Immune response of the hibiscus mealybug, Maconellicoccus hirsutus Green (Homoptera: Pseudococcidae), to oviposition of the parasitoid Anagyrus kamali Moursi (Hymenoptera: Encyrtidae)

Anagyrus kamali Moursi has been recently introduced into the Caribbean as a biological agent against the hibiscus mealybug, Maconellicoccus hirsutus Green. This host has a cellular defense reaction that involves encapsulation and melanization of the endoparasitoid egg. The impact of this immune response on the parasitoid progeny was assessed, as well as the response of the parasitoid countermeasures to overcome it. Under laboratory conditions, significant differences in the immune response were found for different developmental stages of M. hirsutus. The intensity of the immune response varied between second instar, third instar and adult mealybugs. After 30 h, the level of encapsulation was the highest for eggs oviposited in adults: 58% of eggs were encapsulated, followed by third (32%) and second (4%) instars. Three days after oviposition 23, 44 and 86% of the parasitoid eggs oviposited, respectively, in adult, third and second instars were not encapsulated. The unencapsulated parasitoid eggs could hatch and continue their development. Adult mealybugs required 30 h to encapsulate 50% of the eggs, whereas in second and third instars, 50% level encapsulation was never reached. Superparasitism had a saturating effect on the immune system; reduced levels of encapsulation occurred when more than 10 eggs were oviposited in a single mealybug. Wasp larvae were never encapsulated by M. hirsutus.     

BIONOMICS OF MICROPLITIS MEDIATOR (HYM: BRACONIDAE) PARASITIZING PSEUDALETIA SEPARATA (LEP: NOCTUIDAE)*

Abstract This paper deals with the bionomics of Microplitis mediator parasitizing Pseudaletia separata. There were some stings (ca. 3. 1%) in which female wasps oviposited more than one eggs and about 4. 6% stings that there was no egg laid into a host. A negative correlation existed between the host age and percentage of parasitization. When the host suffered more stings by the parasitoid wasps, the percentage of parasitization was higher. But there was no correlation between the parasitoid sex ratio and the host age or the frequency of stings by the parasitoids. The host age is not the major factor affecting the offspring sex ratio of the parasitoid. Some putative sex determinant factors in the parasitoid are discussed and the 2nd instar of the host is recommended as a suitable host age in which parasitization could be performed for the purpose of the parasitoid mass culture.     

Host instar susceptibility and selection and interspecific competition of three introduced parasitoids of the mealybug Paracoccus marginatus (Hemiptera: Pseudococcidae)

Three previously introduced parasitoids (Acerophagus papayae Noyes and Schauff, Anagyrus loecki Noyes and Menezes, and Pseudleptomastix mexicana Noyes and Schauff [Hymenoptera: Encyrtidae]) of the mealybug Paracoccus marginatus Williams and Granara de Willink (Hemiptera: Pseudococcidae) were studied for their host instar susceptibility and sex ratio, host instar selection, and interspecific competition in the laboratory. All three parasitoids were able to develop in the second instars, third-instar females, and adult females of P. marginatus. No progeny emerged from first-instar mealybugs. The proportion of female emergence was increased with increasing host size. Parasitoids selected their host instars for oviposition when they had a choice. Between second- and third-instar hosts, A. papayae and P. mexicana had significantly higher parasitism in second-instar mealybugs, whereas A. loecki had higher parasitism in the third-instar mealybugs. When competed with either one or two parasitoid species, A. papayae was significantly more successful in second-instar hosts and A. loecki was significantly more successful in third-instar mealybugs. P. mexicana was significantly less competitive when with A. papayae in both second and third instars, with A. loecki in third instars and with both A. papayae and A. loecki in second and third instars. Overall, A. papayae provided a better control of the host, when present singly or with the other two parasitoids. This information is important in evaluating the efficiency of A. papayae, A. loecki, and P. mexicana and understanding the outcome of their recovery and establishment in field studies conducted in Florida.     

The effect of host stage and temperature on selected developmental parameters of the solitary endoparasitoid Meteorus gyrator (Thun.) (Hym., Braconidae)

Abstract:  The development of the solitary endoparasitoid Meteorus gyrator was compared in the six larval stages of its host, the tomato moth Lacanobia oleracea , and at five constant temperatures. The host instar at the time of parasitism had a marked effect on the larval developmental period of the parasitoid, such that larvae derived from eggs oviposited in first instar hosts took approximately 18 days to egress, whilst those derived from eggs oviposited in sixth instar hosts took <10 days. The weight of cocoons was greatest when oviposition was into final instar hosts, where female cocoons averaged 12.8 mg, and lowest in those derived from eggs oviposited into first instars (9.2 mg). The parasitoid's larval development rate in third instar hosts increased with temperature increments in a linear fashion up to 25°C , after which development times were only marginally increased. At 10°C, the mean larval development time was approximately 90 days and pupal development 35–40 days, whilst at 25°C development times were 10–11 days for larvae and 6–7 days for the pupae. In the majority of cases, overall development times were marginally longer (<1 day) in females than in males.     

A biological assessment of Apophua simplicipes (Hymenoptera: Ichneumonidae) as a parasitoid of the oblique banded leafroller, Choristoneura rosaceana (Lepidoptera: Tortricidae)

Apophua simplicipes (Cresson) (Hymenoptera: Ichneumonidae) is a common parasitoid of the oblique banded leafroller, Choristoneura rosaceana (Harris) (Lepidoptera: Tortricidae) in organically managed apple orchards in the southern interior of British Columbia, Canada. The biological characteristics of a laboratory colony of A. simplicipes were studied. When held at 15 and 25°C and provided with honey water, individual females survived an average of 60.6±6.1 and 29.8±4.7 days, oviposited 196.7±50.7 and 326.6±51.3 eggs and parasitized a total of 163.4±40.4 and 229.4±35.8 hosts, respectively. Females oviposited into first through fourth instar oblique banded leafrollers, with significantly more parasitism occurring in the first two instars compared to the third and fourth instars. No parasitoid larvae survived past the first larval stage in parasitized fourth instar hosts. Apophua simplicipes did not parasitize larvae of three-lined leafroller, Pandemis limitata (Robinson) (Lepidoptera: Tortricidae) which is sympatric with oblique banded leafrollers in orchards in the southern interior of British Columbia. Female predation and host feeding from wounds on early instars of both leafroller species was observed under laboratory conditions. In addition, early instar hosts exited diet feeding sites in response to the probing activity of the ovipositing wasps. A similar escape reaction in the orchard may cause a leafroller larva to move away from its feeding site, making it more vulnerable to predation or movement off the tree. Apophua simplicipes larvae emerged from fifth and sixth instar hosts. Parasitized oblique banded leafroller hosts consumed significantly less meridic diet than unparasitized female larvae from fifth instar through to parasitoid emergence or leafroller pupation. Our laboratory results suggest that A. simplicipes may reduce field populations of oblique banded leafroller and decrease pest feeding damage.     

Multiparasitism by the pupal parasitoids, Xanthopimpla stemmator (Hymenoptera: Ichneumonidae) and Pediobius furvus (Hymenoptera: Eulophidae) on two African cereal stemborers, Chilo partellus (Lepidoptera: Crambidae) and Busseola fusca (Lepidoptera: Noctuidae)

The outcome of multiparasitism by two pupal parasitoids, the exotic solitary ichneumonid Xanthopimpla stemmator and the indigenous gregarious eulophid Pediobius furvus, was studied using the invasive crambid stem borer Chilo partellus and the indigenous noctuid Busseola fusca as hosts. Two parasitism sequences were observed, where X. stemmator oviposited before P. furvus, and vice versa. In addition, the effect of time between first and second parasitism on parasitoid emergence, development, sex ratio, and number of offspring was assessed. For most treatments, time interval between parasitism had no significant effect on the percentage if pupae producing either P. furvus or X. stemmator. In general, X. stemmator outcompeted P. furvus irrespective of the order of oviposition, time interval between ovipositions and host species. Further studies, especially on host finding capacity of the two parasitoid species are required to determine the competitiveness of the two species under field conditions.     

Intra- and interspecific interactions between Lysiphlebus testaceipes and Lipolexis scutellaris (Hymenoptera: Aphidiidae) reared on Toxoptera citricida (Homoptera: Aphididae)

Classical biological control of the brown citrus aphid Toxoptera citricida Kirkaldy in Florida has involved the release of Lipolexis scutellaris Mackauer. Lysiphlebus testaceipes Cresson is already present in Florida and also parasitizes T. citricida. Because parasitoid-parasitoid interactions may affect the establishment of a newly introduced parasitoid species, intra- and interspecific larval interactions of both parasitoids were studied in the laboratory using T. citricida reared on citrus. Five time intervals were allowed between successive oviposition opportunities. Early developmental times were determined for both parasitoids: eggs of L. testaceipes and L. scutellaris hatched after 54.3 and 61.7 h, while molt to second instar occurred after 73.3 and 87.1 h, respectively, after oviposition at 22 degrees C. At intervals <12 h, both parasitoids had a greater tendency to multiparasitize than to superparasitize, and the tendency to superparasitize or multiparasitize decreased with an increase in time between successive oviposition opportunities. Of the 10 interspecific interactions studied, 5 produced a winning wasp species, 3 of which could be explained by the hypothesis of physical conflict. A combination of development time, age of competing larvae, and oviposition sequence were responsible for the outcomes observed. Neither parasitoid proved to be intrinsically superior when interspecific competition occurs in second- and third-instar T. citricida, indicating there is no reason to suggest that they cannot coexist in Florida citrus groves.     

Multiple Parasitism by Cotesia sesamiae and Cotesia flavipes (Hymenoptera: Braconidae) on Busseola fusca (Lepidoptera: Noctuidae)

Busseola fusca (Fuller) is one of the most important pest of cereals in sub-Saharan Africa. Cotesia sesamiae (Cameron) is the predominant parasitoid attacking B. fusca larvae in many parts of Africa. An exotic parasitoid, Cotesia flavipes Cameron, was introduced into Kenya in 1993 for the control of Chilo partellus (Swinhoe). Laboratory studies indicated that although C. flavipes would search for, and attack B. fusca , it was not able to complete its development in this host. The aim of the present study was to investigate the outcome of multiple parasitism of B. fusca by the two Cotesia species. The study showed that when both parasitoid species stung a B. fusca larva at the same time, both parasitoids emerged from more than half of the host larvae, C. flavipes alone emerged from 17%, and C. sesamiae alone emerged from 9%. When the larvae were parasitized by C. sesamiae first, and then 2 h later by C. flavipes , and vice versa, most of the progeny were C. flavipes . However, when B. fusca larvae were stung by C. sesamiae three days before oviposition by C. flavipes , significantly more C. sesamiae emerged from the larvae. When C. flavipes oviposited first, no larvae produced C. flavipes only. The interaction of parasitoids and the host immune system, and the implications of these results for the biological control of stem borers in East Africa are discussed.     

A new species of Rhynchoidomonas Patton, 1910 (Kinetoplastida: Trypanosomatina) from Operophtera brumata (Lepidoptera: Geometridae)

Summary A new species of Rhynchoidomonas Patton was observed in a single adult male winter moth, Operophtera brumata (L.) from England. Intracellular amastigotes, and extracellular epimastigotes and trypomastigotes with an undulating membrane and free flagellum, were present. All stages had a large, reniform kinetoplast. As transmission of the flagellate between generations of winter moths by ingestion of infected faeces is a virtual impossibility, it is suggested that the flagellate's true host may have been a dipteran parasitoid and that an egg, surface-contaminated with the flagellate, was oviposited into or ingested by a winter moth larva. If the parasitoid had died, this flagellate infection could have been carried over to the adult moth. ac]19830601     

The parasitoidOoencyrtus nezarae (Hymenoptera: Encyrtidae) prefers hosts parasitized by conspecifics over unparasitized hosts

Summary Two laboratory experiments were conducted to examine the ovipositional preferences of the egg parasitoidOoencyrtus nezarae Ishii (Hymenoptera: Encyrtidae) for parasitized and unparasitizedMegacopta punctatissimum Montandon (Hemiptera: Plataspidae). Females that had never oviposited or that had not oviposite for 3 days preferred recently parasitized hosts more than unparasitized hosts. The preference for recently parasitized hosts appeared to be mediated by the punctures in already parasitized hosts made by the ovipositor of the first female. Survival of the parasitoid progeny was lower in recently parasitized hosts than in unparasitized hosts. However, handling time of parasitized hosts was extremely short relative to that of unparasitized hosts, because the superparasitizing female could use the punctures made by the previous females. It is concluded that the females preferred the parasitized hosts over unparasitized hosts because the benefit of saving time and energy for drilling was more than the cost of progeny survival.     

Interactions between the ParasitoidAmetadoria misella(Diptera: Tachinidae) and the Granulovirus ofHarrisina brillians(Lepidoptera: Zygaenidae)

Interactions between a granulovirus (HbGV), a tachinid parasitoidAmetadoria misella,and their host, the western grapeleaf skeletonizerHarrisina brillians,were investigated. In field populations, the occurrence ofA. misellain HbGV-infectedH. brillianspupae was less frequent than would have been expected by random assortment of the virus and the parasitoid. Furthermore, enzyme-linked immunosorbent assay detected granulovirus less frequently and in lower concentrations in parasitized pupae than in nonparasitized pupae. Finally, in the host pupae that tested positive for virus, parasitoids were more likely to survive pupation than hosts. When laboratory-rearedH. brillianslarvae were exposed to naturally occurringA. misellain a field experiment, the parasitoid oviposited more often in older than in younger host larvae and more often in healthy than in HbGV-infected host larvae. These results are consistent with the hypothesis that selective oviposition byA. misellaleads to reduced overlap of the parasitoid and HbGV in hosts, resulting in greater parasitoid survival.     

Biosteres longicaudatus: Developmental dependence on host (Anastrepha suspensa) physiology

Larvae of Anastrepha suspensa that were in the first day of the third instar were parasitized by females of the solitary endoparasitoid, Biosteres longicaudatus. At the end of the 6-hr oviposition period, larvae were ligated posterior to the ring gland so that some larvae had parasitoids anterior to the ligature while in others, the parasitoids were in the abdomen, posterior to the ligature. Ninety-two percent of the parasitoids anterior to the ligature hatched to the first through third instars. Parasitoids posterior to the ligature had a 75% egg hatch to the first instar only. No larval molts to the second or subsequent instars occurred in these parasitoids. Upon parabiosis to 3-day-old, unparasitized host pupae, the ligated larvae pupated and 97% of the first-instar parasitoids in these parabiosed larval abdomens molted to the second instar. Newly laid parasitoid eggs transplanted to 3-day-old pupal hosts had less than one-third of the egg hatch of those transplanted to first-day third-instar hosts. The data implicate the physiological state of the host (vis-a-vis pupation and associated events) as being an important factor in the development of the endoparasitoid.     

Acceptability and suitability of Spodoptera exigua (Hübner) for Cotesia icipe Fernandez‐Triana & Fiaboe on amaranth

The beet armyworm Spodoptera exigua (Hübner) (Lepidoptera: Noctuidae) is a polyphagous insect that is distributed worldwide and was recently reported as an important pest on African indigenous vegetables. Cotesia icipe Fernandez‐Triana & Fiaboe (Hymenoptera: Braconidae) is a recently described parasitoid, reported from various Afrotropical countries. This work investigated the performance of C. icipe on S. exigua infesting Amaranthus dubius Mart. ex Thell. under laboratory conditions. Cotesia icipe was aggressive on the host and successfully oviposited on S. exigua with 70% of parasitoid females ovipositing after 2 hr of exposure. Parasitoid densities significantly affected the parasitism rate and the nonreproductive larval mortality. Parasitism rate was 9.7 ± 0.8% and 59.5 ± 3.1% for a single and cohort of five females released, respectively, when offered 50 host larvae. The cohort female release resulted in significantly higher larval nonreproductive mortality than the single release. However, there was no significant difference between parasitoid release densities in regard to pupal nonreproductive mortality. The larval and pupal mortalities in the presence of C. icipe were significantly higher than the natural mortalities at both parasitoid release densities. The parasitoid sex ratio was female‐biased for the cohort females but balanced when a single female was released. The hind tibia and forewing lengths were not affected by the density of female parasitoids but there were variations according to sex. The implication of these findings on the potential use of C. icipe for biological control of S. exigua in amaranth production systems is discussed.     

Interactions between the Parasitoid Rhopus nigroclavatus (Ashmead) (Hymenoptera: Encyrtidae) and its Mealybug Hosts Tridiscus sporoboli (Cockerell) and Trionymus sp. (Homoptera: Pseudococcidae)

This research investigated age-class-specific parasitism rates of the buffalograss mealybugs Tridiscus sporoboli (Cockerell) and Trionymus sp. by Rhopus nigroclavatus (Ashmead) (Hymenoptera; Encyrtidae), size class preference of this parasitoid, and mealybug–parasitoid interactions through choice and no-choice studies. In the no-choice studies, the mean rates of parasitism by R. nigroclavatus were 45, 20, 0, and 0%, respectively, for mealybugs adult female, third and fourth instars, first and second instars, and eggs. Choice studies indicated that rate of parasitism increased with host size. The mean rates of parasitism on mealybugs in the choice studies were 100% for adult females, 24% for third and fourth instars, 0% for first and second instars, and 0% for eggs. A second set of choice studies investigating mealybug/parasitoid behavior revealed that R. nigroclavatus oviposits in all post-egg mealybug age classes, but first and second instars were less often parasitized than older mealybugs.     

Winter survival and oviposition before and after overwintering of a parasitoid wasp, Ooencyrtus nezarae Ishii (Hymenoptera: Encyrtidae)

Winter survival and oviposition before and after overwintering in Ooencyrtus nezarae, an egg parasitoid of phytophagous heteropterans, were examined in Osaka, Japan. Eggs of Riptortus clavatus parasitized by O. nezarae were kept under natural photoperiod and temperature. When honey was supplied, some female adults emerging from early September to late November overwintered. The percentage of overwintering individuals increased as the date of adult emergence advanced. Most female adults supplied with honey and hosts oviposited soon after emergence, then stopped laying eggs. Female adults emerging in mid‐October and early November laid eggs and then overwintered. The induction of diapause in the field seems to vary greatly depending on host availability. Without honey, the survival time of female adults was very short, whether host eggs were supplied or not. After overwintering, most females began to lay eggs in early May if host eggs were supplied, and they produced both male and female progeny. In the study area, a legume field in Osaka, parasitization by O. nezarae was observed from early July to November.     

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.     

Virus-like Particles in the Calyx of Phanerotoma flavitestacea (Hymenoptera: Braconidae) and Their Transfer Into Host Tissues

Virus-like particles occur in the calyx fluid and cells of the braconid parasitoid, Phanerotoma flavitestacea Fischer. These particles occur in the cytoplasm, nucleus, and cytoplasmic vacuoles of the calyx wall cells as well as in the lumen of the calyx. They were subsequently transferred to the host, Paramyelois transitella (Walker), during oviposition of the parasitoid. Within 2 hours after the wasps had oviposited, similar virus-like particles were found in the host in the cytoplasm of developing muscle tissue, fat body, and nerve tissue. Such particles were absent in unparasitized hosts. The significance of these findings as related to the avoidance of a host response is discussed.     

Age-dependent functional response of Aphidius matricariae (Hymenoptera: Braconidae) on tobacco aphid,Myzus persicae nicotianae (Hemiptera: Aphididae)

This study was performed to investigate the age-dependent functional response of Aphidius matricariae Haliday (Hymenoptera: Braconidae) at varying densities of tobacco aphid, Myzus persicae nicotianae Blackman (Hemiptera: Aphididae). Six densities (2, 4, 8, 16, 32, and 64 numbers) of third instar nymphs of M. persicae nicotianae were daily exposed to one pair of 1-day-old A. matricariae until the parasitoid female died. Each host aphid density was replicated 10 times. The experiments were conducted at constant environmental conditions (25 ± 1 °C, 70 ± 5% RH and 16:8h L:D photoperiod). The logistic regression model demonstrated that the functional response of A. matricariae was type III in the second day and type II in the first day and from third to the end of adult parasitoid lifetime. The handling time (T_h) was lowest in the first day (0.2305 h) and highest in the sixth day (2.9296 h) of the parasitoid’s life. The searching efficiency (a) values were 0.0573, 0.0649, 0.0503, 0.0229, and 0.0077 h^?1 in the first, third, fourth, fifth, and sixth days of the parasitoid’s life, respectively. Also, the b value for type III functional response was 0.0147 in the second day. The values of maximum attack rate (T/T_h) ranged from 104.12 nymphs/24 h in the first day to 19.8 nymphs/24 h in the sixth day of parasitoid’s life. Based on the results, A. matricariae showed a high potential to control the population of tobacco aphid and the highest efficiency gained in the first half of the female parasitoid life.     

Spermatogeny as a life-history index in parasitoid wasps

Reproduction is a major life-history trait but it has been studied mostly in relation to female reproductive effort. Recently, an ovigeny index using the proportion of oocytes ready to be oviposited at eclosion has been proposed for female insect parasitoids. Here, we propose a spermatogeny index for male parasitoid wasps. Prospermatogenic species have an index of 1, have all their spermatozoids mature at emergence and do not produce more later in life. At the other end of the spectrum, synspermatogenic species have no spermatozoids at emergence and produce them later in life. The level of spermatogeny should be linked to several other life-history parameters such as longevity, size, nutrition, distribution of mating opportunities and dispersion before and after mating. Data presented for some parasitoid species support the presence of variability in this male life-history parameter.     

Host discrimination by the endoparasitoid Psyllaephagus pistaciae (Hymenoptera: Encyrtidae): A case of time-dependent ability

Host-discrimination behavior by the koinobiont parasitoid, Psyllaephagus pistaciae Ferrière (Hymenoptera: Encyrtidae), the major biocontrol agent of the common pistachio psylla, Agonoscena pistaciae Burckhardt & Lauterer (Hemiptera: Psyllidae), in Iran, was investigated in the laboratory. The results demonstrated that P. pistaciae quickly detected and avoided freshly parasitized hosts, either after antennating or by probing with their ovipositor. However, this discriminatory ability declined with time, probably influenced by both external and internal markers left by the previous ovipositing female. The results also confirmed that female P. pistaciae responded to changes in host quality associated with the parasitoid's larval development 4 days after the initial parasitization, clearly indicating that the second female could detect the presence of the larvae and adjust her host-selection decision. In addition, psyllid nymphs treated topically with a solution of Dufour's gland were rejected by the parasitoid, showing that Dufour's gland secretion had a significant effect as a host marking chemical. The current study also showed that superparasitism increased the host-mortality and that the rate of encapsulation decreased, suggesting that, when two eggs are laid in the same psyllid nymph, only one parasitoid develops to an adult.