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.     

Source of the host marking pheromone in the egg parasitoid Trissolcus basalis (Hymenoptera: Scelionidae)

After oviposition, Trissolcus basalis females always mark the host's surface, depositing host marking substances for herself and to warn other ovipositing females. The perception of these host marking substances, probably through the antennae, can induce the female to leave and seek healthy hosts. Parasitoid females exposed to conspecific parasitized egg masses left the host egg masses significantly more often than when exposed to non-parasitized egg masses. More egg mass leaving behavior also was observed when the egg masses were treated with Dufour's gland secretion but not when treated with secretion from the common oviducts. The common oviduct has a secretory epithelium that produces electron-dense vesicles, probably containing proteinaceous substances. The secretory cells of the accessory gland, Dufour's gland, contain electron-lucid vesicles, whose secretion appears to be a lipid similarly to that found in pheromone secreting glands. Ultrastructural and behavioral evidence suggests that Dufour's gland is the host marking pheromone source.     

Adult biology ofTelenomus remus [Hymenoptera: Scelionidae] under laboratory conditions

Males ofTelenomus remus Nixon were attracted to the female by a pheromone present for the last 2 days of pupal life and the first few minutes after emergence, but not thereafter. Females in groups lived 6 times longwer than solitary ones; males did not show such a difference. Unmated males lived 8 days, and mated ones 1 day only. The normal sex ratio of 60–70 % females declined to 22 % as the females aged. Under conditions promoting superparasitism, the sex-ratio was close to 50 %. Duration of oviposition was 37 sec. with young females, but increased to over 50 sec. with age. All 3 layers of the host egg mass were parasitized equally. Several females may oviposit simultaneously on the same host batch without showing any aggression. About 60 progeny were produced during the first day of the female's life, but 20 or fewer thereafter.     

Response to host age of the egg parasitoid Anaphes n.sp. (Hymenoptera: Mymaridae)

The mymarid Anaphes n.sp. was observed to parasitize the eggs of the carrot weevil Listronotus oregonensis (LeConte) up to 138 h old at 23±1°C and 18L:6D. Adult emergence of the parasitoid was observed in 85%‐97.5% of the weevil eggs 42–138 h old and host embryogenesis was interrupted in nearly 100% of these eggs. In host eggs of 18 h old, parasitism was very high but not all parasitoids completed their development and their subsequent emergence was generally retarded. An.sp. could parasitize host eggs 162 h old, but such an attack rarely retarded the normal development of the host. The parasitoid was unable to complete its development in sterile carrot weevil eggs.     

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.     

Morphology of the antennal sex-gland in male Trissolcus basalis (Woll.) (Hymenoptera : Scelionidae), an egg parasitoid of the green stink bug,Nezara viridula (Hemiptera : Pentatomidae)

The 5th antennomere, beginning with the scape, of the male Scelionidae antenna has been referred to as a sex-segment due to its morphological and presumed sexual function. A study of the sex-segment of the parasitoid, Trissolcus basalis (Woll.) (Hymenoptera : Scelionidae) revealed a single ventral peg that is involved in releasing a secretion from an exocrine gland located in the antennomere. This is a “class 3” exocrine gland and the secretion appears to be partially proteinaceous since it is susceptible to protease digestion. Although the function of the sex-segment and gland remains unknown, the suggestion that it is involved in courtship is supported. Some sex-segment processes of other Scelionidae are also considered.     

Life history,immature morphology and adult behavior ofTelenomus solitus [Hymenoptera: Scelionidae]

The results of laboratory studies on the life history, morphology of immature stages, effects of food and water on adult longevity, host age preference for oviposition, parthenogenesis, and adult behavior ofTelenomus solitus Johnson are presented.Telenomus solitus completed its development in 10.8 days (egg, 0.3 day; 1^st instar, 2.7 days; 2^nd instar, 3.7 days; and pupa, 4.1 days). The adults lived longest when provided with both honey and water. Without food, they survived less than 24 h. Parasitization of host eggs varied significantly with their age, being highest in 42 h old eggs. Fewer than 10% of eggs older than 60 h were parasitized successfully. The periodicity of oviposition was unimodal; it occurred throughout the 24 h period and peaked in the late photophase. Females ofT. solitus are arrhenotokous. The behavioral sequences during courtship, mating, and oviposition are described.      

Cultivation of teratocytes of the egg parasitoid telenomus heliothidis (hymenoptera: Scelionidae)

Summary A method for in vitro cultivation of teratocytes from the egg parasitoidTelenomus heliothidis (Hymenoptera: Scelionidae) is described. Parasitoid eggs, from which teratocytes were derived, were collected from 24-h-oldHeliothis virescens (Lepidoptera: Noctuidae) eggs previously parasitized byT. heliothidis females. Optimal culture conditions, including species and concentration of serum, were determined experimentally. Thirty percentManduca sexta hemolymph or 10% chicken serum in Hink’s TNH-FH medium were found to generate the most satisfactory number of teratocytes per parasitoid larva. Teratocytes cultivated in vitro showed similar development and morphology to those produced in vivo. However, cultured teratocytes lived approximately 10 times longer than teratocytes in natural hosts and were not dependent upon the presence of the parasitoid larva for normal development.     

Population dynamics ofTelenomus fariai (Hymenoptera: Scelionidae), a parasite of Chagas' disease vectors IX. Larval competition and population size regulation under laboratory conditions

A series of experiments were carried out with the endophagous egg parasite Telenomus fariai on its host Triatoma phyllosoma pallidipennis to determine the possible role of intraspecific competition by the parasite progeny in population regulation of the parasite. Eight parasite densities (1, 3, 5, 10, 20, 30, 40, and 50 individuals per vial) were used, and the design of sequentially sacrificed replicates applied. Survivorship curves for each density indicated smaller number of progeny per host at higher densities, and the shapes of the curves suggested a relatively early mortality process.Morris' linear regression technique for determining within-generation density-dependence was used, and the results showed that only larval mortality could be identified as density dependent. The same technique applied within the larval stage proved that only mortality of larvae in their second, third, and fourth day of development were responsible for population regulation. The applicability of the technique, as well as the relevance of the results for natural population, is discussed.     

Reproductive biology of Fidiobia dominica (Hymenoptera: Platygastridae), an egg parasitoid of Diaprepes abbreviatus (Coleoptera: Curculionidae)

The reproductive biology of Fidiobia dominica Evans (Hymenoptera: Platygastridae) was studied in the laboratory (25.6 +/- 1 degrees C) using host eggs of Diaprepes abbreviatus L. (Coleoptera: Curculionidae). F. dominica readily parasitized D. abbreviatus eggs on both host plant and wax paper substrates. The number of egg masses parasitized and the number of offspring produced were higher when females were offered more than one host egg mass but did not differ when either two or three egg masses were offered. Female parasitoids that were provided with host eggs and a honey food source lived significantly longer than those that were not provided a food source; however, they did not parasitize more D. abbreviatus eggs. Oviposition occurred in host eggs from 0 to 7 d old, and host mortality was relatively consistent for eggs 0-5 d old and lower for eggs 6-7 d old. Successful parasitoid emergence seldom occurred after host eggs were 4 d old, and by 7 d, no adults successfully emerged. Developmental time from egg to adult was 19.3 +/- 0.2 d for males, significantly more rapid than the females (20.4 +/- 0.1 d). The mean longevity of adult females was 8.0 +/- 0.4 d, with a mean oviposition period of 2.7 +/- 0.3 d; males survived 8.1 +/- 0.4 d. The demographic parameters including intrinsic rate of increase (r(m)), generation time (T), and net reproduction (R(o)) were 0.142/d, 22 d, and 22.4 female eggs/d, respectively.     

Flight orientation behavior of Ooencyrtus nezarae (Hymenoptera: Encyrtidae), an egg parasitoid of phytophagous bugs in soybean

We conducted field experiments to examine flight orientation behavior of Ooencyrtus nezarae Ishii, an egg parasitoid of soybean feeding bugs, using sticky traps with synthetic aggregation pheromones of the host, Riptortus clavatus (Thunberg). Experiments were conducted in plots either containing a soybean field or not containing one. When the traps were suspended at heights of 0.5, 1.5 and 3.0 m above a soybean field in Kobe, O. nezarae females were most frequently caught in the 0.5 m traps. In an open field on the same campus where the vegetation was predominantly tall goldenrod, females were more frequently trapped at 1.5 or 3.0 m. In a soybean field in Kumamoto, females were most frequently trapped in the 0.8 m traps, compared to traps placed at 2.0, 4.0 and 6.0 m. In an oat field in the same area, females were trapped most frequently at heights of 0.8 and 2.0 m. Unlike females, only a small number of males were trapped in the fields in both areas. The result that O. nezarae females were caught in traps at higher positions in non‐host habitats than in soybean fields probably reflects differences in host searching behavior in or movement between the habitats. In host habitats, females mainly walk on the plants or fly within the plant canopy to search for hosts. In non‐host habitats, females may not intensively search for hosts within the plants, and directly respond to host pheromones in the traps at heights equal to or a little higher than the plant canopy.     

Biology of Anagrus atomus (Hymenoptera: Mymaridae), an egg parasitoid of the grape leafhopper Arboridia kermanshah (Homoptera: Cicadellidae)

Biology, morphology and oviposition behavior of Anagrus atomus (Linnaeus), an egg parasitoid of the grape leafhopper Arboridia kermanshah Dlabola in Isfahan, Iran, were investigated. Adults were smaller than those so far reported from other regions. Females continuously drummed on plant surfaces with their antennae to search for host eggs. Parasitoid eggs hatched 2–3 days after oviposition, and A. atomus had two larval instars. First instar larvae were sacciform and immobile. Second instar larvae appeared 4 days after oviposition and were very active, and doubled their body length. The prepupal and pupal stages lasted for 1 and 5–6 days, respectively. Adult emergence began 16 days after oviposition, and peaked on day 17.     

In vitro rearing of Ooencyrtus nezarae (Hymenoptera: Encyrtidae), an egg parasitoid of Riptortus pedestris (Hemiptera: Alydidae)

Ooencyrtus nezarae Ishii (Hymenoptera: Encyrtidae) is a polyphagous egg parasitoid of various true bugs, including Riptortus pedestris (Fabricius) (Hemiptera: Alydidae), a major pest of soybean and fruit trees in northeastern Asia. This study was conducted to develop artificial host eggs containing insect haemolymph for mass rearing of O. nezarae. The haemolymph of Chinese oak silkworm (Antheraea pernyi Guérin-Méneville) pupae was found to be most suitable for artificial eggs for the complete development of O. nezarae. Among 764 parasitoid eggs laid in artificial eggs, 49.2% successfully developed to adult parasitoids. The developmental time in artificial eggs was delayed by 3–5 days relative to that observed in natural host eggs; however there was no significant difference in the length of hind tibia between adults that emerged from artificial and natural host eggs. Moreover, new generations that emerged from artificial eggs reproduced on all egg types offered (either natural or artificial eggs). We propose that the artificial eggs developed herein can therefore be an effective means for mass rearing O. nezarae.     

The rearranged mitochondrial genome of Podagrion sp. (Hymenoptera: Torymidae), a parasitoid wasp of mantis

The complete sequence of the mitochondrial genome of Podagrion sp. (Hymenoptera: Torymidae) is described. The mitogenome was 15,845 bp in size, and contained typical sets of mitochondrial genes. The base composition of the Podagrion sp. mitogenome was also biased toward A + T bases (81.8%). The mitochondrial genome of Podagrion sp. has a weak AT skew (0.07) and a strong GC skew (?0.26). Podagrion sp. exhibits a novel rearrangement compared with the ancestral order, including six protein-coding genes (nad3, cox3, atp6, atp8, cox2 and cox1), which have inverted to the minor strand from the major strand. The A + T-rich region of Podagrion sp., which is located between trnN and trnI, have five tandem repeats. The apomorphic rearrangements, including the conserved block “cox3-atp6-atp8-cox2-cox1-nad5-nad4-nad4l-nad6-cob” and the special locations of trnV and trnA, were mapped onto the phylogeny of Proctotrupomorpha.     

Biological studies on two neotropical egg parasitoid species: Trichogramma nerudai and Trichogramma sp. (Hymenoptera: Trichogrammatidae)

The main biological attributes of two Neotropical egg parasitoids, the arrhenotokous Trichogramma nerudai and the thelytokous Trichogramma sp., were assessed under controlled laboratory conditions. Developmental time from egg to adult, and parasitoid survival, fecundity and fertility were studied using life tables. Results showed that T. nerudai had a faster developmental time than Trichogramma sp. (13.014±0.4019 and 13.595±0.4931 days, respectively). Both species showed similar life table statistics, r_m was 0.222 and 0.225 for T. nerudai and Trichogramma sp., respectively. Parasitoid survival averaged 95% for both species. The results obtained are discussed in the context of selecting one of these natural enemies as a potential biological control agent for the European pine shoot moth Rhyacionia buoliana (Lepidoptera: Tortricidae) in pine forests and the codling moth Cydia pomonella (Lepidoptera: Tortricidae) in apple orchards in Argentina.     

Host location by the gerrid egg parasitoidTiphodytes gerriphagus (marchal) (Hymenoptera: Scelionidae)

Mechanisms used by a scelionid egg parasitoid to locate its gerrid host eggs were studied using field experiments. Parasitism by Tiphodytes gerriphaguswas higher in exclosures containing adults of Limnoporus dissortisthan in cages without gerrids. Higher densities of adult gerrids were associated with higher rates of parasitism, but dispersion of aquatic vegetation did not apparently affect rates of parasitism. T. gerriphagusmay use visual cues, chemical cues, or a combination of both to locate their hosts. We discuss the possible influence of T. gerriphaguson the reproductive behavior of its hosts.     

Host-related antennal variation in the polyphagous egg parasite Telenomus alsophilae (Hymenoptera: Scelionidae)

Abstract. The solitary egg parasite Telenomus alsophilae was reared from the eggs of six lepidopteran hosts of two families. One-way analyses of variance showed statistically significant differences among host classes for means of (a) lengths and widths of all female antennomeres; and (b) in twelve of twenty-three means of log-transformed ratios constructed from pairs of antennomere dimensions. Principal component analyses of dispersion matrices of both untransformed and log-transformed antennomere dimensions were conducted to examine patterns of variation among variables. In both analyses PCI was a general component and interpreted as a measure of overall size. One-way analyses of variance of projection scores of OTUs on each of the first five principal components showed significant differences among host classes for means of scores on (a) PCI (both analyses) and (b) PC4 for untransformed data and PC3 for log-transformed data. Differences among hosts classes for scores on the latter two components are interpreted to indicate that antennal variation results from factors in addition to simple size differences among host eggs and the allometric growth of body parts of parasites reared from them.     

Parasitism, development and adult longevity of the egg parasitoid Telenomus nawai (Hymenoptera: Scelionidae) on the eggs of Spodoptera litura (Lepidoptera: Noctuidae)

When Telenomus nawai Ashmead or Trichogramma ostriniae Pang & Chen (Hymenoptera: Trichogrammatidae) was inoculated into intact egg masses of Spodoptera litura (Fabricius) that were covered with a scale-hair layer, T. nawai emerged from 76% of the eggs, while emergence of T. ostriniae was from only 3% of the eggs. When the hair layer was removed before inoculation, the rate by the latter increased to 21%. These observations confirmed that the hair layer effectively protects S. litura egg masses from attack by T. ostriniae, and that T. nawai parasitism is more effective. In order to assess the feasibility of T. nawai as a biological control agent for S. litura, parasitism rate, development time and adult longevity were examined in the laboratory. Emergence of T. nawai was observed in more than 95% of 0- and 1-day-old separated eggs of S. litura, but the rate decreased to 60% and 0% for 2- and 3-day-old eggs, respectively. The emergence rates of T. nawai were near 95% for temperatures in a range from 25 degrees C to 35 degrees C, but decreased to 80% at 20 degrees C. No parasitoids emerged at 15 degrees C. The developmental periods decreased as temperature increased from 20 degrees C to 35 degrees C. The developmental threshold occurred at 13.7 degrees C and 13.9 degrees C, and the effective accumulative temperatures from egg to adult emergence were 149.3 and 147.1 degree-days for females and males, respectively. Mean longevity of the adult females decreased with increasing temperature; 87.0 days at 15 degrees C and 9.5 days at 35 degrees C. The feasibility of the use of T. nawai for controlling S. litura is discussed.     

Host location in Oomyzus gallerucae (Hymenoptera: Eulophidae), an egg parasitoid of the elm leaf beetle Xanthogaleruca luteola (Coleoptera: Chrysomelidae)

Eggs of the elm leaf beetle Xanthogaleruca luteola are often heavily attacked by the chalcidoid wasp Oomyzus gallerucae. We studied the chemical signals mediating interactions between the egg parasitoid, its host, and the plant Ulmus campestris. Olfactometer bioassays with O. gallerucae showed that volatiles of the host-plant complex attract the parasitoid. In order to determine the source of attractive volatiles within this host-plant-complex, we tested separately the effect of odours of eggs, gravid elm leaf beetle females, faeces of the beetles and elm twigs (with undamaged leaves and leaves damaged either mechanically or by feeding of the beetles). Odours of faeces of the elm leaf beetle were attractive, whereas neither volatiles from eggs nor from gravid females acted as attractants. Volatiles from undamaged or damaged plants did not elicit a positive reaction in O. gallerucae, whereas volatiles from feeding-damaged plants onto which host eggs had been deposited were attractive. This latter result suggests that it is not feeding but deposition of host eggs onto elm leaves that induces the production of plant volatiles attractive to the egg parasitoid. Investigations of the search patterns of O. gallerucae within the habitat by laboratory bioassays revealed that the egg parasitoid encounters host eggs by chance. Contact kairomones from faeces were demonstrated to be important in microhabitat acceptance, while contact kairomones isolated from the host eggs are relevant for host recognition. Received: 12 February 1997 / Accepted: 29 April 1997     

Metaphycus parthenolecanii sp. n. (Hymenoptera: Encyrtidae), a parasitoid of Parthenolecanium corni (Hemiptera: Coccidae) in Iran

Metaphycus parthenolecanii Japoshvili sp. n. (Hymenoptera, Chalcidoidea, Encyrtidae), a parasitoid of the European fruit lecanium Parthenolecanium corni (Bouché, 1844) (Hemiptera, Coccoidea, Coccidae) in Iran, is described and illustrated.

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