Source and characterization of an egg recognition kairomone of Telenomus heliothidis, a parasitoid of Heliothis virescens

ABSTRACT. The ability of Telenomus heliothidis Ashmead (Hymenoptera; Scelionidae) to recognize its host, the eggs of Heliothis virescens (F.) (Lepidoptera; Noctuidae), was determined in laboratory studies. The material present in the accessory gland of female H. virescens adults acted as an egg recognition kairomone for T. heliothidis. Host eggs which lacked the kairomone were not attacked, while glass beads the size of H. virescens eggs and coated with the accessory gland material were. The parasitoid failed to respond to the kairomone unless it was associated with a target the size and shape of normal hosts. The kairomone is heat stable, but was deactivated by proteolytic enzymes. The accessory gland contained 200 mg/ml protein which suggests that the kairomone is proteinaceous.     

Characterization of a cDNA encoding a putative chitinase from teratocytes of the endoparasitoid Toxoneuron nigriceps

Females of insect endoparasitoids inject substances along with the egg at oviposition, which can regulate or induce changes in a number of host physiological processes to benefit the developing parasitoid. These changes can be caused by substances such as venoms, calyx fluids, or symbiotic‐associated virus particles (polydnavirus), which are injected by females along with their eggs, and by substances secreted by parasitoid‐derived tissues (teratocytes) or the developing parasitoid larvae. Teratocytes (dissociated cells of the serosal membrane after parasitoid eclosion) release substances that have roles (i) in parasitoid nutrition, (ii) in the digestion of host tissues, and (iii) in the regulation of host development. Teratocytes of Toxoneuron nigriceps (Viereck) (Hymenoptera: Braconidae) have been implicated in the arrestment of the host development and in the regulation of circulating levels of host ecdysteroids. Here we describe the cDNA of a teratocyte‐secreted chitinase and its expression during parasitoid development, and discuss its putative role in this host–parasitoid association.     

霍氏啮小蜂的卵巢发育和卵子发生

霍氏啮小蜂是一些鳞翅目和双翅目昆虫蛹的寄生蜂。本文描述该寄生蜂从亚洲玉米螟羽化后0,6 12,18,24,30,36,42,48,54,60,66和72 h的卵巢发育和卵子发生过程。霍氏啮小蜂雌性生殖系统为典型的膜翅目姬小蜂类型,包括2个卵巢,每个卵巢通常有8-9根卵巢管,另有1对输卵管附腺,1个受精囊和1个受精囊腺等。卵黄和卵子形成于羽化后12 h,刚羽化的雌蜂不含成熟卵(成熟卵指数OI=0),成熟卵出现在羽化后24 h,成熟卵数量随后不断增加至数量较为稳定。卵巢管内卵数与雌蜂的体长和头宽呈正相关,相关系数分别为0.859和0.907。研究结果表明霍氏啮小蜂属卵育型(synovigeny)寄生蜂,本文还讨论了卵育型霍氏啮小蜂的卵成熟策略。     

Volatile and Contact Chemicals Released by Nezara viridula (Heteroptera:Pentatomidae) Have a Kairomonal Effect on the Egg Parasitoid Trissolcus basalis (Hymenoptera: Scelionidae)

The responses of females of the egg parasitoid Trissolcus basalis (Wollaston) (Hymenoptera: Scelionidae) to volatile and contact chemicals from its host Nezara viridula (L.) (Heteroptera: Pentatomidae) were investigated in a Y-tube olfactometer and under open arena conditions. In the Y-tube tests, volatiles from virgin males and from females in a preovipositional state attracted T. basalis females, while volatiles from host virgin females did not. In an open arena, traces left by N. viridula adults in different physiological conditions function as contact cues inducing the wasps to remain longer in the arena and to change the pattern of their walking behavior. However, only contact kairomones from N. viridula mated females in a preovipositional condition induced an arrestment response characterized by an increase in patch searching time and turning rates and a reduction in linear speed. The chemical ecological implications of these results on this host–parasitoid association are discussed.     

The Importance of the Parasitoid Egg in Host Discrimination by Cephalonomia stephanoderis , an Ectoparasitoid of the CoVee Berry Borer Hypothenemus hampei

A laboratory study was performed to examine the importance of the parasitoid egg in host discrimination by Cephalonomia stephanoderis Betrem (Hymenoptera: Bethylidae), an ectoparasitoid of the coVee berry borer, Hypothenemus hampei (Coleoptera: Scolytidae). C. stephanoderis was able to distinguish between parasitized and unparasitized hosts, and preferred to oviposit on the latter. This discriminative ability appeared to be innate, not learned. W hen the parasitoid egg was removed artificially the host was accepted for parasitism once more, suggesting that the presence of the egg is the most important factor in host discrimination. However, it was not possible to exclude the eVect of a chemical marker present either on or within the host, and/or associated with the presence of the parasitoid egg.     

Effects of parasitization of Trichoplusia ni by Chelonus sp.

ABSTRACT. Parasitization of Trichoplusia ni (Huebner) (Lepidoptera: Noctuidae) by Chelonus sp. (Hymenoptera: Braconidae), an egg-larval parasitoid, leads to precocious cocoon spinning of the host in the fourth (penultimate) stadium followed by parasitoid emergence from the prepupa. We have investigated the mechanism by which Chelonus sp. disrupts host development. The developing larva and fluids injected by the adult female separately from the egg, are not the source of these effects, but it remains a possibility that the teratocytes, originating from the trophamnion of the parasitoid egg, are responsible. The titre of the juvenile hormone esterase activity in the haemolymph of the parasitized fourth instar host is similar to that in the initial period of the final instar of normal T. ni, but lacks the postwandering peak of activity. The increased JHE activity leads to a reduced JH titre early in the fourth stadia. This indicates that disruption of host development occurs within 12h after apolysis to the fourth stadium, if not before. Anti-juvenile hormone activity is not detected in extracts of parasitized T. ni. The morphological and behavioural changes associated with precocious development of the T. ni host are prevented by applications of juvenile hormone I, juvenile hormone II and the juvenoid, Ro 10–3108, but not juvenile hormone III and the juvenoid R 20458. However, these applications fail to prevent the onset of juvenile hormone esterase activity, another marker of precocious development. These observations indicate that simple anti-juvenile hormone activity may not be the mechanism of disruption of host development. Development of the parasitoid is disrupted by application of Ro 10–3108 and juvenile hormones I, II and III, but timing of eclosion is only affected by application of juvenile hormone I, juvenile hormone II and Ro 10–3108. This observation may indicate a discrimination by the parasitoid between its own juvenile hormone III and the host's juvenile hormone II.     

Effect of biotic and abiotic factors on reproductive parameters of the synovigenic endoparasitoid Venturia canescens

Abstract.  The main effects and interactions of adult age, access to food and host deprivation, on the egg load of Venturia canescens (Gravenhorst) (Hymenoptera: Ichneumonidae), a larval parasitoid of Ephestia kuehniella (Lepidoptera: Pyralidae) and other pyralids, were studied in the laboratory. Intraspecific variation in the number of ovarioles was also studied. There was a positive and significant correlation between wasp size and both egg load and ovariole number, with the reproductive system of large wasps containing significantly more ovarioles and mature eggs than small wasps. Newly emerged adult wasps contained 27 ± 2.4 ovulated (mature) eggs in their lateral oviducts. Access to food and host deprivation were the only conditions under which egg load increased with parasitoid age. Wasps that had access to hosts immediately after emergence showed a significant decline in their egg complement, irrespective of food presence. Under conditions of both host and food deprivation, there was practically no alteration of egg load with parasitoid age. A three-way analysis of variance revealed that egg load varies significantly with food or host access but not with parasitoid age. All interactions among the three factors were significant. It is confirmed that there is no egg resorption in V. canescens and that egg production stops in the absence of food.     

Improved rearing and release procedures forCotesia melanoscela (Hym.: Braconidae), an early season parasitoid ofLymantria dispar (Lep.: Lymantriidae)

Rearing and release procedures forCotesia melanoscela (Ratzeburg) (Hymenoptera: Braconidae), an early-season parasitoid ofLymantria dispar (L.), were evaluated in 1995 and 1996 at the Beltsville Agricultural Research Center, Beltsville, Maryland, USA. Mass rearing procedures were developed during the winter/spring of 1995/1996 to produce diapausingC. melanoscela cocoons for experimentation. Program efficiency as measured by percent adult emergence was estimated to be 84% with a measured sex ratio of 48:52 M:F, at an estimated cost per 1 000 cocoons of U.S. $5.26 for materials and a labor requirement of approximately 6 hours. A host larvae:female parasitoid ratio of 100:1 was found to be most efficient in terms of number of parasitoid cocoons produced per female parasitoid, but a ratio of 100:2 was most efficient in terms of number of cocoons per rearing cup. OverwinteringC. melanoscela cocoons under natural conditions for 25 weeks (November placements) before expectedL. dispar egg hatch (mid-April) resulted in excellent synchrony with host egg hatch. Alternatively, holdingC. melanoscela cocoons in cold storage for 23 or more weeks at 5°C resulted in adult parasitoid emergence beginning 10 days after removal from cold storage, and proper timing of removal would result in similar synchrony of adult emergence with host egg hatch. These alternative release strategies give managers increased program flexibility.     

The ovipositing female of Ooencyrtus telenomicida relies on physiological mechanisms to mediate intrinsic competition with Trissolcus basalis

Ongoing studies by our group showed that the outcome of the intrinsic competition between two solitary egg parasitoids, Trissolcus basalis (Wollaston) (Hymenoptera: Scelionidae) and Ooencyrtus telenomicida (Vassiliev) (Hymenoptera: Encyrtidae), is dominated by O. telenomicida. In this article we investigated the role played by the ovipositing O. telenomicida female in the suppression of a T. basalis competitor. Laboratory experiments were conducted by allowing an O. telenomicida female to puncture the eggs of Nezara viridula (L.) (Heteroptera: Pentatomidae) with her ovipositor (= no oviposition) or to parasitize them. The results show that O. telenomicida relies on some physiological mechanisms to mediate its interspecific intrinsic competition with T. basalis. In fact, the emergence of T. basalis was strongly reduced in host eggs that were parasitized either before or after being punctured by O. telenomicida at fixed time intervals (5, 15, 30, or 45 h). The low percentage of emergence of T. basalis (ranging from approximately 4–20%) was a consequence of the delay and growth rate reduction of larval development. Furthermore, the percentage of eclosion of N. viridula nymphs was negatively affected by the O. telenomicida female’s punctures (96% from healthy host eggs, 4% from punctured host eggs). Host eggs punctured or oviposited in by O. telenomicida showed alterations in the ooplasm including some melanized‐like areas near the hole made with the ovipositor; such alterations indicate that the adult parasitoid releases substances that affect the host eggs survival. These results suggest that the O. telenomicida female influences both the physiological interspecific parasitoid‐parasitoid interaction, as well as the host‐parasitoid interaction, providing, for the first time in egg parasitoids, evidence that physiological suppression of some competitive egg parasitoids is mediated by the ovipositing female.     

Effects of seasonal variation in host quality and availability on parasitism by the egg parasitoid Telenomus coloradensis

Hemlock looper, Lambdina fiscellaria (Guenée) (Lepidoptera: Geometridae), is one of the most important defoliator in North American forests. The common egg parasitoid Telenomus coloradensis Crawford (Hymenoptera: Platygastridae) plays a significant role as a natural control agent, with parasitism levels in spring typically higher than in fall. The objectives of this study were to quantify changes in host acceptance and reproductive performance of the parasitoid in relation to (1) host egg fertilization in fall, (2) host diapause status, (3) host embryonic development in spring, and (4) host deprivation during summer. Our results indicate that T. coloradensis do not have the capacity to develop in unfertilized host eggs, whereas early‐diapausing eggs are more suitable for the parasitoid than post‐diapausing eggs. Furthermore, the host physiological suitability decreases with embryonic development in spring. Finally, a host deprivation period during the summer tends to negatively affect the parasitic activity of T. coloradensis. These laboratory results confirm previous hypotheses concerning T. coloradensis seasonal ecology and contribute to a better understanding of the effect of hemlock looper egg physiology and availability on the reproductive potential of T. coloradensis.     

Influence of host deprivation on egg load and oviposition behaviour of Brachymeria intermedia, a parasitoid of gypsy moth

Abstract. Mated female Brachymeria intermedia (Hymenoptera: Chalcididae) deprived, since emergence, from pupae of their host Lymantria dispar (Lepidoptera: Lymantriidae), accumulated eggs but had a very low rate of hostacceptance. Parasitoids that were host-deprived after encountering pupae early in life also accumulated eggs, but maintained a high acceptance rate. Thus early exposure to hosts promoted active reproductive behaviour. Total egg production depended on the total number of pupae encountered, indicating that B.intermedia adjust their egg production to host availability. Hence, in B. intermedia both the physiological state of the parasitoid (age and egg load) and the informational state (in this case host-availability and experience) interact to shape oviposition behaviour.     

Lethal and sublethal effects of fenitrothion and deltamethrin residues on the egg parasitoid Trissolcus grandis (Hymenoptera: Scelionidae)

Effects of fenitrothion and deltamethrin, the most commonly used insecticides in Iran for controlling Eurygaster integriceps Puton (Heteroptera: Scutelleridae), in wheat and barley were assessed on adults and preimaginal stages of egg parasitoid Trissolcus grandis Thompson (Hymenoptera: Scelionidae). Adult parasitoids exposed to field recommended concentrations of the insecticides suffered 100% mortality within 24 h. LC50 values of fenitrothion and deltamethrin for T. grandis were 8.1 and 3.9 microg (AI) /ml, respectively. Both insecticides and the preimaginal stage of exposure had a significant influence on the level of adult emergence from host eggs treated with field recommended rates. Fenitrothion and deltamethrin reduced the emergence rates by 18 and 34.4%, respectively, compared with the control. However, neither insecticide significantly affected the longevity or reproductive capacity of emerged females, or the sex ratio of their progeny. This study revealed that application of these insecticides should be avoided in early season to conserve natural or released populations of T. grandis. Both insecticides seemed to be detrimental to the parasitoid and need to be applied cautiously through season.     

Host recognition by the egg–larval parasitoid Chelonus inanitus: effects of physical and chemical cues

Parasitoid wasps are widely used as biocontrol agents. For successful deployment, it is important to know how they search for and recognize their hosts. They often use chemical cues in host searching and recognition. Little attention has been paid to the use of physical cues, particularly when combined with chemical cues, though a combination of cues may improve searching efficiency. Chelonus inanitus L. (Hymenoptera: Braconidae) is a solitary, egg–larval parasitoid wasp that parasitizes various lepidopteran species, including serious crop pests. In this study, effects of physical and/or chemical cues on host recognition were investigated in C. inanitus by observing its antennal searching, ovipositor probing, and the duration of arrestment. In a physical cue test, egg-mass models with glass beads differing in size were used, and a single-egg model with a specific size of bead. Egg-mass models with beads of 0.4–1.0 mm in diameter arrested C. inanitus, in accordance with egg size and arrangement of the host. In a chemical cue test, extract from egg masses of the host Spodoptera litura (Fabricius) (Lepidoptera: Noctuidae), equivalent to half of a host egg mass and extract diluted 100-fold were used, and the undiluted extract stimulated C. inanitus. Untreated egg-mass models (0.2-mm-diameter glass beads) and diluted egg extract arrested C. inanitus less effectively; however, when glass beads were treated with the diluted extract, arrestment increased to the same degree as the undiluted extract. These results suggest that C. inanitus uses both physical and chemical cues to recognize eggs of its host, S. litura, and that these stimuli act synergistically.     

Source and characterization of host recognition kairomones of Tetrastichus hagenowii, a parasitoid of cockroach eggs

ABSTRACT. Recognition of Periplaneta americana (L.) oothecae by the egg parasitoid Tetrastichus hagenowii (Ratzeburg) (Hymenoptera: Eulophidae) is mediated by mucopolysaccharides from the salivary glands, and calcium oxalate from the colleterial glands of the roach. Calcium oxalate, the most active compound isolated, only elicits stereotyped host recognition behaviour from the parasitoid when perceived in the context of a curved surface. Although empty oothecae contain calcium oxalate, female T. hagenowii appear to determine the unsuitability of these oothecae soon after drumming begins and abandon them. The potential usefulness of host recognition kairomones in biological control is discussed.     

Increasing host age does not have the expected negative effects on the fitness parameters of an egg parasitoid

Host age is an important determinant of host acceptance and suitability for egg parasitoids. As host embryonic development advances, the quality of resources available to the parasitoid offspring typically declines, usually resulting in reduced acceptance levels by foraging females and lower offspring fitness. We examined the ability of the parasitoid Telenomus podisi Ashmead (Hymenoptera: Scelionidae) to parasitize and develop in Podisus maculiventris (Say) (Hemiptera: Pentatomidae) eggs of different ages. In laboratory experiments, we measured the effect of host age (6, 24, 48, 72, 96, or 120 h old) on parasitism rate and offspring fitness parameters such as survival, development time, sex ratio, and size. Contrary to our expectations, parasitism rate did not differ between host age treatments, nor did sex ratio allocation, offspring size, or the fecundity of newly emerged female offspring. However, parasitoid offspring had a longer development time with increasing host age. This trend was stronger for males than for females, which we suggest could reduce the degree of protandry among offspring emerging from older host eggs, thus increasing the rate of virginity upon leaving the emergence patch and resulting in more frequent off‐patch mating by female offspring in nature. Overall, our results suggest that all stages of P. maculiventris embryonic development are suitable for acceptance and development of T. podisi. Unlike most species of egg parasitoids, T. podisi has evolved mechanisms to utilize host resources, regardless of host developmental stage, with relatively minor fitness consequences.     

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.     

Fitness-related changes in laboratory populations of the egg parasitoid <Emphasis Type="Italic">Trichogramma galloi</Emphasis> and the implications of rearing on factitious hosts

In this study, we used the specialist egg parasitoid Trichogramma galloi Zucchi, 1988 (Hymenoptera: Trichogrammatidae) to test whether development over multiple generations on a factitious host would later influence performance on the target pest. We also investigated possible adaptations to laboratory rearing conditions by monitoring over 50 and 46 generations life history traits of T. galloi populations maintained on the natural [Diatraea saccharalis (Fabr., 1794) (Lepidoptera: Crambidae)] and factitious host [Ephestia kuehniella (Zeller, 1879) (Lepidoptera: Pyralidae)], respectively. We detected a possible sign of adaptation to rearing conditions in early generations on the emergence of the population maintained on the natural host. We also verified a reduction in fecundity and emergence of parasitoids maintained on the factitious host when fitness was evaluated on the natural host, confirming the prediction that the use of factitious hosts can lower fitness on the target pest. The importance of these results for the effectiveness of biological control programs is discussed.     

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.     

Sexual attractiveness shared by both sexes mediates same‐sex sexual behaviour in the parasitoid wasp Telenomus triptus

The mating behaviour of a quasi‐gregarious egg parasitoid Telenomus triptus Nixon (Hymenoptera: Platygastridae), which exploits egg masses of a stink bug Piezodorus hybneri (Heteroptera: Pentatomidae), is examined in the laboratory. In this parasitoid wasp, male adults that emerge earlier stay at the natal egg mass and mate with subsequently emerging females. In the present study, a male adult that encounters the emergence of another male always waits for it to egress, and then mounts the newly emerging male. To examine why males of T. triptus show same‐sex sexual behaviour, male adults are presented with a parasitized host egg mass or a freshly killed wasp. Male adults are observed to remain at host egg masses from which only male wasp(s) had emerged. In addition, male adults attempt to copulate with freshly killed young male wasps. It is suggested that newly emerging male wasps are targets of same‐sex sexual behaviour because they possess cues for male sexual behaviour similar to the cues of females. Both the sex and age of freshly killed wasps affect the frequency of the sexual behaviour of male adults: females are more attractive than males, although their attractiveness declines with age. When the mating opportunity is restricted to the natal egg mass, the costs of failing to notice newly emerging female adults should be extremely high. Therefore, males are forced not to discriminate the sex, resulting in same‐sex sexual behaviour.     

Field collection of egg parasitoids of Pentatomidae and Scutelleridae in Northwest Italy and their efficacy in parasitizing Halyomorpha halys under laboratory conditions

The invasion of Halyomorpha halys (Stål) (Hemiptera: Pentatomidae) has caused severe economic damage in crops in North America and Europe, motivating research to identify its natural enemies, both in native and invaded areas. In its Asian native range, the main natural enemies are egg parasitoids, among which the most effective are Trissolcus japonicus (Ashmead) and Trissolcus mitsukurii (Ashmead) (Hymenoptera: Scelionidae) in China and Japan, respectively. In Europe, biology, host range, and impact of most native scelionid species are not well‐known. The present study aimed to investigate (1) presence and abundance of scelionid species that parasitize native Pentatomidae and Scutelleridae eggs in Northwest Italy, and (2) their ability to develop on H. halys eggs. During 4‐year field surveys, egg masses were collected and reared until bug nymph or adult parasitoid emergence. Then, the obtained scelionid females were tested for their ability to parasitize H. halys eggs in laboratory no‐choice experiments. Egg masses of all collected bug species were parasitized, and Telenomus spp. (Hymenoptera: Scelionidae), Trissolcus belenus (Walker), and Anastatus bifasciatus (Geoffroy) (Hymenoptera: Eupelmidae) were the most common parasitoids. In the laboratory, Trissolcus kozlovi Rjachovskij was the only species to significantly produce offspring from fresh H. halys eggs, whereas all tested Trissolcus species significantly induced host egg abortion (non‐reproductive effects). This study provides knowledge of the parasitoid species associated with native bugs, and represents a starting point to investigate the intricate interactions between native and exotic parasitoids recently found in northern Italy. These egg parasitoids could potentially be effective biocontrol agents of H. halys.