of the males encountered females while they were walking. In all pairs both sexes contacted with antennae before copulation, however, in some occasions copulation did not proceed just after antennal contact. On the contrary, in all pairs observed, the male licked the elytra of the female with his mouth palpi just before copulation (licking); immediately afterwards, the male grabbed the female with his forelegs and mounted her suggesting a chemically mediated mate recognition by males.     

‘Males second’ strategy in the allocation of sexes by the parasitic wasp,Gryon japonicum

Summary Patterns of the sex ratio allocation of Gryon japonicum (Ashmead) (Hymenoptera: Scelionidae), a solitary egg parasitoid of Riptortus clavatus (Thunberg) (Heteroptera: Alydidae), were investigated in the laboratory, and the result was checked against the field data on the sex composition of the parasitoid. When five host eggs were presented simultaneously to each of the females of G. japonicum in a laboratory experiment, they had a strong tendency to lay a male egg in second host egg and female eggs in the others. However, when four host eggs were presented to each female more than 3 h after the completion of oviposition on a host egg, most of the females laid male eggs in the third oviposition, i.e. the second host eggs after the experimental interruption of oviposition. These results indicated that there was a mechanism for G. japonicum to produce a male egg in the second host egg in consecutive ovipositions, and that the mechanism was reset by more than 3 h intervals of oviposition. By this mechanism, G. japonicum is thought to produce the precise sex ratio in response to the size of a host egg batch. Field data on the size of a host egg batch and the sex composition of the parasitoid in a host egg batch supported this view.     

Increasing the availability of food and the reproduction ofPoecilus cupreus in a cereal field by strip-management

The spruce bud moth,Zeiraphera canadensis Mutt & Free. (Lepidoptera: Tortricidae), displayed a bimodal activity pattern in two young white spruce,Picea glauca (Moench) Voss, plantations. During the day, when temperatures were highest and relative humidities lowest, most moths remained in the lower crown, presumably in response to the risk of hygrothermal stress and/or predation. Flight activity peaks occurred near dawn and dusk. Moths moved higher in the crowns shortly after sunset and many were observed flying above the tree canopy. This activity peak was associated with oviposition and mating. Oviposition started at sunset and continued for approximately 3 h. Females probed oviposition sites with their ovipositor before ovipositing, suggesting that they use sensillae on the ovipositor to evaluate potential oviposition sites. Most mating occurred between 22∶00 and 04∶00 h. Males hovered adjacent to branches containing females before alighting. After alighting, males usually approached to within 2–5 cm of females and then remained motionless for a short time before attempting mounting, indicating that some short-range communication usually occurs prior to mating. Reduced flight activity between midnight and dawn was not due to low temperatures, which were above the minimal temperatures for flight for males (10°C) and females (12°C) determined in the laboratory.