Dynamic egg maturation strategies in an aphid parasitoid

Abstract.  Females of the parasitoid Aphelinus albipodus both lay more eggs on younger stages of their aphid host, Aphis glycines , and contain more mature eggs in their ovaries when held with these younger host stages. This suggests that egg maturation is more rapid in the presence of younger host stages. Three factors explaining the difference in the egg loads of A. albipodus females exposed to various host stages are examined. These are (i) the number of host-feeding meals; (ii) the number of eggs laid; and (iii) the host stage utilized. Together, these factors explain 69% of the variance in A. albipodus egg load. The number of host-feeding meals taken by females is a strong predictor of egg load. More host meals are taken on young hosts, suggesting that host feeding contributes to the trend of faster egg maturation in the presence of younger hosts. Host stage has a strong impact on egg load even when the effect of host feeding is accounted for. There is no evidence for an effect of the oviposition rate on egg load. The results indicate that egg maturation by A. albipodus is dynamic; females mature eggs faster when in the presence of preferred hosts. It is hypothesized that this allows A. albipodus females to more closely match their reproductive effort to reproductive opportunities.     

Influence of host age and host deprivation on longevity, egg load dynamics and oviposition rate in Microplitis rufiventris

Adult size, longevity, egg load dynamics and oviposition ofMicroplitis rufiventris Kok. which began their development in the first, second, third （preferred hosts） or fourth （non-preferred hosts） instar larvae of Spodoptera littoralis （Boisd.） were studied. The parasitoid size was largely determined by the initial host size at parasitism. Non-ovipositing females derived from older hosts lived for longer periods than those derived from younger ones. However, the ovipositing females, irrespective of their size, lived for almost the same periods. At emergence, the oviducts of adult females contain a significant amount of mature eggs available for oviposition for a few hours on eclosion day. Egg load increases during the early phase of adult life. The amount of additional mature eggs and rate of egg maturation per hour was greater for wasps derived from preferred hosts compared with those in females derived from non-preferred hosts. The pattern of egg production in M. rufiventris females depended on the availability of hosts for parasitization. Host-deprived females depleted the egg complement with aging; the longer the host deprivation, the lower the oviduct egg load. Marked reduction in both realized or potential fecundity of host-deprived females was observed following host availability. Host privation for more than 3 days induced a marked deficit fecundity pattern through the female＇ s life. The realized fecundity was determined by the interaction among host availability, the number of eggs that are matured over the female＇ s life span, oviposition rate and host size from which the female was derived. These results suggest that： （i） M. rufiventris wasp is a weak synovigenic species; （ii） the maturation of additional eggs is inhibited once the maximum oviduct egg load is reached; （iii） the egg load of the newly emerged female is significantly less than the realized fecundity; and （iv） because M. rufiventris females oviposit fewer eggs when they begin depleting their egg supply at 3 days, augmentative releases will require release immediately following emergence to ensure the highest parasitization rate in the field.     

Determinants of egg load in the soybean aphid parasitoid Binodoxys communis

We used a series of laboratory studies to investigate factors contributing to variability in egg load of the parasitoid Binodoxys communis (Gahan) (Hymenoptera: Braconidae), a biological control agent of the soybean aphid, Aphis glycines Matsumura (Hemiptera: Aphididae). The change in egg load in newly emerged females over time was determined in response to three treatments: post‐eclosion temperature, sugar meals, and host density. Binodoxys communis females emerge with an average egg load of approximately 40 mature eggs that increases to approximately 200 eggs within 24 h of emergence. The egg maturation rate over this time period is higher when females are held at 26 °C than at 18 °C. And although the egg load of sugar‐fed females was slightly higher than that of starved females, this difference was not statistically significant. Binodoxys communis females that were held with 150 hosts for 8 h laid more eggs than those that were held with 30 hosts, and they also matured more eggs over the subsequent 16 h than those held with 30 hosts or no hosts at all. However, we detected no difference in the egg maturation pattern between B. communis females held with the low host density and that of control females held with no hosts at all. Thus, we conclude that enhanced egg maturation in the higher host‐density treatment is more likely explained by a rapid replenishing of partially depleted ovaries than a host‐induced stimulus of egg maturation per se. Taken together, these results suggest a strategy of maintaining a high egg load and thus avoiding or mitigating the negative effects of egg limitation.     

Superparasitism in the solitary ectoparasitoid Aptesis nigrocincta: the influence of egg load and host encounter rate

Theory predicts that the acceptance of hosts already parasitized by a conspecific will depend both on egg load and the availability of hosts. In the present laboratory study, we tested the effect of egg load and host encounter rate on the propensity of superparasitism in the solitary parasitoid Aptesis nigrocincta Gravenhorst (Hymenoptera, Ichneumonidae), a synovigeneous ectoparasitoid of prepupae of the European Apple Sawfly. Parasitoid females carry few voluminous eggs at a time and the egg maturation rate is less than one egg per day. Egg load was manipulated by giving females access to hosts one week prior to the start of treatments and host availability by giving females access to either one host cocoon every day or every other day. In the first treatment where females had a high egg load of 5.3 egg in their ovaries and encountered host cocoons at low rates, we found that parasitized hosts were accepted to the same degree as healthy hosts. In females with significantly decreased egg load (3.8 eggs) encountering hosts at the same rate we found a slight but non-significant decrease in the acceptance of parasitized hosts compared with healthy hosts. In contrast, A. nigrocincta females accepted significantly fewer parasitized hosts at a high host encounter rate that would lead them to the point of egg limitation in the near future. Within the range of egg loads tested, the host encounter rate appears to be the most important determinant for a females decision to oviposit onto hosts already parasitized by a conspecific.     

Egg size manipulation in the migrant skipper, Parnara guttata guttata (Lepidoptera: Hesperiidae), in response to different host plants

Female adults of the migrant skipper, Parnara guttata guttata, lay different-sized eggs on different host plants in different generations. Moreover, P. g. guttata maintains large egg size variation either in the population or in the individual. Why such phenotypic plasticity in egg size is maintained has not been clarified. In the present study we performed a series of experiments to verify whether or not females of P. g. guttata discriminate between the different host plants, i.e., rice plant Oriza sativa in the first and second generations and cogon grass Imperata cylindrica in the third (overwintering) generation and manipulate egg size accordingly on these host plants. When a caged female was allowed to lay eggs alternatively on soft-leafed rice plant and tough-leafed cogon grass, the size of the eggs laid on cogon grass was significantly larger than that on rice plant. When a caged female was allowed to lay eggs on the two host plants that were supplied on alternate days, the size of eggs laid on cogon grass was also significantly larger than that on rice plant. A preliminary experiment using crude extract from the plants suggested that females did not manipulate egg size in response to chemical stimulants alone. The skipper female was able to lay different-sized eggs instantaneously after assessing the host plant. However, the reaction norm to different host plants differed among females. Eggs of various sizes were matured in each ovariole, and egg size variation at the lowest part of the ovariole ranged from the size of the larger eggs laid on cogon grass to that of the smaller eggs laid on rice plant, although how the female chose the appropriate-sized egg at each occasion is unknown.     

Nocturnal activity and resource utilization in the aphid hyperparasitoid, Dendrocerus carpenteri

Abstract.

• 1 Females of the aphid hyperparasitoid Dendrocerus carpenteri (Curtis) search successfully for hosts during both day and night. Oviposition numbers per host patch did not differ significantly between day and night.
• 2 D.carpenteri females also displayed a nocturnal flight activity, showing that they are not only capable of searching on a given host plant but also of dispersing between host plants.
• 3 Nocturnal oviposition activity was mainly influenced by egg load. Females with a high egg load laid more eggs at night than females with a comparatively low egg load. Thus, D.carpenteri females may use nocturnal foraging to compensate for the lack of oviposition opportunities during day.
• 4 D.carpenteri females which foraged continuously for hosts both day and night (= for 24 h per day) benefitted from an 1.4-fold increase in lifetime reproductive success when compared to females which foraged only by day (= for 16 h per day).
• 5 The benefit of night foraging for this species is a significantly increased reproductive success.

     

Host plant pollen influences calling behavior and ovarian development of the sunflower moth,Homoeosoma electellum

Summary Females of the sunflower moth, Homoeosoma electellum held in the presence of pollen, or an ethanolic pollen extract, from the sunflower Helianthus annuus initiated calling behavior at a significantly younger age following emergence than those provided sucrose only. Furthermore, females with pollen subsequently spent more time calling, and had an increased rate of egg maturation, than those held without pollen. These effects were attributed to a kairomone as females held in the presence of, but denied direct access to, pollen behaved in the same manner as those in contact with pollen. The importance of this life history strategy for the exploitation of temporarily available resources, essential to the survival of neonate larvae, and on the dispersal of adults, is discussed.     

Do Golden Egg Bugs (Phyllomorpha laciniata: Heteroptera, Coreidae) Require Conspecifics for Oviposition?

Golden egg bug (Phyllomorpha laciniata) females lay eggs on the bodies of conspecifics of both sexes. We investigated to what extent reproduction depended on the availability of conspecifics as oviposition substrate and the acceptability of the host plant as an alternative oviposition substrate in the absence of conspecifics. Mated females were placed in experimental enclosures each containing a sprig of fresh host plant. Each experimental female was subjected to one of three treatments: isolated from conspecifics (solitary), paired with another female, or paired with a male. Solitary females laid a few eggs on the host plant but then stopped laying eggs, and solitary females laid significantly fewer eggs than those enclosed with another female or a male. Females enclosed with a male laid no more eggs than those enclosed with a female. When two previously isolated females were later enclosed together, they soon renewed oviposition. Females in nature contained significantly more oviducal eggs than did females that were enclosed with other females for a short period. Thus the availability of suitable conspecifics as oviposition substrate stimulates the deposition of mature eggs, and reproduction depends on the presence of conspecifics of either sex as oviposition substrate.     

Egg maturation dynamics of the parasitoid Microplitis rufiventris: starvation speeds maturation in early life

The number of mature eggs carried by a female parasitoid at any given moment (egg load) is a fitness‐related parameter affecting reproductive potential and impacting upon host population dynamics. Microplitis rufiventris Kokujev (Hymenoptera: Braconidae) is a solitary koinobiont endoparasitoid wasp of several noctuid pests, including Spodoptera littoralis. The number of mature eggs carried by females at emergence is approximately 50. The rate of egg maturation is strongly affected both by feeding status and access to host larvae. In early adult life, egg maturation rates are lower for 6–72 h in fed wasps compared with food‐deprived wasps. When given access to hosts, honey‐fed wasps live for approximately 9 days with high lifetime fecundity (226 eggs). By contrast to early adult life, the total realized fecundity is positively affected by feeding status, where water‐fed and starved females have 140 and 107 eggs, respectively. Egg resorption is most pronounced in the later life of females. The results suggest, in addition to confirming the effect of honey‐feeding on total fecundity, that fecundity of starved wasps includes rapid egg maturation early in life, which potentially could improve the performance of the parasitoid as a biological control agent.     

Egg load of lab-cultured <Emphasis Type="Italic">Anaphes iole</Emphasis> and effects of mate presence and exposure time on load depletion

Anaphes iole Girault (Hymenoptera: Mymaridae) is a solitary egg parasitoid of Lygus bugs (Heteroptera: Miridae) in North America. This research considered factors that might impact the egg load of lab-cultured A .iole females, reared from Lygus hesperus Knight egg patches. The following hypotheses were tested: (1) egg load was related to body size and not affected by female age, and (2) egg load depletion was not affected by mate presence and time (in days) that females were exposed to host patches. Initial egg load averaged 48 mature eggs and no immature eggs were detected in the ovarioles of dissected females. Egg load was neither related to body size (hind tibia or forewing length) nor affected significantly by age (0, 1 or 2days old honey-fed females). Mate presence (females with or without males) and exposure time (1, 3 or 5days on the same host patch) had no effect on egg load depletion. Females usually depleted most of their egg load within 24h. From 86 to 92% of females contained less than six mature eggs and no immature eggs after 1, 3 or 5days of exposure to host patches. The results of this study suggest that A. iole females are certainly pro-ovigenic and initial egg load does not correlate with body size or age. Since mated and unmated females deplete most of their egg load in 24h, time-efficient production of progeny may result when ovipositing parasitoids are exposed to suitable hosts for just a few days.The United States Government has the right to retain a non-exclusive, royalty-free license in and to any copyright of this article. This article reports the results of research only. Mention of a commercial or proprietary product does not constitute an endorsement of the product by the United States Department of Agriculture.     

Fecundity and oviposition ofEucelatoria bryani, a gregarious parasitoid ofHelicoverpa zea andHeliothis virescens

We examined longevity, fecundity, and oviposition strategies ofEucelatoria bryani Sabrosky (Diptera: Tachinidae), a gregarious endoparasitoid ofHelicoverpa zea (Boddie) andHeliothis virescens (F.) (Lepidoptera: Noctuidae). Longevity of adult femaleE. bryani was not related to body size. In contrast to longevity, largerE. bryani females had greater potential fecundity than smaller females, as determined by the number of embryonated eggs present in the common oviduct. However, female parasitoid size did not affect primary clutch size (number of eggs deposited in a host). Because embryos in eggs located in the ovisac were larger than those located elsewhere in the common oviduct, maximum primary clutch size may be physiologically limited by the number of fully mature eggs a female has available at one time.E. bryani females adjusted primary clutch size in response to host size, for bothH. zea andH. virescens. This adjustment appears to be adaptive because females did not overexploit hosts by depositing more larvae than a host could support. Adult emergence was not related to host size. Although host weight positively influencedE. bryani progeny weight, increases in progeny size with host size were counterbalanced by increases in primary clutch size with host size.     

Intraspecific variability in egg maturation patterns and associated life‐history trade‐offs in a polyembryonic parasitoid wasp

1. Life‐history theory predicts a trade‐off between the resources allocated to reproduction and those allocated to survival. Early maturation of eggs (pro‐ovigeny) is correlated with small body size and low adult longevity in interspecific comparisons among parasitoids, demonstrating this trade‐off. The handful of studies that have tested for similar correlations within species produced conflicting results. 2. Egg maturation patterns and related life‐history traits were studied in the polyembryonic parasitoid wasp, Copidosoma koehleri (Hymenoptera: Encyrtidae). Although the genus Copidosoma was previously reported to be fully pro‐ovigenic, mean egg loads of host‐deprived females almost doubled within their first 6 days of adulthood. 3. The initial egg‐loads of newly emerged females were determined and age‐specific realised fecundity curves were constructed for their clone‐mate twins. The females' initial egg loads increased with body size, but neither body size nor initial egg load was correlated with longevity and fecundity. 4. The variation in initial egg loads was lowest among clone‐mates, intermediate among non‐clone sisters and highest among non‐sister females. The within‐clone variability indicates environmental influences on egg maturation, while the between‐clone variation may be genetically based. 5. Ovaries of host‐deprived females contained fewer eggs at death (at ～29 days) than on day 6. Their egg loads at death were negatively correlated with life span, consistent with reduced egg production and/or egg resorption. Host deprivation prolonged the wasps' life span, suggesting a survival cost to egg maturation and oviposition. 6. It is concluded that adult fecundity and longevity were not traded off with pre‐adult egg maturation.     

Onion fly,Delia antiqua,oviposition and mating as influenced by insect age and dosage of male reproductive tract extract (Diptera: Anthomyiidae)

One hundred percent of virgin female onion flies,Delia antiqua, receiving 1/20 of a male equivalent of an aqueous extract of mature male reproductive tract remained unmated in the presence of males and began laying unfertilized eggs at a normally mated rate of about 20 eggs/female/day. The 50% behavioral response (BR₅₀) fell between 1/40 and 1/20 of a male equivalent. Sex peptide responses are not always all-or-none. Some females receiving extract at 1/40 male equivalent oviposited at an intermediate rate. Moreover, at low sex peptide dosages, some females were fully activated ovipositionally but were receiptive to mating. A low level of sex peptide was present in 1-day-old males. Sex peptide titer rose with age until plateauing by 6 days posteclosion. Males began mating at 3 days, when they first had ample mature sperm; 50% of 6-day-old males mated. The mean number of females inseminated per male exposed to an excess of virgin females over 24 h was 4.3±0.6 (±SE). Presence of mature eggs was not always a prerequisite for mating, although probability of insemination was correlated with egg maturation. One-day-old preovipositional females receiving 1/20 of a male equivalent of extract began ovipositing when they had mature eggs at 5–6 days old. Therefore, sex peptide may act early and permanently or have a long half-life and affect behaviors once females reach sexual maturity. Male flies provide females with an excess of sex peptide in many cases.D. antiqua males transferred ca. 5–10 times more sex peptide than necessary to activate females fully. We suggest this excess is related to the speed of female response. It is yet unclear whether sex peptide potency or titer in Diptera has become exaggerated by intra- or intersexual selection.     

The relationship between mating and oogenesis in monarch butterflies (Lepidoptera: Danainae)

We studied the relationship between the timing of mating and oogenesis in monarch butterflies (Danaus plexippus) to determine (1) the potential for male nutrient input into eggs and (2) whether mating stimulates egg development. Most females mated soon after they started maturing eggs. One and 2 days after mating, females contained the same number of mature oocytes as virgin females of the same age, while 3 days after mating they contained more mature oocytes than did virgins. These results confirm the potential for male-derived nutrients to augment oocyte production, but indicate that mating is not required for oocyte maturation to occur.     

椰子木蛾的产卵节律及其对寄主植物的产卵选择性

【背景】椰子木蛾是近年来新入侵我国棕榈科植物的害虫,研究其产卵习性可为监测和防治该虫提供参考。【方法】在室内条件下,观察、记录了椰子木蛾雌成虫的产卵节律及其对不同寄主植物的产卵选择性。【结果】椰子木蛾最高日产卵量可达34.4粒·头-1,且主要集中在羽化后4 d内产卵,占总产卵量的54.1%;产卵活动主要发生在夜间23:00到次日8:00;在椰子、蒲葵、大王棕、槟榔和散尾葵等寄主植物上的产卵量无显著差异,为89.3~147.7粒·头-1,但产卵位置存在差别。【结论与意义】椰子木蛾雌成虫具有较强的繁殖能力和产卵节律性,且在不同寄主植物上的产卵量一致。     

Egg maturation and oviposition strategy of a capital breeder,Cleorodes lichenaria,feeding on lichens at the larval stage

Abstract 1. Capital breeders rely solely on resources acquired before breeding, whereas income breeders may obtain considerable amounts of resources following the commencement of reproduction. In income breeders oviposition occurs over a longer time period with a small number of eggs ready to be laid at the start of adult life, whereas capital breeders reproduce more rapidly and contain higher numbers of mature eggs at eclosion relative to potential fecundity. 2. This study explored egg maturation and oviposition strategy of a nocturnal geometrid moth, Cleorodes lichenaria. It was predicted, based on the long larval period of C. lichenaria and the known biology of other geometrids, that this lichen‐feeding geometrid moth concentrates egg production at the beginning of its breeding period and that external nutrients have a limited role on its fecundity. 3. Approximately 46% of eggs were ready to be laid in newly hatched females and approximately 40% of their potential fecundity was realised during the first night, after which the number and the weight of eggs decreased steadily. Feeding or drinking did not increase the fecundity of females. Pupal mass correlated positively with realised fecundity and ovigeny index. 4. Reproduction of C. lichenaria is based solely on larval‐derived resources. It is suggested that females are able to resorb thoracic tissues to increase fecundity and that this ability is probably linked to relatively long lifespan and low ovigeny index of this species, characteristics rarely observed in other geometrid moths.     

Superparasitism in the solitary ectoparasitoid Aptesis nigrocincta: the influence of egg load and host encounter rate

Theory predicts that the acceptance of hosts already parasitized by a conspecific will depend both on egg load and the availability of hosts. In the present laboratory study, we tested the effect of egg load and host encounter rate on the propensity of superparasitism in the solitary parasitoid Aptesis nigrocincta Gravenhorst (Hymenoptera, Ichneumonidae), a synovigeneous ectoparasitoid of prepupae of the European Apple Sawfly. Parasitoid females carry few voluminous eggs at a time and the egg maturation rate is less than one egg per day. Egg load was manipulated by giving females access to hosts one week prior to the start of treatments and host availability by giving females access to either one host cocoon every day or every other day. In the first treatment where females had a high egg load of 5.3 egg in their ovaries and encountered host cocoons at low rates, we found that parasitized hosts were accepted to the same degree as healthy hosts. In females with significantly decreased egg load (3.8 eggs) encountering hosts at the same rate we found a slight but non‐significant decrease in the acceptance of parasitized hosts compared with healthy hosts. In contrast, A. nigrocincta females accepted significantly fewer parasitized hosts at a high host encounter rate that would lead them to the point of egg limitation in the near future. Within the range of egg loads tested, the host encounter rate appears to be the most important determinant for a females decision to oviposit onto hosts already parasitized by a conspecific.     

Maternal effects on progeny size, number and body color in the desert locust, Schistocerca gregaria: Density- and reproductive cycle-dependent variation

The effects of rearing density and maternal age on the progeny size, number and coloration of the desert locust, Schistocerca gregaria, were investigated. Isolated-reared females deposited smaller, but more eggs than crowd-reared females. The former produced smaller and more eggs with age, whereas the latter showed a tendency to produce larger and fewer eggs over time. A similar tendency was also observed with virgin females, indicating that mating or the presence of males was not important. The first egg pod produced by each mated crowd-reared female contained significantly smaller and more eggs than did the subsequent egg pods. The former often produced many green hatchlings (0-100%) characteristic of solitarious forms, whereas the egg pods deposited after the first pod produced predominantly black hatchlings typical of gregarious forms. Adults were highly sensitive to a shift in rearing density and quickly modified the quality and quantity of their progeny depending on the density encountered. The number of eggs per pod was influenced not only by the mother's rearing density but also by rearing density of the grandmother. The present results demonstrated that the characteristics of progeny are influenced not only by the crowding conditions experienced by the mother and grandmother but also by the mother's reproductive cycle.     

Mating behaviour of the corn stalk borerSesamia nnonagrioides

The mating behaviour of the corn stalk borerSesamia nonagrioides (Lef.) [Lepidoptera:Noctuidae] was studied under laboratory conditions at 25 ± 1 °C, 65 ± 5% r.h. and 16:8 (L:D) regime. The females began calling during the first scotophase following emergence, the peak of calling occurred during the second scotophase and thereafter decreased. Maximum calling was observed between the fourth and sixth hour of the scotophase. The calling pattern varied with age. During the first and second scotophase most of the females were calling continuously, while during the third and fourth scotophase periodic calling was observed. The mean onset of calling was advanced and the mean length of calling per day was increased slightly as the female became older. Females held for 72 h in continuous darkness exhibited an endogenous circadian rhythm of calling. Matings began during the first scotophase following emergence and the peak of mating was observed during the second scotophase. Mated females did not remate. Few males mated more than once during the following scotophases.     

Fecundity and Longevity of Green Vegetable Bug,Nezara viridula, Following Parasitism byTrichopoda giacomellii

The effects of parasitism by the ArgentinianTrichopoda giacomellii(Blanchard) on reproduction and longevity of its host,Nezara viridula(L.) are reported. Parasitoid larvae suppress egg maturation, reducing by 70% the fecundity of mature female hosts during the period of larval development. Egg viability was not affected, but mating frequency was reduced by approximately 50%. When parasitized as newly eclosed adults, 84% of females fail to reproduce. In male hosts, fertility and mating frequency were not affected during the period of larval parasitoid development. In male and reproductively immature female hosts, death was coincident with, or occurred shortly after parasitoid emergence (2–4 days); in mature females, death occurred on average 2 weeks after larval parasitoid emergence. Host mortality occurred as a consequence of tissue damage incurred as the parasitoid larvae emerged from the host. Some individuals survived parasitism though no further reproductive activity (mating or oviposition) occurred. The effectiveness ofT. giacomelliias a biological control agent is discussed in relation to its impact on reproduction and survival of its host and contrasted with the action of otherTrichopodaspecies.