Interspecific larval competition between two egg parasitoids in refrigerated host eggs of Riptortus pedestris (Hemiptera: Alydidae)

In a previous study, we found that soybean fields could be supplemented with refrigerated eggs of Riptortus pedestris (Fabricius) (Hemiptera: Alydidae) to enhance parasitism. As a part of a study to evaluate the effect of host egg refrigeration on parasitism, host acceptance behavior and interspecific larval competition between Gryon japonicum (Ashmead) (Hymenoptera: Scelionidae) and Ooencyrtus nezarae Ishii (Hymenoptera: Encyrtidae) were studied in multiparasitized unrefrigerated and refrigerated eggs. O. nezarae showed complete host acceptance behavior when offered refrigerated host eggs that were preparasitized by G. japonicum. Adult emergence rate of O. nezarae was 43 and 74% when the interval between the first and second oviposition was 0 and 4 days, respectively, and was not different between refrigerated and unrefrigerated eggs. Refrigeration did not change host acceptance behavior of G. japonicum, but adult emergence declined from 80% in unrefrigerated eggs to 37% in refrigerated eggs that were pre-parasitized by O. nezarae on the same day. No negative effects of refrigeration on sex ratio, adult longevity, and adult size of the both parasitoids were found. Generally host egg refrigeration did not negatively affect host acceptance behavior of the both parasitoids on preparasitized eggs or larval competition between the two parasitoids in multiparasitized host eggs with exceptions in the development time and emergence rate of G. japonicum. Therefore, host egg refrigeration may not interrupt interactions between the parasitoid populations in the field.     

Enhancement of Riptortus clavatus (Thunberg) (Hemiptera: Alydidae) egg parasitism by a component of the bug's aggregation pheromone

Egg parasitism of the bean bug, Riptortus clavatus (Thunberg) (Hemiptera: Alydidae), was surveyed using individual egg bags in a sweet persimmon orchard and at Gyeongsang National University (GNU) campus, Korea in 2006. The effect of (E)-2-hexenyl (Z)-3-hexenoate (E2HZ3H), one component of R. clavatus aggregation pheromone, on the parasitism enhancement was tested at GNU campus and in a soybean field in 2005 and 2006, respectively. In 2006, two egg parasitoid species, Ooencyrtus nezarae Ishii (Hymenoptera: Encyrtidae) and Gryon japonicum (Ashmead) (Hymenoptera: Scelionidae), emerged from R. clavatus eggs. Parasitism by O. nezarae was 9.4–48.3% in mid August to mid September in GNU campus and 6.7–30.0% at the orchard. Total parasitism by G. japonicum (2.5%) at both sites throughout the experimental period was lower than that by O. nezarae (12.5%). This survey revealed nearly no activity of the two species after October at both sites. G. japonicum was solitary and O. nezarae could be either solitary or gregarious. From a single R. clavatus egg, one female or one male G. japonicum emerged. However, an average of 4.3 O. nezarae emerged from one host egg. It took 12.6 d for G. japonicum and 12.0 d for O. nezarae to emerge from R. clavatus eggs in the laboratory. Treatment with E2HZ3H increased parasitism by O. nezarae in both years, but did not increase parasitism by G. japonicum. This suggests that E2HZ3H can be used as a kairomone to reduce the density of R. clavatus in the fields where natural parasitism by O. nezarae is high.     

Evaluation of cold-stored eggs of Dolycoris baccarum (Hemiptera: Pentatomidae) for parasitization by Trissolcus nigripedius (Hymenoptera: Scelionidae)

Trissolcus nigripedius Nakagawa (Hymenoptera: Scelionidae) is an egg parasitoid of Dolycoris baccarum L. (Hemiptera: Pentatomidae), a polyphagous insect pest of many crops including soybean. As a method for mass-rearing to augment the parasitoid, cold storage of host eggs were evaluated in the laboratory. After 0 (control), 8, 20, 60, 90, and 120 days of refrigeration, host eggs were given to adult female T. nigripedius. Host acceptance behaviors of the parasitoid, categorized as drumming, oviposition, and marking, on the refrigerated eggs and biological attributes of offspring were assessed. Most of the attributes examined were affected by the refrigeration of host eggs. But host eggs could be refrigerated for up to two months without significant change in emergence rate and sex ratio and with 90% of parasitism. In addition, the second generation of the parasitoid was not negatively affected at all. However, frozen eggs of D. baccarum can not be used for rearing the parasitoid since parasitism rate decreased to 44% on host eggs frozen for 8 days. Refrigeration of D. baccarum eggs could be useful for mass-rearing and augmentation of T. nigripedius to control D. baccarum without reduction in the quality of parasitoid’s progeny. Furthermore, refrigerated eggs could be supplemented in the field to boost the population of T. nigripedius since immature D. baccarum can not hatch after 20 days of refrigeration.     

‘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.     

Biology and field performance of gryon clavigrallae (Hymenoptera: Scelionidae), an egg parasitoid of Clavigralla spp. (Hemiptera: Coreidae) in India

The biology and impact of Gryon clavigrallae Mineo, an egg parasitoid of Clavigralla scutellaris Spinola and C. gibbosa (Westwood), was investigated. The calculated developmental threshold temperatures for females and males were 15.6 degrees C and 15.8 degrees C, respectively. Emergence exceeded 94% at temperatures between 22 and 30 degrees C. Adult females lived on average 28-96 days when fed with honey. Without food, adults lived < 6 days. Mean fecundity was 56.4 eggs per female. A significant trend of lower fecundity after longer periods of host deprivation was observed. Gryon clavigrallae females successfully oviposited in host eggs of all ages though eggs < 4 days old were preferred. Total host handling times were significantly longer on C. gibbosa eggs (23.5 min) than on C. scutellaris eggs (12.0 min). Females readily distinguished parasitized from non-parasitized host eggs. Superparasitism was observed when few or no unparasitized eggs were available. Eggs of the two Clavigralla species can be separated by surface structure and condition after eclosion. Clavigralla scutellaris laid significantly larger egg clusters than C. gibbosa (19.9 versus 10.5 eggs per cluster). Gryon clavigrallae was present as soon as the first Clavigralla spp. egg clusters were found on pigeonpea. The percentage of egg clusters parasitized increased early in the season with egg cluster density and remained high (up to 83%) despite fluctuations in host density. Overall, G. clavigrallae parasitized 40 and 58% of C. gibbosa and C. scutellaris eggs. The percentage of egg clusters parasitized and the number of eggs parasitized per cluster increased significantly with egg cluster size. The overall parasitoid sex ratio was highly female biased but varied with the number of eggs parasitized per cluster.     

THE PREFERENCE AND DENSITY RESPONSES OF EGG PARASITOID,OOENCYRTUS NEZARAE ISHO,TO THE EGGS OF SOYBEAN STICK BUGS,RIPTORTUS CLAVATUS THUNBERG AND PIEZODORUS HYBNERI GMELIN*

Abstract The preference and density response test9 of egg parasitoid, Ooencyrtus nezarae Ishii, to the eggs of soybean stick bugs, Riptortus clavatus Thunberg and Piezodorus hybneri Gmelin, were carried out in this study. In the free choice test, O. nezarae emerged from both eggs of stick bug prefer R. clavatus eggs only in the various ratios of host egg densities. In non‐choice test, O. nezarae emerged from P. hybneri eggs parasitize both R. clavatus and P. hybneri eggs, showing a high parasitism on host eggs. However, O. nezarae reared on R. clavatus eggs had a significantly lower parasitism on the eggs of P. hybneri. Olfactometer test showed that the odor emitted from R. clavatus eggs had a great influence on the host searching and parasitizing behavior of O. nezarae. The number of R. clavatus and P. hybneri eggs parasitized by O. nezarae gradually increased with the increase of host egg densities. However, the parasitism of O. nezarae to R. clavatus eggs decreased with the increase of host density. The parasitism of O. nezarae are significantly lower (P < 0.05) on host egg P. hybneri under the laboratory condition, especially for O. nezarae emerged from R. clavatus eggs. This research results implied that in the soybean field, there were some optimal or spatial barriers for O. nezarae to find R. clavatus eggs efficiently.     

Effects of egg age of Spartocera dentiventris (Berg) (Hemiptera: Coreidae) on parasitism by Gryon gallardoi (Brethes) (Hymenoptera: Scelionidae)

To evaluate the effect of egg age of Spartocera dentiventris (Berg) on the parasitism by Gryon gallardoi (Brethes), groups of 12 eggs of 2, 3, 4, 5, 6, 7, 8, and 12 days old were exposed to one female of G. gallardoi on a test tube, with the wasp observed for 2h subsequently. Later, the eggs were individualised and incubated (25 +/- 1 degrees C; 12h photoperiod) until adult emergence. To identify the preferred age, groups of 12 eggs of 1, 3, 5, 7 and 12 days of age were simultaneously exposed to a parasitoid female on an arena for 2h. These groups were also individualised and incubated until parasitism was confirmed. On all ages tested parasitism was at about 88.5 +/- 2.64%, with a reduction in adult parasitoid emergence with age increasing (P < 0.01). The development time increased with increasing egg age (P < 0.01), with males developing faster. Parasitoid sex ratio for all ages tested was female-biased (0.49:1), with the proportion of males invariant with host age (P > 0.05). When the egg groups were exposed simultaneously, the female preferred eggs one and three days old. The results indicate a preference of G gallardoi for younger eggs of S. dentiventris, with host egg age exerting a negative effect on development time and offspring emergence of the parasitoid.     

Interactions between the winter cherry bug <Emphasis Type="Italic">Acanthocoris sordidus</Emphasis> (Hemiptera: Coreidae) and its egg parasitic wasps

Detailed research on the relative strength of the influence of parasitoids on hosts has contributed to the development of biological control techniques. Gryon philippinense (Ashmead) is a major egg parasitoid of the winter cherry bug, Acanthocoris sordidus (Thunberg). However, available data are insufficient to determine which egg parasitoids have the strongest effect on mortality of A. sordidus or why G. philippinense is the major egg parasitoid of A. sordidus. In this study, G. philippinense attained a much higher level of parasitism on A. sordidus eggs compared to four other parasitoid species. Mortality induced by G. philippinense on A. sordidus egg clusters occurred in an all-or-none manner, whereas mortality caused by the four other parasitoids occurred in a random manner. Compared to Ooencyrtus spp., G. philippinense required a significantly shorter time to parasitize an A. sordidus egg and also parasitized significantly more A. sordidus eggs within 24 h. These findings demonstrate that a short handling time allows G. philippinense to be the most significant egg parasitoid of A. sordidus.     

Effects of storage at low-temperature of various heteropteran host eggs on the egg parasitoid, <Emphasis Type="Italic">Trissolcus semistriatus</Emphasis>

Various heteropteran host eggs stored under two low-temperatures were tested in the laboratory for their usability in the production of sunn pest egg parasitoid, Trissolcus semistriatus. Parasitism, adult emergence rate and development time were assesed on stored and fresh eggs of Eurygaster integriceps, Dolycoris baccarum, Graphosoma lineatum and Eurydema ornatum. Masses of fresh host eggs in microcentrifuge tubes were maintained at +6 °C and −20 °C. Every 30 days, 50 eggs of host species were exposed to parasitism by T. semistriatus. The host eggs stored at 6 °C remained viable to parasitism by T. semistriatus up to 2 months, while those of stored at −20 °C were parasitized at high rates up to 4 months, alhough succesful parasitism rates decrease with time. However, it was indicated that both fresh or stored E. ornatum eggs were not preferred to parasitism by the parasitoid. A longer development time from egg to adult was observed in stored eggs under two tested storage techniques when compared with fresh eggs.     

Factors affecting sequential sex allocation in the parasitoid wasp Gryon japonicum

Female wasps of the solitary egg parasitoid Gryon japonicum (Hymenoptera: Platygastridae) allocate male and female offspring in a particular sequence to successive hosts. Male eggs are typically laid in the second host, and the sex allocation sequence is reset after a certain period of time. The present study aimed to examine the underlying mechanism to hold information and reset the sequence by using eggs of Riptortus pedestris (Heteroptera: Alydidae) as hosts. After completion of initial oviposition, a female wasp was treated by cold anesthesia for 1 h, exposure to a parasitized host for 3 h, or being kept at 15°C in darkness for 24 h, and then presented with three host eggs. Cold‐anesthetized females did not reset the sex allocation sequence, indicating that cold anesthesia did not block the mechanism of holding information about oviposition order. Frequent encounters with parasitized hosts were also insufficient to reset the sequence. However, being kept in cool, dark conditions significantly affected resetting, suggesting that low temperature lengthened the time required to reset the sequence. This implies that it is probable that the mechanism to hold information and reset sex allocation sequence in G. japonicum involves metabolism.     

Ecotoxicological effect of insecticides on Ooencyrtus nezarae (Hymenoptera: Encyrtidae) reared from refrigerated and unrefrigerated Riptortus pedestris (Hemiptera: Alydidae) host

Due to increased field occurrence of Riptortus pedestris (Fabricius) (Hemiptera: Alydidae) on various crops including soybean, persimmon and apple in recent years in Korea, demand for insecticide applications to control the stink bug has increased. Acute toxicity of eight pesticides on Ooencyrtus nezarae Ishii (Hymenoptera: Encyrtidae), a major egg parasitoid of R. pedestris, was compared in the laboratory. Fenitrothion, spinosad, cyfluthrin, etofenprox and carbosulfan caused 100% mortality of O. nezarae within 24 hours by topical application or exposure to residue. Fenitrothion was also highly toxic to the parasitoid when ingested orally. In a previous study, release of refrigerated inviable eggs of R. pedestris was found to increase field parasitism; therefore, we evaluated the sublethal effect of fenitrothion when O. nezarae parasitised refrigerated or unrefrigerated host eggs. Although parasitism rates on both kinds of eggs significantly decreased when O. nezarae were provided with host eggs sprayed with fenitrothion, no difference in parasitism rate, adult emergence, sex ratio, development time and longevity of O. nezarae was found between the refrigerated or unrefrigerated host eggs when the insecticide was treated either before or after oviposition. There was no significant sublethal effect when parasitised host eggs were treated with the insecticide. From these results, all the insecticides tested showed high, acute toxicity against O. nezarae with relatively lower sublethal effects. Refrigeration of host eggs did not affect the susceptibility of O. nezarae to insecticides.     

Susceptibility of Nezara viridula (L.) (Hemiptera: Pentatomidae) Egg Masses of Different Sizes to Parasitism by Trissolcus basalis (Woll.) (Hymenoptera: Platygastridae) in the Field

Egg masses of Nezara viridula (L.) are commonly parasitized by Trissolcus basalis (Woll.), and we investigated the role of size of egg masses on parasitization by T. basalis. Sentinel egg masses were exposed to parasitism in the field for 6–7 days, when they were collected for evaluation of parasitoid emergence. We recorded the number of eggs per egg mass, the number of emerged hosts, and the number of empty and parasitized eggs. We calculated the proportion of attacked host egg masses (DE), the proportion of parasitized eggs per attacked egg mass (PE), and total parasitism (PI). The total number of egg masses exposed to parasitism was 330. The minimum, mean, and maximum egg mass sizes were 25, 75.2, and 111, respectively. DE and PE varied widely between different fields, and they were independent of egg mass size. In 14.2% of all parasitized egg masses, we found simultaneous emergence of T. basalis and N. viridula independently of host egg mass size. PE exhibited low variability compared with PI and DE, which were linearly related. PI and DE values from other field studies are consistent with the linear relationship, suggesting that PI is mostly related to the proportion of the DE. This also suggests that total parasitism is independent of egg mass size, of possible differences in plant species, and T. basalis density and strains.     

Parasitization by Edovum puttleri (Hymenoptera: Eulophidae) in relation to host density in the field

Abstract.

• 1 The relationship between parasitization by Edovum puttleri Grissell and density of eggs of the Colorado potato beetle, Leptinotarsa decemlineata Say (Coleoptera: Chrysomelidae), was studied on two spatial scales (eggs mass and 6 m² cage).
• 2 For both scales, rates of parasitism were generally inversely related to host density for periods ranging from 2 to 8 days after parasitoid release. Thereafter, parasitism became independent of host density.
• 3 The initial inverse-density relationship and subsequent shift to density independence may result from several factors: (1) ambient temperatures, (2) the parasitoid's limited egg production, (3) differential times of exposure of egg masses to parasitoids, and/or (4) the parasitoid's patterns of host feeding and oviposition.
• 4 Although overall levels of parasitism were relatively low, total mortality of L.decemlineata eggs (including nonviable and cannibalized eggs, and those killed by parasitoid feeding) in parasitized egg masses was consistently high (?70–90%).

     

Reciprocal influences and costs of parasitism on the development of Corcyra cephalonica and its endoparasitoid Venturia canescens

Many endoparasitoids develop successfully within a range of host instars. Parasitoid survival is highest when parasitism is initiated in earlier host instars, due to age-related changes in internal (physiological) host defences. Most studies examining fitness-related costs associated with differences in host instar have concentrated on the parasitoid, ignoring the effects of parasitism on the development of surviving hosts that have encapsulated parasitoid eggs. A laboratory experiment was undertaken examining fitness-related costs associated with encapsulation of Venturia canescens (Hymenoptera: Ichneumonidae) eggs by fifth (L5) instar larvae of Corcyra cephalonica (Lepidoptera: Pyralidae). Growth and development of both host and parasitoid were monitored in C. cephalonica larvae containing 0, 1, 2, or 4 parasitoid eggs. Adult size and fecundity of C. cephalonica did not vary with the number of eggs per host. However, there was a distinct increase in host mortality with egg number, although most parasitoids emerged from hosts containing a single egg. The most dramatic effect on the host was a highly significant increase in development time from parasitism to adult eclosion, with hosts containing 4 parasitoid eggs taking over 2.5 days longer to complete development than unparasitized larvae. The egg-to-adult development time and size of adult V. canescens did not vary with egg number per host, as demonstrated in a previous experiment using a different host (Plodia interpunctella). The results described here show that there are fitness-related costs to the host associated with resistance to parasitism.     

Biological attributes of Ooencyrtus nezarae Ishii (Hymenoptera: Encyrtidae) reared on refrigerated eggs of Riptortus pedestris (= clavatus) Fabricius (Hemiptera: Alydidae)

Storing host eggs at low temperatures has been used to mass rear parasitoids of stink bugs, including Riptortus pedestris Fabricius (Hemiptera: Alydidae), a major soybean pest in Korea and Japan. However, no information on the effect of cold storage on parasitization by Ooencyrtus nezarae Ishii (Hymenoptera: Encyrtidae), one of the major parasitoids of R. pedestris, has been published. In this study, we examined biological attributes, including parasitism rate, development time, sex ratio, adult size, and longevity, of O. nezarae when the adult parasitoids were provided with host eggs refrigerated for 0, 15, 30, 60, and 120 days at 2.0 °C. None of the attributes of the first or second generation of O. nezarae was negatively affected by host egg refrigeration up to 30 days. In addition, O. nezarae could parasitize refrigerated host eggs successfully for the first four days of post-refrigeration period when they were kept at 26.3 °C and 78.7% RH conditions. Therefore, refrigeration of R. pedestris eggs can be a good method to mass rear O. nezarae.     

Demonstration of interspecific competition between two sympatric egg parasitoids of Riptortus pedestris (Fabricius) in laboratory condition

Ooencyrtus nezarae Ishii (Hymenoptera: Encyrtidae) acts as a facultative hyperparasitoid of Gryon japonicum (Ashmead) (Hymenoptera: Platygastridae) sympatric parasitoid of Riptortus pedestris (Fabricius) (Hemiptera: Alydidae). A longer period of host egg exploitation by both parasitoid species would be beneficial for O. nezarae, while G. japonicum tends to be successful when the parasitoids have only a short exploitation period. We demonstrated the interspecific competition by measuring parasitism in nine combinations of host densities (10, 20, and 40 eggs) and exploitation times (1, 3, and 5 days). To reflect the gregarious-solitary dichotomy of the two species, three O. nezarae and one G. japonicum mated females were compared in addition to a one-to-one competition design. We found that O. nezarae was the better competitor when exploitation time was longer than 1 day, irrespective of host density. Total parasitism rate and progeny emergence of O. nezarae were 1.6–2.8 and 4.7–7.3 times higher than for G. japonicum in three-to-one competition design, respectively. Although G. japonicum females were more effective in host finding (as shown by their higher per capita rate of parasitism when exploitation time was short), their progeny suffered high mortality from the larval interspecific competition inside multiparasitized host eggs. These results suggest that gregarious O. nezarae is the superior competitor when host eggs are available for longer period of time while solitary G. japonicum is superior when host resources are available for only a limited time.     

Teratocytes of the solitary endoparasitoid Meteorus gyrator (Hymenoptera: Braconidae): morphology, numbers and possible functions

Abstract. Laboratory studies investigated the development of teratocytes derived from the eggs of the parasitoid Meteous gyrator (Thun.) in its host, the tomato moth Lacanobia oleracea (L.). At hatching, each parasitoid egg produced an average of approximately 1000 teratocytes, but this number declined to approximately 400 during the course of parasitism. The teratocytes increased in size markedly, such that 7 days after egg hatch their mean diameter was approximately four times that of the cells immediately after dissociation. The haemolymph of parasitized hosts had reduced phenoloxidase activity, and teratocytes inhibited phenoloxidase activity when coincubated with plasma from nonparasitized hosts. The injection of teratocytes into nonparasitized fifth‐instar L. oleracea larvae suppressed growth and induced a supernumerary moult in some larvae. A number of parasitism‐specific proteins were detected in the haemolymph of parasitized hosts, and incubation of teratocytes in culture media indicated that these cells were a source of at least two of these proteins.     

Host preference: parasitism, emergence and development of Trissolcus semistriatus (Hym., Scelonidae) in various host eggs

Abstract: Trissolcus semistriatus is an important egg parasitoid of the sunn bug, Eurygaster integriceps. Conservation and the supportive augmentation of populations of egg parasitoids appears to offer high potential for the biological control of sunn bug in an integrated pest management system. The present study was conducted to determine the host preference of T. semistriatus on various heteropteran host eggs. When each host species was examined separately, the eggs of E. integriceps, Dolycoris baccarum, Graphosoma lineatum, Carpocoris pudicus and Holcostethus vernalis, were parasitized at high levels, namely 88.0, 83.6, 94.8, 87.3 and 80.8%, respectively, whereas a smaller portion of Eurydema ornatum eggs were parasitized (24.0%). There was no parasitism in Nezara viridula eggs by T. semistriatus. Similar high rates were recorded for the percentage of adult emergence in each host species, with the exception of E. ornatum. The average development time was shorter for males than for females. The average development period for both male and female was significantly longer in eggs of E. ornatum than in those of the other host species. These results and observations recorded during the experiments showed that G. lineatum and D. baccarum were available, cheap hosts and it was demonstrated that these hosts could be used for the mass production of egg parasitoids.     

Comparative assessments of Gonatocerus ashmeadi and the ‘new association’ parasitoid Gonatocerus tuberculifemur (Hymenoptera: Mymaridae) as biological control agents of Homalodisca vitripennis (Hemiptera: Cicadellidae)

Egg age preference, competitive ability, and behavior of Gonatocerus tuberculifemur (‘new association’ parasitoid) and Gonatocerus ashmeadi (‘old association’ parasitoid) were investigated in the laboratory to determine if one species exhibited competitive superiority. When searching concurrently for Homalodisca vitripennis egg masses, G. ashmeadi consistently outperformed G. tuberculifemur by parasitizing 25–53% more eggs under three different experimental systems in the laboratory with varying host densities, egg ages, and exposure times. G. ashmeadi parasitism in control vials containing one parasitoid ranged from 81–97% across all egg ages. G. tuberculifemur in control vials parasitized 60–66% of eggs 1 and 3 days old, and just 18% of eggs 5 days old. G. ashmeadi produced 5–16% more female offspring than G. tuberculifemur for all experimental conditions. In comparison to G. ashmeadi, G. tuberculifemur was observed off leaves with host eggs 20% more frequently and it oviposited 15% less frequently. G. ashmeadi and G. tuberculifemur when confined together allocated ∼1% of behaviors to antennating or aggressively chasing competitors off egg masses, and up to 2% of behaviors to antennating host egg masses and/or ovipositing into eggs from the opposite side of the leaf. These latter behaviors did not occur when parasitoids were confined alone with host eggs.     

Density dependence of Anaphes sordidatus (Hymenoptera: Mymaridae) parasitism on eggs of Listronotus oregonensis (Coleoptera: Curculionidae)

Listronotus oregonensis (LeConte) (Coleoptera: Curculionidae) oviposits in carrot leaves and the larvae feed in carrot roots. Its eggs are parasitized by Anaphes sordidatus (Girault) (Hymenoptera: Mymaridae) throughout its oviposition period. This parasitoid is the major biotic mortality factor for L. oregonensis. Parasitism by A. sordidatus was studied for three years in sequentially sown plots of carrots, Daucus carota var. sativa. Over the scason, significantly fewer L. oregonensis eggs were oviposited in later sown carrots than in earlier sown carrots because oviposition started later in late sown plots of carrots. A positive temporal density-dependent relationship was observed each year between parasitism rates and host densities. This positive density dependence occurred in early and mid-summer for earlysown carrots where host density reached 1–2 host eggs per plant but disappeared in late summer when host density decreased while parasitism remained high. Latesown carrots had low host egg density (0.2 host egg per plant) and contributed little to the total number of eggs. In these late sown plots, parasitism increased rapidly to over 80% but no density dependence was observed. Spatially, few statistically significant regressions were found but all indicated a positive spatial density-dependence. Most non-significant regressions occurred because the range of egg density was too small between plots for a given date.