Egg Deposition Behavior in the Haplodiploid Sawfly Athalia rosae ruficornis Jakovlev (Hymenoptera: Symphyta: Tenthredinidae)

The egg deposition behavior of the turnip sawfly, Athalia rosae (Hymenoptera: Symphyta), is described. Both unmated and mated females lay eggs individually inside of fresh young leaves of cruciferous plants. During an oviposition event, females exhibit a distinct pause in abdominal contractions just before the actual egg deposition act. Unmated females show a longer pause (11.31 s on average) than mated females (4.38 s on overall average). By employing an eye color mutation, the sex of the eggs laid by females was ascertained. Females mated once lay mostly fertilized (diploid female) eggs initially but begin to lay a considerable number of unfertilized (haploid male) eggs later in life. The laying of an unfertilized egg is associated with a longer pause (6.98 s on average) than the laying of a fertilized egg (3.76 s on average). These results are in contrast to previous reports on apocritan Hymenoptera, where the presence of a pause or a longer pause during oviposition was associated with the deposition of fertilized eggs rather than unfertilized eggs. The possibility that mated Athalia rosae females control fertilization and its implications for sex allocation strategies are discussed.     

Life history characteristics of three types of females in Brachionus calyciflorus Pallas (Rotifera) fed different algae

This study describes the life history characteristics of amictic, unfertilized mictic and fertilized mictic females of the rotifer Brachionus calyciflorus cultured individually on two different algae at 0.1 mg ml^–1 food concentration and 27 °C. The duration of the juvenile period of amictic females was significantly shorter on Chlorella pyrenoidosa Chick than on Scenedesmus obliquus Kütz or both algae together. The duration of the juvenile period of unfertilized mictic females was significantly longer, and the number of eggs produced by amictic females was significantly larger on Chlorella pyrenoidosa than on S. obliquus. When fed the same type of alga, the duration of the juvenile period of the fertilized mictic females was the longest among the three types of females, and the durations of the reproductive period of the amictic females and the post-reproductive period of the fertilized mictic females were longer than, or equal to those of the other two types of females, respectively. The number of eggs produced by an unfertilized mictic female was the largest among the three types of females, and that of amictic females was larger than or equal to that of fertilized mictic females, depending on the type of diet.     

Fusion of fertilized and unfertilized sea urchin eggs : Maintenance of cell surface integrity

Here we report that immediately after the fusion of a fertilized and an unfertilized egg, the two halves of the fused egg retain their respective cell surface organizations. Long microvilli are present on that area of the surface contributed by the fertilized egg, and the unfertilized portion remains comparatively smooth. Cortical granules are absent in the cortex contributed by the fertilized egg, whereas these organelles are present in the cortex of the unfertilized portion. There are distinct boundaries formed by the presence or absence of long microvilli and of undischarged cortical granules. However, following the synchronous prophase of the two nuclei, the original fertilized and unfertilized portions are no longer distinguishable. The observations indicate that the unfertilized portion of the fused egg is capable of maintaining its original surface properties but can, during prophase, undergo changes equivalent to those that take place at fertilization.     

Periodicity of oviposition,feeding, and calling by mated femaleHeliothis virescens in a field cage

This paper describes behaviors exhibited by mated, 3-day-old female Heliothis virescensin the presence of flowering cotton infield cages in mid-July. Females exhibit discrete periods of egg deposition, calling, and feeding during scotophase. Females exhibit behaviors such as walking while wing fanning, wing buzzing, antennal movement, bending of abdomen, and dragging of the ovipositor on the substrate before egg deposition. These behaviors are interspersed with flight and calling. The number and duration of some behaviors differ when the female is ovipositing on cotton plants versus cage screen. Egglaying occurs in several major bouts through the scotophase with periods of inactivity or other activities interspersed. Most eggs are laid before 0300 h. Significant calling is observed after 0100 h and continues until about 0430 h. Although females exhibit some feeding near sunset, the major feeding activity is seen just prior to and after sunup.     

Biology of Leipothrix dipsacivagus (Acari: Eriophyidae), a candidate for biological control of invasive teasels (Dipsacus spp.)

The present study describes key aspects of the biology of Leipothrix dipsacivagus, an eriophyid mite that is under study as a biological control candidate of Dipsacus fullonum and D. laciniatus (Dipsacaceae). Preliminary host-specificity tests have shown that it can develop and reproduce only on Dipsacus spp. (teasels). Studies were conducted in a laboratory at 26 ± 2^oC with 16 h of light per day. Mites for the stock colony were collected from D. laciniatus in Klokotnitsa, Bulgaria and reared on rosettes of D. laciniatus in the laboratory. Unfertilized L. dipsacivagus females reared in isolation from the juvenile stage produced male offspring only, while progeny of fertilized females were of both sexes, suggesting arrhenotokous parthenogenesis with haplodiploid sex determination. Experiments were designed to compare male progeny from fertilized females to males from unfertilized females and to compare males and females from fertilized females. Male progeny of virgin mothers had significantly longer durations of active immature stages and total egg-to-adult period than male progeny of fertilized females. Female progeny had significantly longer durations of egg incubation, active immature stages and egg-to-adult period than male progeny from fertilized mothers. Adult longevity was significantly greater in females than in males. Fertilized females produced significantly more eggs per day and overall than virgin females. The results of this study suggest that fertilization status of L. dipsacivagus females can affect both their own fecundity and the development of their male progeny.     

Microhabitat selection and audible sexual signalling in the wolf spider Hygrolycosa rubrofasciata (Araneae, Lycosidae)

We studied the microhabitat selection and male sexual signalling behaviour in the wolf spider Hygrolycosa rubrofasciata (Ohlert). Males strike dry leaves with their abdomen, producing an audible sexual drumming signal, and females use this signal to choose mating partners. In the field we followed male drumming rate and microhabitat selection using both the mark–recapture method and direct observations. In the laboratory we conducted an experiment on male microhabitat and drumming substrate selection. We found that in the field males were not distributed randomly among the habitat; fewer males were found in areas that had high sedge cover, low elevation, and low dry leaf cover. In the laboratory experiment, males spent more time on dry leaf substrate. Drumming rate in the field was positively correlated with dry leaf cover and in the laboratory males clearly preferred dry leaves as drumming substrate. Temperature was positively correlated with male drumming rate, and with male and female mobility. We conclude that in H. rubrofasciata male distribution and sexual signalling rate, and thus mating success, are greatly affected by environmental factors. Therefore, males may be sexually selected to make effective use of their signalling habitat by active microhabitat choice. Received: 27 December 1999 / Received in revised form: 14 February 2000 / Accepted: 14 February 2000     

草地贪夜蛾生殖行为及其昼夜节律研究

草地贪夜蛾Spodoptera frugiperda是当前跨国迁飞性重大害虫,云南是其重要虫源地及其向内地迁移的重要通道。生殖行为是性选择和物种进化的重要基础,基于生殖行为的害虫监测和防治策略也有重要价值。本研究模拟自然条件下可能出现的择偶情况,采用永久配对和每日更换配偶两种处理,对草地贪夜蛾成虫一生(约10 d)的生殖行为和节律进行了连续观测。结果表明,该蛾在羽化当日(0日龄)即出现召唤和求偶行为但无交配发生,在1日龄开始出现交配行为并在2日龄开始产卵,从2日龄到6日龄期间一直保持了较高的生殖活动,之后开始下降。雄虫求偶与雌虫召唤在节律上有较高的同步性,交配高峰多出现在召唤和求偶高峰之后。每日更换配偶与永久配对相比,雌蛾表现出较高的求偶频率和持续时间,较高交配比率和频率。雌虫在遇到新配偶或与新配偶交配后,会延缓产卵或下调产卵速率。从性选择理论来看,这些行为策略将有助于雌虫获得物质或基因利益,值得进一步探究。交配次数、召唤时长与雌虫寿命之间相关性不显著,但前5 d产卵速率与雌虫寿命间存在显著负相关,即早期产卵量较高的雌虫寿命较短,符合早期繁殖-后期存活基因拮抗假说。     

Oviposition in <Emphasis Type="Italic">Heliconius erato</Emphasis> (Lepidoptera,Nymphalidae): how Essential Is Drumming Behavior for Host-Plant Selection?

Quantitative studies on the behavioral events involved in oviposition by phytophagous insects are scant. One of these events is drumming behavior, a rapid extension of forelegs after landing on a host, which remains largely under studied. In this study, quali- and quantitative analyses of Heliconius erato phyllis (Lepidoptera: Nymphalidae) oviposition events and drumming behavior in relation to both preferred (Passiflora misera and P. suberosa) and non-preferred (P. caerulea and P. alata) hosts were performed. For the first time, drumming behavior is described by images made during the butterfly oviposition process. The same set of females was assigned to both single- and multiple-choice tests under insectary conditions, and their behaviors toward each host were recorded on video. The resulting images were analyzed frame by frame. Both frequency and duration of behavioral events related to oviposition varied on preferred vs non-preferred hosts. On preferred hosts, behaviors that were associated with egg deposition (inspecting flight, drumming and abdomen touching) were pronounced; subsequently, oviposition occurred within a few seconds. On non-preferred host plants, behaviors that did not predict oviposition (flying and resting) predominated, and oviposition on these hosts was negligible. Drumming occurred on all plants and resulted in a faster decision-making process on high-quality hosts (prompt acceptance) as well as on lethal hosts (prompt rejection) compared to the host that confers poor larval performance (delayed rejection). Thus, drumming is crucial for decision-making related in host-plant selection by H. erato phyllis, and influences other behaviors involved in the oviposition process.     

The reproductive program of female Varroa destructor mites is triggered by its host, Apis mellifera

Reproducing Varroa females begin oviposition on a host larva by laying an unfertilized (male) egg, followed by fertilized (female) offspring. Using transfer experiments, we examined whether the sequence of sexes in the brood cell is triggered by a host stimulus. When reproducing Varroa females were transferred from white-eyed pupae (worker brood) into freshly capped worker brood cells, 77% (n = 22 fertile mites after the transfer) began a new reproductive cycle by laying a male egg. The proportion of brood cells with male offspring was similar to naturally infested brood cells. Varroa females transferred into brood cells with young pupae reproduced, but only 6% (n = 16 fertile mites after the transfer) produced male offspring. This was significantly different from male production in naturally reproducing Varroa females and those transferred into freshly capped brood cells. We conclude that a host stimulus present in freshly capped brood cells triggers both the start of reproduction and the sequence of sexes.     

The courtship and spawning behaviors ofCanthigaster valentini (Tetraodontidae)

Synopsis The sharpnose puffer,Canthigaster valentini, occurs in male-dominated harems of territorial females at Lizard Island (Australia). Spawning occurs daily and year-round. Males only spawn with the females of their social group, inside the female's territory. Females signal their readiness to begin courtship with a display that exposes their egg swollen abdomen. Courtship is long and males do not develop specific courtship colors or perform unique displays. Eggs are fertilized in a demersal nest and there is no parental care.     

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.     

The effect of host-plant quality on the survival of larvae and oviposition by adults of an ant-tended lycaenid butterfly, Jalmenus evagoras

Abstract.

• 1 Juveniles of the Australian lycaenid butterfly, Jalmenus evagorus (Donovan), secrete to ants a solution of sugars and amino acids, primarily serine. The attendant ants protect the larvae and pupae from parasites and predators.
• 2 The effect of caterpillar nutrition on the defence provided by ants was investigated. Potted food plants of Acacia decurrens were either given water containing nitrogenous fertilizer or were given water alone. Fertilized plants had a higher nitrogen content than unfertilized plants.
• 3 Fifth instar larvae of J.evagoras feeding on fertilized plants attracted a larger ant guard than those feeding on unfertilized plants. In the absence of caterpillars, ants were not differentially attracted to fertilized and unfertilized plants.
• 4 In the presence of ants, over a 10-day period, larvae on fertilized plants survived better than larvae on unfertilized plants. In the absence of ants larvae survived equally on fertilized and unfertilized plants. It is concluded that larvae on fertilized plants attracted a larger ant guard, and thereby survived better, than larvae on unfertilized plants.
• 5 Adult females of J. evagoras preferred to lay egg batches on fertiized, rather than unfertilized plants, but they did not lay larger egg batches.

     

Western corn rootworm (Diabrotica virgifera virgifera LeConte) (Coleoptera: Chrysomelidae) oviposition can be released by decapitation

Nervous system control of western corn rootworm (Diabrotica virgifera virgifera LeConte, WCR) (Coleoptera: Chrysomelidae) oviposition was studied using decapitation. Gravid females (n = 364) were decapitated with scissors and floated in water-filled Petri dishes. Oviposition by individuals was observed at 30 min intervals for 4 hr after decapitation; cumulative oviposition was tallied at 48 hr post-treatment. Overall, 82.7% of females laid eggs within 48 hr. Oviposition commenced quickly; 78.4% of females laid eggs during the observation period, 81.8% of these began egg laying within 30 min of decapitation. Egg-laying females deposited a total of 66.8 ± 2.1 eggs (mean ± SEM); this was 85.1% of the total mature egg load. Dissection revealed that 31.9% of n = 301 laying females and 14.3% of 63 non-laying females had a rupture of the common oviduct, manifest as an egg-filled hernia containing a mean of 8.17 ± 1.3 eggs (range: 1–83). Among n = 237 females that laid eggs during observations, females with hernias laid significantly fewer eggs (35.8 ± 4.2) than intact females (48.0 ± 2.7) during the 4-hr interval. There was no difference in the mean proportion of hatch for eggs collected from the same n = 10 females before (0.88 ± 0.03) or after decapitation (0.84 ± 0.04). Rapid oviposition following decapitation suggests that WCR egg laying is under constant descending neural inhibition; the motor programme controlling egg laying must reside posterior to the head. Decapitation can be used to quickly collect mature, fertilized WCR eggs.     

Carbohydrate accumulation and utilization by oocytes of Rhodnius prolixus

The processes of accumulation and mobilization of carbohydrate stores in eggs of Rhodnius prolixus were analyzed. During oogenesis, the total amounts of glycogen, glucose, and trehalose increased with an accumulation of proteins, especially when oocytes grew from 1.0 to 1.5 mm in length. At 2.0 mm length, when oocytes were ready for oviposition, nutrient reserves did not increase appreciably and trehalose content decreased. Mating did not affect the final content of carbohydrates or proteins in oocytes of mated and virgin females. A trehalase activity was detected in follicles containing vitellogenic oocytes, 1.0 and 1.5 mm length, in both mated and virgin females. This activity was extremely low in chorionated, 2.0-mm oocytes. After oviposition, glycogen content decreased in fertilized eggs, but not in unfertilized ones, and some was present in newly hatched nymphs. Glucose content remained constant in unfertilized eggs, but increased in fertilized ones, while total protein amount was constant in both groups after egg laying.     

Chemical signals mediating interactions betweenGaleruca tanaceti L. (Coleoptera, Chrysomelidae) and its egg parasitoidOomyzus galerucivorus (Hedqvits) (Hymenoptera, Eulophidae)

Chemical signals mediating interactions betweenGaleruca tanaceti and its egg parasitoidOomyzus galerucivorus (Hymenoptera, Eulophidae) were studied. Neither odor of gravid females ofG. tanaceti nor volatiles of their feces were attractive to the parasitoid. However, the presence of the beetles’ feces on a substrate arrested the parasitoid and elicited frequent antennal drumming. Thus, feces may contain a kairomone important for host finding. Odors of damaged and undamaged host plants had no effect on the parasitoids.O. galerucivorus did not detect its host eggs at close range but encountered them by chance. Neither the structure nor the dark color of the egg surface play a key role in host recognition, but chemicals of the extrachorion which could be isolated by dichloromethane. Fractionation of the dichloromethane extract by TLC revealed a single active fraction which induced host recognition behavior. Since the eggs ofG. tanaceti contain anthraquinones and anthrones which are active as feeding deterrents against predators, we hypothesized that reproductive success ofO. galerucivorus is due to sequestration of these protective compounds. However, GC-MS analyses revealed that there was no transfer of them from the host egg into the adult parasitoid.     

Reproductive mode of the geographic parthenogenetic mayfly Ephoron shigae, with findings from some new localities (Insecta: Ephemeroptera, Polymitarcyidae)

The burrowing polymitarcyid mayfly Ephoron shigae is distributed widely in Japan. Some populations are bisexual, others are unisexual, and the distributions of the two types overlap broadly. Experimental evidence of parthenogenetic reproduction, long suspected in unisexual populations, is presented here, based on a comparative analysis of the developmental rate of fertilized and unfertilized eggs. The developmental rate of fertilized eggs from 20 mated females in a bisexual population was 98.4% ± 0.73% (mean ± SD), and no unfertilized eggs from 20 virgin females in that population developed. The developmental rates of unfertilized eggs in two unisexual populations were 89.0% ± 4.59% and 84.2% ± 1.96%, respectively. This article presents experimental evidence of geographic parthenogenesis in E. shigae and provides support for the previous interpretation. In addition, we discuss the relationship between the sex ratio of each population and the developmental rate of fertilized versus unfertilized eggs from the females in those populations.     

Effect of host deprivation on egg quality,egg load,and oviposition in a solitary parasitoid,Chetogena edwardsii (Diptera: Tachinidae)

Solitary parasitoids are limited to laying one egg per host because larvae compete within hosts. If host encounter rate is low, females should not increase the number of eggs/host in response. The tachinid fly, Chetogena edwardsii,was used to evaluate the effect of host deprivation on egg accumulation, oviposition behavior, and egg quality in a solitary parasitoid. Females deprived of hosts for 2– 7 days accumulate about 1 day's supply of eggs. Egg output of deprived females once hosts are restored does not differ from that of control females. Deprived females retain one egg in the uterus where it undergoes embryogenesis. Maggots emerging from retained eggs are more likely to survive in hosts molting in 40 h or less after receipt of an egg than are maggots emerging from eggs fertilized shortly before oviposition. Egg retention is a consequence of host deprivation that permits females to broaden the range of hosts they can exploit to include soon-to-molt hosts and possibly multiply parasitized hosts.     

Genetics of female functional virginity in the parthenogenesis-Wolbachia infected parasitoid wasp Telenomus nawai (Hymenoptera: Scelionidae)

A lepidopteran egg parasitoid species Telenomus nawai consists of two distinct populations with different reproductive modes. One is a completely thelytokous population consisting of females only, whereas the other displays arrhenotokous reproduction where fertilized eggs develop into diploid females and unfertilized eggs into haploid males. Thelytoky in T. nawai is caused by a bacterial symbiont, the parthenogenesis-inducing (PI) Wolbachia. Recent theoretical studies have shown that when a PI-Wolbachia is spreading in a population, mutations that allow uninfected females to produce more male offspring will spread rapidly eventually becoming fixed. The consequence of such a mutation is that sexual reproduction is no longer successful in infected females. Here we determine the genetic basis of the females' inability to reproduce sexually by introgressing the genome of a thelytokous line into an arrhenotokous line. The results suggest that the mutations are recessive and inherited either as a single-locus major gene with some modifiers, or as two partially linked loci.     

Are individuals from thelytokous and arrhenotokous populations equally adept as biocontrol agents? Orientation and host searching behavior of a fruit fly parasitoid

Hymenopteran parasitoids generally reproduce by arrhenotoky, in which males develop from unfertilized eggs and females from fertilized eggs. A minority reproduce by thelytoky, in which all-female broods are derived from unfertilized eggs. Thelytokous populations are potentially of interest for augmentative biological control programs since the exclusive production of females could significantly lower the costs of mass rearing. Behavioral traits are a major component of parasitoid efficacy. Here, we examined orientation and host searching behavior in thelytokous and arrhenotokous populations of the fruit fly parasitoid Odontosema anastrephae Borgmeier (Hymenoptera: Figitidae). Orientation behavior to various odorant sources was studied in a two-choice olfactometer. No major differences were found between thelytokous and arrhenotokous wasps for this behavior. However, when host-searching behaviors were analyzed, some differences were found. Thelytokous females arrived sooner, foraged longer, and remained longer on non-infested guavas than arrhenotokous females. Individuals of both forms exhibited similar stereotyped behavioral sequences vis-à-vis guava treatments, with only slight deviations detected. Our results suggest that individuals from selected thelytokous and arrhenotokous O. anastrephae populations have similar abilities to search for tephritid larvae, supporting the use of thelytokous strains for augmentative releases.     

Quantitative analysis of ovipositional behavior: Effects of a host-plant chemical on the onion fly (Diptera: Anthomyiidae)

Behavioral responses of female onion flies, Delia antiqua (Meigen), to hostplant cues were quantified during encounters of individual flies with onion plants and onion foliar surrogates. The behavioral repertoire of such females included sitting, grooming, running up and down foliar surfaces, extension of the proboscis such that the labellum contacted foliar and soil surfaces, movements of the tip of the abdomen over surfaces (surface probing), subsurface probing of soil crevices with the ovipositor, and oviposition. Sequences of behaviors preceding oviposition were probabilistic rather than highly stereotyped but generally followed the order given above. Foliar surrogates were used to determine the effects of n-dipropyl disulfide (Pr₂S₂) on the sequence of behaviors leading up to oviposition. The addition of a Pr₂S₂-treated surrogate to a cage increased the frequency of alighting on that surrogate but also increased alighting on a nearby foliar surrogate without Pr₂S₂. After alighting, females encountering surrogates treated with Pr₂S₂ had shorter latencies to proboscis extension and surface probing, spent less time sitting and grooming, and had runs of shorter duration. These females were also more likely to make the transition from probing of surfaces of foliage and soil to subsurface probing of soil crevices and oviposition. Thus, rather than mediating a particular step in the behavioral sequence, Pr₂S₂ played a role throughout the sequence leading up to oviposition. Collectively, these data and past studies on the onion fly support the hypothesis that egglaying is triggered by a temporal summation of inputs to the central nervous system from various sensory modalities rather than strict behavioral chaining, with each transition effected by some unique cue.