Sex allocation and effects of superparasitism on secondary sex ratios in the gregarious parasitoid, Trichogramma chilonis (Hymenoptera: Trichogrammatidae)

The role of sex-controlling behaviour at oviposition in generating primary sex ratios, and the effect of larval competition on secondary sex ratios, were studied in the gregarious endoparasitoid, Trichogramma chilonis. The production of a fertilized (female) egg is indicated by the incorporation of a pause in abdominal movements during oviposition, while the absence of it indicates the production of an unfertilized (male) egg. During each ovipositional bout, the first male egg is deposited at the second oviposition, and thereafter at intervals of about eight eggs. This simple pattern enables the wasps to adjust their progeny sex ratios under local male competition to a wide range of host size. Inexperienced wasps do not distinguish between parasitized and healthy hosts. Immature mortality is not significantly different between the sexes when a host is attacked by a single wasp, while females suffer higher immature mortality than males when superparasitism occurs.     

Naturally occurring egg parasitoids of the genus Trichogramma (Hymenoptera: Trichogrammatidae) in a pomegranate orchard in Tunisia

Four Trichogramma species were found in a pomegranate orchard in Gabès, an arid region of Tunisia, from parasitized eggs of Ectomyelois ceratoniae Zeller (Lepidoptera: Pyralidae), an economically important insect pest. Identification based on assessment of male genitalia and internal transcribed spacer 2 ( ITS 2) sequences showed that they were T. bourarachae Pintureau and Babault, 1988, T. oleae Voegelé and Pointel, 1979, T. cacoeciae Marchal, 1927 and T. evanescens Westwood, 1833. Trichogramma evanescens is reported for the first time in Tunisia. Trichogramma cacoeciae was the largely dominant species in the analyzed samples, whereas T. bourarachae was present in a minor portion of 1.38%. The implications of these results for attempts at controlling E. ceratoniae are discussed.     

The effect of temperature and humidity on the bionomics of six African egg parasitoids (Hymenoptera: Trichogrammatidae)

The life table statistics of six native Kenyan species/strains of Trichogramma and Trichogrammatoidea were established using a factitious host Corcyra cephalonica, Stainton (Lepidoptera: Pyralidae), at eight different temperatures (10, 15, 20, 25, 28, 30, 32 and 35 degrees C) and two humidity levels (40-50 and 70-80%). The objective was to select insects with superior attributes for augmentative release against lepidopteran pests in horticultural crops. Both temperature and humidity affected developmental time and life table parameters of the parasitoids but temperature played a more critical role. Developmental time was inversely related to temperature. The intrinsic and finite rates of increase increased with temperature up to 30 degrees C. Both net reproduction rate and intrinsic rate of increase were higher at the lower humidity. Temperature inversely affected generation time of parasitoid strains regardless of the relative humidity. Two strains of Trichogramma sp. nr. mwanzai collected from both low and medium altitudes and Trichogrammatoidea sp. nr. lutea from the mid-altitudes, were better adapted to both low and high temperatures than the other strains, as indicated by the high intrinsic and net reproductive rates, at both humidity levels. These three strains appear to be promising candidates for augmentation biocontrol against the African bollworm Helicoverpa armigera in Kenya.     

Pollen as a source of food for egg parasitoids of the genus Trichogramma (Hymenoptera: Trichogrammatidae)

The value of corn pollen as a food source for Trichogramma brassicae Bezdenko (Hymenoptera: Trichogrammatidae) was assessed in the laboratory by testing the effect of different pollen diets on the longevity, capacity of parasitism and other life-table parameters. T. brassicae females that fed on corn pollen and water (wet filter paper dusted with pollen) lived significantly longer (4.97 days) than females that fed on water alone (2.67 days), but significantly shorter than those feeding on honey alone (8.37 days) or corn pollen and honey (8.23 days), respectively, in the absence of host eggs. The presence of European corn borer eggs (Ostrinia nubilalis Huebner, Lepidoptera: Pyralidae) significantly increased longevity. With host eggs available, female T. brassicae that fed on corn pollen and water lived significantly longer (4.9 days) than on water alone (2.60 days), but significantly shorter than on honey alone (12.33 days) or corn pollen and honey (12.17 days), respectively. The cumulative lifetime fecundity was 82.53 offspring pupae when fed on pollen and water, which was significantly higher than 61.70 for water alone but lower than feeding on honey alone (95.70) or corn pollen and honey (99.97), respectively. Compared to water alone, corn pollen and water increased the net reproduction rate (R₀), intrinsic rate of increase (r_m), and finite rate of increase (λ), and reduced the doubling time (T₂), but had no effects on the emergence rate, sex ratio, and mean generation time (T_c). In another set of experiments, 13 out of 30 females reduced the number of pollen grains offered to them. The average number of pollen grains utilized per female in this 48-h test 2.46±0.40 (mean±SE). Hence, T. brassicae used corn pollen as food that significantly increased longevity and fecundity of the tested females. Pollen and honey supplied a complete diet for T. brassicae, compared to pollen and water or water alone.     

Strategic ejaculation in the egg parasitoid Trichogramma turkestanica (Hymenoptera: Trichogrammatidae)

Abstract.  1. Several parameters influence sperm allocation by males, including their size and sperm stock, intra-specific variability, quality of females', as well as the risk and intensity of sperm competition.
2. Models predict that males should invest the maximum ejaculate size when sperm competition intensity is low. As sperm competition intensity increases, males should decrease the number of sperm transferred during mating.
3. This decrease in sperm transfer to females occurs because the benefits gained by males with each extra unit of expenditure on sperm decrease. When sperm supply is not unlimited, males could expect a better return by keeping some or all sperm for mating under lower competition intensity.
4. In this study, the ejaculate size of males that were kept in groups of one, five or 10 males prior to mating, has been investigated in the haplodiploid egg parasitoid Trichogramma turkestanica Meyer (Hymenoptera: Trichogrammatidae).
5. As predicted by theory, the number of sperm transferred decreased significantly with an increase in the number of rivals.
6. This is the first study showing strategic sperm allocation depending on sperm competition intensity in a parasitoid.     

Trichogramma species (Hymenoptera: Trichogrammatidae) parasitoids of Lepidopteran eggs in Lara State, Venezuela

This work was conducted to determine the occurrence of Trichogramma wasp species in Lara State, Venezuela. Lepidopteran egg samples collected from various crop leaves were observed daily under laboratory conditions for emergence of Trichogramma species adult wasps. Trichogramma were also obtained from traps containing eggs from the hosts Sitotroga cerealella (Olivier) (Lepidoptera: Gelechiidae) and Spodoptera frugiperda (J. E. Smith) (Lepidoptera: Noctuidae). In the laboratory, specimens of Trichogramma species were mounted on microscope slides to show male genitalia and other morphological characters used for its identification. Trichogramma atopovirilia Oatman y Platner emerged from S. frugiperda eggs collected in a cornfield at La Palomera, Humocaro Alto and at El Parchal, Humocaro Bajo, Municipio Morán. T. atopovirilia also emerged from S. frugiperda eggs collected in a cornfield at Sabana Grande and from S. cerealella eggs used as traps in a pepper culture at Tintinal, Municipio Andrés Eloy Blanco. T. exiguum Pinto y Platner emerged from S. frugiperda eggs used as traps or collected in cornfield at Totumito and at La Palomera, Humocaro Alto. T. pretiosum Riley emerged from Phthorimaea operculella Zeller eggs collected on stored potatoes at Monte Carmelo, Sanare. The identification of Trichogramma species in Lara State would be useful in the development of biological control programs for lepidopteran pests.     

Thermal requirements of Fidiobia dominica (Hymenoptera: Platygastridae) and Haeckeliania sperata (Hymenoptera: Trichogrammatidae), two exotic egg parasitoids of Diaprepes abbreviatus (Coleoptera: Curculionidae)

Diaprepes abbreviatus is an exotic root weevil occurring in southern US. It is a highly polyphagous species which can complete its entire life cycle on citrus and several woody ornamental plants. The lack of native egg parasitoids for this weevil in citrus orchards has triggered efforts to evaluate candidate egg parasitoids from the Caribbean Region into Florida. The egg parasitoids Fidiobia dominica and Haeckeliania sperata are two exotic natural enemies of D. abbreviatus recently introduced in the US in a classical biological control program. The thermal requirements of both parasitoids were studied in the laboratory. The upper development threshold (UDT) of F. dominica was 30.0°C, its maximal development rate (MDR) occurred at 27.6°C, its lower development threshold (LDT) was 9.6°C and its thermal constant (K) for development from egg to adult was 293.1 DD. For H. sperata, UDT was 35.0°C, MDR occurred at 31.0°C, LDT was around 15°C and K was 188.1 DD. Based on these results, both species would be able to complete 17 to 18 generations annually in southern Florida. However, host availability during critical periods could severely impair the ability of these egg parasitoids to establish and successfully control D. abbreviatus in areas where winter temperatures fluctuate around 12°C, the LDT for this pest.     

Identification of egg parasitoids of rice leafhoppers and planthoppers (Hemiptera: Cicadellidae and Delphacidae) of economic importance in Taiwan,part 2: Trichogrammatidae (Hymenoptera)

Trichogrammatid egg parasitoids (Hymenoptera: Trichogrammatidae) of rice leafhoppers and planthoppers (Hemiptera: Cicadellidae and Delphacidae) known from Taiwan are identified taxonomically and illustrated, and their host associations are determined by rearings using sentinel eggs of the key pest species. The previously published records of such egg parasitoids in Taiwan are critically analyzed in the unfortunate absence of almost any voucher specimens. Pseudoligosita nephotetticum (Mani) is shown to be the most common egg parasitoid of the two leafhopper and three planthopper pests of rice in Taiwan: the zig-zag leafhopper Maiestas dorsalis (Motschulsky) (a new host record), the green rice leafhopper Nephotettix cincticeps (Uhler), and the delphacids Laodelphax striatella (Fallén) (small brown planthopper), Nilaparvata lugens (Stål) (brown planthopper), and Sogatella furcifera (Horváth) (whitebacked planthopper). Paracentrobia (Brachistella) andoi (Ishii) syn. nov., originally described from Japan as Japania andoi Ishii, and Paracentrobia (Brachistella) garuda Subba Rao syn. nov. from Thailand are synonymized under Paracentrobia (Brachistella) bicolor (Girault) from northern Queensland, Australia; Pseudoligosita tachikawai (Yashiro) syn. nov., originally described as Oligosita tachikawai Yashiro from Japan, is synonymized under Pseudoligosita nephotetticum. Lectotypes are designated for Japania andoi and Oligosita shibuyae Ishii.     

Parasitism,longevity and progeny production of six indigenous Kenyan trichogrammatid egg parasitoids (Hymenoptera: Trichogrammatidae) at different temperature and relative humidity regimes

Six native Kenyan species/strains of Trichogramma and Trichogrammatoidea, recovered from Helicoverpa armigera were evaluated at six different temperatures (10, 15, 20, 25, 30 and 35°C) and two relative humidity levels (40–50 and 70–80%) with the aim of selecting strains adapted to warmer temperature regimes. The species/strains were collected from low (<700 m), medium (between 700 and 1200 m) and high altitude (>1200 m) locations and were evaluated for parasitism, adult longevity, progeny production and progeny sex ratio at the different environmental regimes. Eggs of the factitious host, Corcyra cephalonica Stainton (Lepidoptera: Pyralidae) were used in the investigations. Temperature and humidity interactions affected parasitism and progeny production. The highest parasitism at the two humidity levels was at 25 and 30°C for all the strains evaluated. Adult longevity was also significantly affected by the interaction of temperature and relative humidity and was longer at the lower than higher relative humidity. Survival followed a type I survivorship curve at lower temperatures and a type III survivorship curve at the higher temperatures. Trichogramma sp. nr. mwanzai from low altitude, Trichogramma sp. nr. mwanzai from medium altitude and Trichogrammatoidea sp. nr. lutea also from medium altitude lived longer than other strains at all the temperatures and relative humidity levels evaluated, including the warmest regimes of 30 and 35°C. These strains appear promising as candidates for augmentative biocontrol of H. armigera in Kenya.     

Description and behavioural biology of two Ufens species (Hymenoptera: Trichogrammatidae), egg parasitoids of Homalodisca species (Hemiptera: Cicadellidae) in southern California

The morphology, distribution, and hosts of two egg parasitoids, Ufens principalis Owen sp. n. and U. ceratus Owen sp. n. (Hymenoptera: Trichogrammatidae), are described for the first time. These species are compared to U. niger (Ashmead), the only species of Ufens s. str. currently described from the Nearctic, and diagnostic differences are presented. The behavioural biology of U. principalis, and U. ceratus to a smaller extent, is also described for the first time. Ufens principalis exhibited a rapid and long-distance response in the form of directional flight toward freshly laid eggs of Homalodisca species, its primary hosts in southern California. Parasitism involved aggregations of female U. principalis on fresh Homalodisca egg masses, which remained attractive to U. principalis for a relatively short time. The level of oviposition by U. principalis females was low during most of the day and peaked before sunset in tandem with a peak in Homalodisca oviposition. Oviposition behaviour of U. principalis is described and the distribution of ovipositor probe durations showed that most probes were generally of very short duration. Mating of both Ufens species occurred on the egg mass, with males showing aggressive behaviour towards each other as they competed for emerging females. Ufens ceratus males displayed greater aggression towards other males than U. principalis males. By contrast, fights among U. principalis males involved more individuals and lasted longer than corresponding fights between U. ceratus males.     

Egg parasitoids (Hymenoptera: Mymaridae and Trichogrammatidae) of Kolla paulula (Walker) (Hemiptera: Cicadellidae) in Taiwan

Pseudoligosita nephotetticum (Mani) (Hymenoptera: Trichogrammatidae) and Gonatocerus (Cosmocomoidea) sp. (Hymenoptera: Mymaridae) have been identified for the first time as egg parasitoids of the sharpshooter leafhopper Kolla paulula (Walker) (Hemiptera: Cicadellidae: Cicadellinae: Cicadellini) in Taiwan. K. paulula has been recently identified as a candidate vector of the phytopathogenic bacterium Xylella fastidiosa, the causative agent of Pierce's disease of grapes and similar diseases of other affected plants in Taiwan, and thus has become of particular economic importance. Also provided is a summary of the known records of egg parasitoids (Mymaridae and Trichogrammatidae) of other leafhoppers from the tribe Cicadellini in the world, as well as taxonomic notes on P. nephotetticum.     

Effect of pesticides on Colpoclypeus florus (Hymenoptera: Eulophidae) and Trichogramma platneri (Hymenoptera: Trichogrammatidae), parasitoids of leafrollers in Washington.

Pesticides were evaluated for their effect on two parasitoid species, Colpoclypeus flouts and Trichogramma platneri, that are potential biological control agents of leafrollers in apple orchards. Organophosphate and carbamate insecticides were highly toxic to both parasitoids in topical applications, but foliar residues of some products were nontoxic after 7 d. At reduced rates, topically applied pyrethroids were low in toxicity to C. florus were highly toxic to T. platneri, and foliar residues were nontoxic after about 7 d. Imidacloprid and abamectin were highly toxic when applied topically to both parasitoids but were not toxic as 1-d-old residues. Insect growth regulators did not cause mortality either as topical applications or residues; however, diflubenzuron caused severe sublethal effects, completely blocking the production of C. florus offspring. Biorational pesticides, such as soap, oil, and B. thuringiensis products, caused no toxicity to C. florus but had a direct impact on T. platneri as topical applications through physical immobilization. The potential to integrate different pesticides with biological control of leafrollers and the need for a step-wise approach to evaluate the impact of pesticides against natural enemies is discussed.     

Costs and benefits of polyandry in the egg parasitoid Trichogramma evanescens Westwood (Hymenoptera: Trichogrammatidae)

Polyandry implies costs (i.e., time, energy, predation risk, etc.) especially in short-lived parasitoid species but females of several hymenopteran parasitoid species, mostly gregarious, do mate with multiple males. Several hypotheses have been proposed to explain the benefits of polyandry but controversy remains, especially in facultative gregarious species that bridge the gap between solitary and gregarious development. In this study, we investigated the possibility that polyandry may bring material benefits to Trichogramma evanescens Westwood (Hymenoptera: Trichogrammatidae) females, a short-lived and facultative gregarious egg parasitoid. Females mated several times with different males both at emergence and throughout their life. No significant difference was found in the offspring sex ratio and the fecundity of multiple mated and single mated females and pre-mating duration increased with the female’s age. The longevity of females did vary significantly with the number of matings but only in the presence of hosts. Female T. evanescens received enough sperm from one mating to allocate an optimal offspring sex ratio and we found no evidence of either nutritional resources or convenience polyandry in this species. Polyandry in facultative gregarious parasitoids might be an adaptive strategy to minimize the risk of mating with males that have already emptied their sperm bank or to accumulate sperm from several partially sperm-depleted males. Polyandry may also increase the probability of non-sib mating in patches exploited by several females.     

Effect of temperature and host species on parasitism,development time and sex ratio of the egg parasitoid Trichogrammatoidea lutea Girault (Hymenoptera: Trichogrammatidae)

The developmental biology of Trichogrammatoidea lutea Girault (Hymenoptera: Trichogrammatidae) was studied at six constant temperatures (18, 21, 24, 27, 30 and 35 °C) on eggs of three lepidopteran host species: Helicoverpa armigera (Hübner) (Noctuidae), Chilo partellus (Swinhoe) (Crambidae) and Cadra cautella (Walker) (Pyralidae). T. lutea did not complete development at 35 °C on any of the three host species. Parasitism levels were highest on H. armigera at 27 °C (58%), C. cautella at 27 and 30 °C (31% and 28%) and C. partellus between 24 and 30 °C (13–17%). Realized progeny of T. lutea per parasitized host egg was influenced by host size. The number of progeny of T. lutea per parasitized host egg was highest on H. armigera, followed by C. partellus and lowest on C. cautella. The sex ratio was female biased on C. partellus, female biased on C. cautella with the exception of 21 °C and close to 1:1 on H. armigera. The rate of development from egg to pupa and egg to adult was fastest on H. armigera and slowest on C. partellus. Lower thresholds for development and degree days (DD) of T. lutea from egg to adult were 12.8 °C and 105.4 DD on H. armigera, 11.3 °C and 141.6 DD on C. partellus and 12.9 °C and 118.2 DD on C. cautella, respectively. Based on these results, H. armigera is the most suitable host for mass rearing of T. lutea for biological control of Lepidoptera pests because of the relatively high parasitism levels, short development time, greater clutch size and balanced sex ratio. C. cautella may also be used although longer exposure times might be required due to lower parasitism levels.     

Egg parasitoids (Hymenoptera: Mymaridae and Trichogrammatidae) of okra leafhopper,Amrasca biguttula (Ishida) (Hemiptera: Cicadellidae), in southern Japan

The okra leafhopper, Amrasca (Sundapteryx) biguttula (Ishida) (Hemiptera: Cicadellidae), is an economically important pest of okra, Abelmoschus esculentus (L.) Moench (Malvaceae), in Japan. During 2019 and 2022 surveys for its egg parasitoids (Hymenoptera) were conducted on Kyushu and Ryukyu islands in southern Japan. Five species of Mymaridae, Anagrus (Anagrus)?atomus (L.), A. (A.) japonicus Sahad, A. (A.) turpanicus Triapitsyn and Hu, Arescon enocki (Subba Rao and Kaur), and Stethynium empoascae Subba Rao, as well as one of Trichogrammatidae, Pseudoligosita nephotetticum (Mani), were recorded from eggs of A. biguttula on okra in Japan. Of these, A. (A.) turpanicus, identified using a combination of molecular and morphological methods, is newly recorded from Japan; A. (A.)?atomus, A. (A.) turpanicus and P. nephotetticum are for the first time reported from eggs of okra leafhopper.     

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.     

Taxonomy of the genusTrichogramma (Hymenoptera,Chalcidoidea, Trichogrammatidae)

Résumé Dans le genreTrichogramma, il existe d'une part un certain nombre d'espèces et, d'autre part, beaucoup de formes biologiques caractérisées seulement par leur comportement. Les espèces bien distinctes par leur morphologie sont:T. evanescens Westwood,T. embryophagum (Hartig),T. semblidis (Aur.,T. minutum Riley,T. japonicum Ash. etT. retorridum (Gir.). La valeur d'une espèce deTrichogramma est établie par le mode de reproduction, la variation de la coloration, la durée du cycle évolutif et les différences des soies des antennes chez les males, et tout cela pour une température haute et constante (30°C). La connaissance des modifications physiologiques et des changements de coloration d'une espèce en fonction de l'habitat est importante pour les études taxonomiques qu'il faut exécuter avec un matériel vivant. La tache d'un taxonomiste deTrichogramma est biologiquement difficile, mais son importance pour l'agriculture est considérable.      

Cold storage of the egg parasitoids Trissolcus basalis (Wollaston) and Telenomus podisi Ashmead (Hymenoptera: Scelionidae)

Adults of Trissolcus basalis and Telenomus podisi were stored either at 15 or 18 °C after their immature development had been completed at 18 or 25 °C. Longevity of the parasitoids in the storage temperatures was evaluated, as well as fecundity and longevity following their return to 25 °C after different periods in reproductive diapause. Temperature during immature development influenced female longevity and highest mean longevity was obtained for females that developed to the adult stage at 25 °C and then were stored at 15 °C (ca. 13 months for T. basalis and 10 months for Te. podisi). For adults of T. basalis that developed at 25 °C, storage periods of 120 or 180 days at 15 or 18 °C did not affect fecundity. The fecundity of T. basalis females that developed at 18 °C and were stored for 120 days at 15 or 18 °C was not affected; however, after remaining for 180 days, fecundity was reduced in ca. 30 and 50%, respectively. Storage of Te. podisi adults at 15 or 18 °C significantly reduced fecundity. It is concluded that adults of T. basalis can be stored in the adult stage at 15 or 18 °C between two soybean crop seasons for mass production purposes, aiming the biological control of stink bugs.     

Trichogramma zahiri (Hymenoptera: Trichogrammatidae) an egg parasitoid of the rice hispa Dicladispa armigera (Coleoptera: Chrysomelidae) in Bangladesh

Trichogramma zahiri Polaszek sp. n. is described from Bangladesh. It has been collected at several localities from the major pest of rice Dicladispa armigera (Olivier) on which it has an important controlling impact. Trichogramma zahiri is compared with known Trichogramma species from the region, and diagnostic differences are presented. Data on developmental period, adult longevity, egg-laying frequency, host egg age preference and seasonal parasitism rate are presented and discussed. Other records of parasitoids of D. armigera are briefly reviewed and discussed.     

Unisexual broods and sex ratios in a polyembryonic encyrtid parasitoid (Copidosoma sp.: Hymenoptera)

Summary We examined brood sex ratios of an undetermined species of Copidosoma. Most broods (65%) were unisexual, with a greater proportion of female broods. Some mixed broods contained extremely small proportions of wasps of the opposite sex. Our results imply that the female-biased sex rations in this species cannot be explained by local mate competition theory.