Egg distribution patterns in the almond seed wasp,Eurytoma amygdali

The egg distribution patterns ofEurytoma amygdali Enderlein (Hymenoptera, Eurytomidae), which oviposits singly in green, developing almonds, were studied in the laboratory and in the field. In the laboratory, individual females were caged with a number of almonds and the eggs deposited in each fruit were counted. In the field, eggs were censused in almonds of different varieties at regular intervals, over four seasons (1988–91). In the susceptible ‘Retsou’ variety, eggs were uniformly distributed among fruits, both in the laboratory and in the field, as long as the mean number of eggs per almond was ≤2.5. When the mean number of eggs per almond was higher the egg distributions were random. This suggests that, up to a certain level of infestation, females were able to assess egg load of fruits and oviposit in the less infested ones. The main factor enabling the females to discriminate and select the less infested almonds for oviposition is probably a host-marking pheromone. In 1990, the same patterns of egg distribution were observed in samples taken from 5 other almond varieties that are not as susceptible as Retsou and have thicker pericarp and harder endocarp. In 1991 however, when fruits developed more rapidly than in 1990, egg distributions in 3 of these 5 varieties (Truoito, Ai, and Marcona) were not uniform. Although the mean number of eggs per fruit was low (1.2–2.0), many fruits of these varieties contained no eggs. This suggests that, in some less susceptible varieties, egg distribution might also be affected by certain fruit parameters, unfavourable for oviposition, related to the pericarp thickness and endocarp hardness.     

Patch marking in the aphid hyperparasitoid,Dendrocerus carpenteri: the information contained in patch marks

The aim of this study was to test if females of the aphid hyperparasitoid Dendrocerus carpenteri (Curtis produce patch marks. We tested if these marks inform a foraging female of the identity of the producer of the mark (the female herself or a conspecific female) and on the producer's success or failure in finding hosts in the marked area. We also tested if the responses to patch marks differ depending on the size and/or egg load of the female. On average, females walked shorter paths and spent less time in previously explored areas in comparison to control areas only if the area had first been explored successfully (host found) by a conspecific female. If no host had been found or if the area had been explored by the same female previously, no differences between average values were recoreded. However, egg load also seemed to influence foraging decisions in those experiments where average values were not different between previously explored and control areas. Females with a low egg load tended to spend less time in previously explored areas than females with a high egg load. Average values therefore somehow obscured the individual responses to pathc marks. The results suggest that at least D. carpenteri females with a low egg load continuously apply a marking pheromone while walking. This pheromone seems to contain information on the identity of the producer. In addition, different pheromones seem to be applied depending on whether or not hosts are present in the area.     

Size specific demography of three species of Anastrepha fruit flies

Individuals of three Anastrepha species: A. obliqua, A. ludens, and A. serpentina (Diptera: Tephritidae), were sorted according to pupal weight in cohorts of large and small flies. Demographic parameters and reproductive patterns and heterogeneity were determined for each cohort. Large flies of the three species presented greater expectation of life and gross fecundity rates. A. ludens was the species with the longest life span (expectation of life of large adults was 110 days) and the greatest gross fecundity rates (1597 eggs/female for the large flies). While, A. obliqua had the shortest mean age of reproduction (33 days), and the greatest daily egg production (14 eggs/female/day). Net fecundity was similar in these two. A. serpentina had lower fecundity rates.Reproductive information for each size and each species include: age-by-parity relations, fraction of sexually mature life in which females lay eggs, and frequency distribution of individual egg production. Results demonstrate that even under constant laboratory conditions and using standard artificial hosts, there is a great deal of life history variation among these Anastrepha species and among other tephritid fruit flies.     

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.     

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.     

Influence of Host Pheromone on Egg Parasitism by Scelionid Wasps: Comparison of Phoretic and Nonphoretic Parasitoids

The kairomonal activity of the attractant pheromone for the spined soldier bug, Podisus maculiventris (Hemiptera: Pentatomidae), was investigated by exposing fresh pentatomid egg masses in field traps with or without synthetic pheromone. Predominantly two parasitoids were recovered from exposed eggs of P. maculiventris and Euschistus obscurus: Telenomus podisi Ashmead (a generalist pentatomid egg parasitoid) and Telenomus calvus Johnson (a phoretic specialist on Podisus eggs) (Hymenoptera: Scelionidae). The incidences of T. podisi from P. maculiventris and E. obscurus eggs placed in pheromone-baited and nonbaited traps were not significantly different, suggesting that this oophagous wasp does not use the spined soldier bug attractant pheromone as a kairomone. However, T. calvus was reared almost exclusively from egg masses of P. maculiventris placed inside pheromone-baited traps. These results suggest that T. calvus females orient to volatile chemicals emitted by spined soldier bug males as a searching strategy to find areas likely to contain host eggs, in addition to the previously discovered strategy of using the pheromone to guide their phoretic behavior. The data also indicate that T. calvus can distinguish between the egg masses of these pentatomid hosts at close range.     

A novel approach to the assessment of fruit damage caused by <Emphasis Type="Italic">Carposina sasakii</Emphasis> (Lepidoptera: Carposinidae) using pheromone trap catches in Korean orchards

This study reports a model that utilizes pheromone trap catches to assess the fruit damage caused by Carposina sasakii. The model consisted of four steps: (1) obtaining influx population density using pheromone traps, (2) estimating the actual female population within a defined area using an estimated conversion rate, (3) calculating the total number of eggs using the oviposition model of C. sasakii, and (4) estimating the proportion of fruits infested with eggs (potential damaged fruits) using the relationship between mean egg density per fruit and the proportion of fruits infested with eggs. The relationship between mean egg density ([`(x)] \bar{x} ) per fruit and variance (s ²) was well described by Taylor’s power law, and its parameters were successfully incorporated into the equation that estimates the relationship between mean egg density and the proportion of fruits infested with eggs. In peach orchards, the model accurately predicted the proportion of fruits infested with eggs at the beginning of C. sasakii emergence in early season, but overestimated it in the mid-season. The fitting ability of the model outputs largely increased when the factor of oviposition behavior of C. sasakii was incorporated into the simulation processes, applying the allocation module of total eggs between peaches and apples.     

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.     

Egg resorption behaviour by the solitary egg parasitoid Anaphes nitens under natural conditions

We investigated under outdoor conditions and inside a climate chamber: (i) whether Anaphes nitens Girault (Hymenoptera: Mymaridae), an egg parasitoid of the Eucalyptus snout beetle, Gonipterus scutellatus Gyllenhal (Coleoptera: Curculionidae), experienced egg resorption, and (ii) how various treatments (location, food, and/or host supply) and body size influenced egg load, egg resorption, fecundity, and longevity. One‐day‐old females were mated and randomly assigned to five groups: (A) honey + hosts, (B) water + hosts, (C) only honey, (D) only water, and (E) control females killed at emergence. We compared the egg load of the newly emerged females, which represent the control group (egg load = ovarian eggs present at emergence), with the lifetime egg load (i.e., ovarian eggs + emerged offspring + not emerged offspring) of the females with various host and diet treatments, by dissection of the ovaries to find evidence of egg resorption. All groups reared outdoors had fewer eggs than the control, while indoors there was no significant difference. Outdoors, starved but host‐provided wasps (B) experienced the highest reduction of the lifetime egg load (51%). Groups without access to food (B + D) resorbed more eggs than groups provided with honey (A + C). Females with honey and hosts (A) had the highest lifetime fecundity, but those with water and hosts (B) showed a higher daily realized fecundity. Host‐deprived females with access to food (C) attained the longest lifespan. Our results suggest that under stress conditions, such as low temperature and food shortage, A. nitens females practice egg resorption, probably to save energy.     

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.     

Facilitation in Ovipositional Behavior of Bactrocera tryoni Flies

Mature female Queensland fruit flies, Bactrocera tryoni Froggatt, released into canopies of field-caged trees containing host fruit, did not discriminate between fruit having or lacking two resident conspecific females when selecting fruit on which to alight. When released individually onto a fruit, mature females showed no greater or lesser propensity to initiate boring with the ovipositor into fruit having or lacking a resident female engaged in pre-ovipositional or post-ovipositional behavior. However, boring propensity of introduced females was significantly enhanced or facilitated when released onto fruit on which a conspecific female was in the act of ovipositing. Introduced females were significantly more prone to initiate boring into high-ranking fruit (nectarines) than low-ranking fruit (lemons) and to do so when having a high rather than a low egg load. Neither fruit type nor egg load of introduced females had a significant effect on the occurrence of encounters between introduced and resident females, on the occurrence of contests, or on the propensity of introduced females to initiate boring in the presence of a resident female. Among introduced females, there was a consistent trend to initiate boring earlier in the presence than in the absence of a resident female. We discuss circumstances, likely to be narrow, under which boring propensity of an intruding female might be facilitated in the presence of a resident female.     

Exploring the foraging environment of a natural enemy of Callosobruchus maculatus: Spatial egg distribution in stored cowpea

Knowledge of the spatial distribution of stored product insects may reduce the dependency on chemicals for control of these insects. Biological control, for instance, could be improved based on such knowledge. In this paper we describe the three-dimensional spatial oviposition pattern of Callosobruchus maculatus in stored cowpea. Individual C. maculatus females oviposited in clusters of 70±15 (SD) eggs. These clusters were variable in shape. In any cluster 90 to 95% of the eggs fitted into a volume of 19.1±3.5 cm³. The egg density was highest (0.6 eggs bean^–1) at the center of a cluster and decreased towards the periphery. A statistically significant relationship existed between the number of eggs n in a cluster and the cluster volume, V(cm³): V=11.5+0.11n. We also investigated the spatial egg distribution of beetles which emerged from egg clusters such as those produced by individual females. Their oviposition was not confined to one specific area but was scattered throughout the bean mass. A point pattern analysis showed that the density of the `parent' cluster had no effect on the spatial egg pattern. These results give insight into the foraging environment which the egg parasitoid Uscana lariophaga, a promising candidate for biological control of C. maculatus, is facing. We argue that the probability p of encountering at least one other bean with eggs after a parasitization is a function of the number n of beans that are visited: p=1–0.42 (0.37)^(n–1).     

Habitat assortment of sexes and water balance in a dioecious grass

Summary Eggs of the carabid beetle Pterostichus oblongopunctatus kept under optimum conditions in petri dishes had a mean mortality of 8% (n=200), while the mortality of eggs incubated in fresh litter was 83% (n=250). Heating the litter up to 70° C and thus eliminating other organisms from the litter fauna reduced mortality to 18% (n=250). When eggs were deposited by the females themselves in a perspex cylinder containing an undisturbed soil sample, the mean egg mortality was 70% (n235), but when predators had been eliminated the mortality was only 7% (n235). The assumption of Grüm (1975) that egg mortality can be neglected does not hold for P. oblongopunctatus. The predators of the eggs must be common species, but it is not yet known to which family they belong.     

Recognition and avoidance of previously laid egg masses by the oblique-banded leafroller (Lepidoptera: Tortricidae)

Investigations were conducted on oviposition by laboratory-reared oblique-banded leafrollers, Choristoneura rosaceana Harris. In paperbag oviposition chambers, females deposited their egg masses in a significantly nonrandom, uniform manner in 16 of 20 replicates. Inpetri dish oviposition chambers, gravid females were exposed to egg masses recently deposited by other females. As determined by the magnitude of the mean direction vector of the second egg mass in relation to the first, females detected and avoided egg masses laid by other females. In addition, they avoided waxed paper from which a recently oviposited egg mass had been removed. This phenomenon is most likely due to a pheromone laid down with each egg mass.     

Egg Load and Body Size of Lab-cultured Cotesia marginiventris

The egg load of lab-cultured Cotesia marginiventris (Cresson) (Hymenoptera: Braconidae), a solitary koinobiont endoparasitoid of noctuid caterpillars, was determined in this study. Information on egg load may provide clues to more efficient in vivo rearing of C. marginiventris. I tested the hypothesis that egg load, defined as the number of mature oöcytes (i.e., fully chorionated eggs) found in adult females, was related to body size. Cotesia marginiventris females possessed two ovaries and two ovarioles per ovary; mature eggs were found in ovaries and oviducts. Newly-emerged females held an average of 149 mature eggs. Immature eggs were slightly visible in the distal portions of the ovarioles; they were not counted. Egg load was marginally related to body size (i.e., hind tibia length). The results of this study suggest that (1) body size can sometimes predict egg load or potential fecundity of lab-cultured C. marginiventris and (2) an efficient rearing system that exploits the potential fecundity of C. marginiventris might involve using young females and allowing them to oviposit in new hosts, each day, for up to a week.     

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.     

Larval host plant affects fitness consequences of egg size variation in the seed beetle Stator limbatus

Egg size variation often has large effects on the fitness of progeny in insects. However, many studies have been unable to detect an advantage of developing from large eggs, suggesting that egg size variation has implications for offspring performance only under adverse conditions, such as during larval competition, periods of starvation, desiccation, or when larvae feed on low-quality resources. We test this hypothesis by examining the consequences of egg size variation for survivorship and development of a seed-feeding insect, Stator limbatus, on both a low-quality (Cercidium floridum) and a high-quality (Acacia greggii) host plant. Our results are consistent with the hypothesis. S. limbatus larval performance was affected by egg size only when developing on the poor-quality host (C. floridum); larvae from large eggs survived better on C. floridum than those from small eggs, while there was no evidence of an effect of egg size on progeny development time, body weight, or survivorship when larvae developed on A. greggii. These results indicate intense selection for large eggs within C. floridum-associated populations, but not in A. greggii-associated populations, so that egg size is predicted to vary among populations associated with different hosts. Our results also support this hypothesis; females from a C. floridum-associated population (Scottsdale) laid larger eggs than females from an A. greggii-associated population (Black Canyon City).     

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.     

Parentage analyses of freshwater goby Rhinogobius sp. OR using microsatellite DNA markers and egg developmental stages of eggs in the nest

Parentage analyses of the paternally caring goby Rhinogobius sp. OR were performed using microsatellite DNA markers and examination of developmental stages of eggs collected from five nests with parental males in the wild. Four of five nests had egg masses with eggs at the same developmental stages from single females. In one nest, the egg mass with eggs at three different developmental stages originated from four females, and eggs of whole developmental stages were observed within the egg mass of each female. This observation suggests that in this goby embryonic stage is an inaccurate indicator of the number of mates.     

Proximate Factors Regulating Maternal Options in the Eggplant Lace Bug, Gargaphia solani (Heteroptera: Tingidae)

The eggplant lace bug, Gargaphia solani, was used to investigate the proximate factors regulating maternal care and a noncaring, condition dependent strategy called egg dumping. We hypothesize that the act of delaying oviposition while searching for a dumping opportunity suppresses oogenesis and triggers guarding behavior. We examined several predictions of this hypothesis by measuring: (1) whether females do delay oviposition in the absence of dumping stimuli, (2) whether females in transition between egg dumping and egg guarding are capable of expressing either reproductive option, (3) the effect of nymphal interactions and antennal ablation on the duration of maternal care, and (4) oogenesis in guarding and dumping females. We found that females without a dumping opportunity wait, on average, 30 h longer to oviposit than females exposed to a dump mass. Females that had initiated their own egg mass could resume eggdumping if they had laid less than half of their eggs but were unlikely to abandon their eggs when most had been laid. Maternal care in G. solani can be prolonged if interactions with nymphs are artificially prolonged. Females require antennae to maintain maternalcare. Presumably antennae transduce cues from eggs and nymphs. Dissections of dumping and guarding females 72 h after their first oviposition demonstrated that dumpers continue to produce primary oocytes after a dumping event but guarders terminate oogenesis whilecaring for their first brood. We interpret all of these results within the context of the hypothesis that juvenile hormone titers regulate the expression of both egg dumping and egg guarding.