The reproductive strategy of the gregarious parasitoid,Pteromalus puparum (Hymenoptera: Pteromalidae)

Summary Host size of Pteromalus puparum, a gregarious pupal parasitoid, shows a wide inter- and intraspecific variation. Experiments were made to study the regulation of the number and sex ratio of progeny per host by the parasitoid. The parasitoid could discriminate inter- and intraspecific size differences of the host and regulate the number of eggs according to the host size when a single female attacked the host. The sex ratio of progeny (proportion males) was about 0.1. The number of progeny laid by the female agreed with the energetically most efficient number og eggs in order to maximize total weight of progeny per host but not with the reproductively most efficient number of eggs to maximize the total fecundity of the progeny. The parasitoid laid smaller number of eggs in a half buried host, but the number was much larger than a half of those in a fully exposed host. When more than one female attacked a single host, the number and sex ratio of progeny per host increased with the number of females attacking the host, but the number of progeny per female decreased. The change of the sex ratio agreed with the prediction of the local mate competition model.     

Fecundity and oviposition ofEucelatoria bryani, a gregarious parasitoid ofHelicoverpa zea andHeliothis virescens

We examined longevity, fecundity, and oviposition strategies ofEucelatoria bryani Sabrosky (Diptera: Tachinidae), a gregarious endoparasitoid ofHelicoverpa zea (Boddie) andHeliothis virescens (F.) (Lepidoptera: Noctuidae). Longevity of adult femaleE. bryani was not related to body size. In contrast to longevity, largerE. bryani females had greater potential fecundity than smaller females, as determined by the number of embryonated eggs present in the common oviduct. However, female parasitoid size did not affect primary clutch size (number of eggs deposited in a host). Because embryos in eggs located in the ovisac were larger than those located elsewhere in the common oviduct, maximum primary clutch size may be physiologically limited by the number of fully mature eggs a female has available at one time.E. bryani females adjusted primary clutch size in response to host size, for bothH. zea andH. virescens. This adjustment appears to be adaptive because females did not overexploit hosts by depositing more larvae than a host could support. Adult emergence was not related to host size. Although host weight positively influencedE. bryani progeny weight, increases in progeny size with host size were counterbalanced by increases in primary clutch size with host size.     

The reproductive strategy of the gregarious parasitoid,Pteromalus puparum (Hymenoptera: Pteromalidae)

Summary Pteromalus puparum is a gregarious parasitoid of lepidopterous pupae. To determine in which phase of a host outbreak superparasitism occurs, field investigations were carried out on seasonal prevalence of the host, Papilio xuhtus, and parasitism by P. puparum in a citrus grove in Fukuoka, Japan in 1974. Host pupation occurred from May to November and the parasitoid attacked the host throughout this period. Pupal density increased rapidly after mid-August and the percentage parasitism decreased during this period. A high level of parasitism was attained after one or more parasitoid generations in mid-September. The superparasitism was observed after mid-September when the parasitoid attained extremely high density, and resulted in an increase in the proportion of males, high mortality, and a decrease in the size of the progeny.     

Reproductive performance of Nesolynx thymus (Hymenoptera: Eulophidae), in relation to age of Musca domestica (Diptera: Muscidae)

An investigation was undertaken to record the influence of host age on the reproductive performance of Nesolynx thymus (Hymenoptera:Eulophidae). This is an indigenous, gregarious, ecto-pupal parasitoid of certain dipteran insects, including the tachinid fly, Exorista bombycis (Louis) which is a well-known endo-larval parasitoid of the silkworm, Bombyx mori. Each gravid N. thymus female was allowed to parasitise 1–4-day-old puparia of Musca domestica L. for 2 days at a parasitoid–host ratio of 1:20. The parasitised host puparia were observed for progeny recovery, sex ratio, female longevity and fitness (adult size). In addition, reproductive performance of the parasitoid progeny was assessed by allowing its females to parasitise for 2 days, 3-day-old puparia of E. bombycis at a parasitoid–host ratio of 1:4. There was a significant negative correlation between host age and parasitisation rate, parasitoid developmental duration, sex ratio and female longevity while the correlation was positive between host age and parasitoid recovery per host puparium. Similarly, negative correlation was observed between host age and morphometric parameters (body length, head width and wing span of male and female and length and width of female abdomen) of the progeny adults. Host age did not have any impact on the reproductive performance of progeny adults when allowed to parasitise puparia of E. bombycis.     

Modelling age‐ and density‐structured reproductive biology and seasonal survival of Apanteles galleriae Wilkinson (Hym., Braconidae)

Abstract: The biology of Apanteles galleriae Wilkinson, an important biological control agent of wax moths, is well described in the literature. We developed models simulating the functional response of fecundity and female progeny proportion of adult females as a function of age, host and/or parasitoid density to integrate current knowledge. Daily pattern of age‐related fecundity and female progeny proportion of different parasitoids was also examined. We investigated the effect of sex, mating status and seasonal time on adult longevity. We derived survival data of females in two different seasonal periods. We found that 50% of adult life span is important for an efficient fecundity. The patterns of host and/or parasitoid density‐dependent fecundity and sex ratio varied considerably. The highest fecundity and female progeny proportion occurred with one parasitoid and one female equivalent host. Longevity of adults was affected by sex and month periods, but mating status did not affect their longevity. Female survival was greater between December and May relative to June and November. Our results indicated that age, host and parasitoid density, and the timing of rearing influenced the life processes of parasitoids.     

Effects of host age and host availability on developmental period,adult size,sex ratio,longevity and fecundity inLariophagus distinguendus Förster (Hymenoptera: Pteromalidae)

The effects of host age on parasitoid reproductive capacity are studied using the pteromalid parasitoid Lariophagus distinguendusFörster and its bruchid hosts, Callosobruchus chinensis (L.) and C. maculatus (F.). A series of experiments were performed to investigate relationships between age and size of host parasitized and the developmental period of pre-imaginal progeny, sex ratio, female size, longevity, fecundity and oviposition rate. There was no effect of host size on preimaginal parasitoid developmental period. Sex ratio varied from less than 5% females from young (small) hosts to 60% females from mature (large) hosts. Adult size, female longevity, fecundity, and oviposition rate were also positively related to host age. Females provided mature hosts lived longer than those provided either young hosts or no hosts, possibly because of an increased ability to host-feed from the larger hosts. The implications of these findings to parasitoid population reproductive capacity and host-parasitoid synchrony are discussed.     

Asymmetric size effect of sexes on reproductive fitness in an aphid parasitoid Aphidius ervi (Hymenoptera: Aphidiidae)

Most studies on size–fitness relationships focus on females and neglect males. Here, we investigated how body size of both sexes of an aphid parasitoid, Aphidius ervi Haliday, affected the reproductive fitness. Reproductive fitness was generally positively correlated with body size for both sexes in this species. Large individuals of both sexes had greater longevity, large males fathered more progeny, and large females had higher fecundity, parasitism, and greater ability in host searching and handling. We demonstrated in this study that size effects of males and females were asymmetric on different reproductive fitness parameters. With increasing body size females gained more than males in longevity and fecundity while males gained more than females in the number of female progeny. Regardless of female size, large males sustained a female-biased population longer than small males. These results suggest that male body size should also be considered in the quality control of mass-rearing programs and the evaluation of parasitoid population growth.     

Biology of Australian and United States strains of Trissolcus basalis,a parasitoid of the green vegetable bug,Nezara viridula

Strains of Trissolcus basalis (Wollaston) Hymenoptera: Scelionidae, an egg parasitoid of the green vegetable bug, Nezara viridula (L.), were obtained from three regions of Australia and from Homestead, Florida (USA). Percent parasitoid emergence was approximately the same for freezer-stored (-75°C) and fresh eggs, and freezer-stored eggs were suitable hosts for a longer time. Optimum host age, oviposition and emergence patterns, mean number of progeny per female, sex ratio and longevity were examined and certain comparisons were made between the Australian and USA parasitoids. Genetic crosses made between males and females of each strain revealed that reproductive isolation did not occur. Higher fecundity of the Australian strains may make them more successful than indigenous (USA) strains for controlling pest populations of N. viridula in the southern USA.     

Reproductive performance of <Emphasis Type="Italic">Nesolynx thymus</Emphasis> (Hymenoptera: Eulophidae) as influenced by host (<Emphasis Type="Italic">Musca domestica</Emphasis>) size

An investigation was conducted to understand the influence of host size on the reproductive performance of Nesolynx thymus, an ecto-pupal parasitoid of the uzi fly, Exorista bombycis, inflicting 10–20% loss to silkworm crops in the southern silk producing states of India. Two day-old adults of N. thymus were allowed to parasitize pupae of M. domestica, categorized into big and small at a parasitoid-host ratio of 1:20. After parasitoid adult eclosion, data on parasitization rate, developmental duration, male, female and total adult recovery per pupa and per female, sex ratio, adult size and adult longevity were recorded. The performance of first generation adults as influenced by host size was also recorded by allowing the first generation adults to parasitize three day-old pupae of E. bombycis. The parameters female and total progeny production per female, sex ratio, longevity, body length and wing span of males, females, head width of male, and abdomen length of female increased significantly with host size. No impact of host size was observed on performance of first generation adults.     

Reproductive attributes of the eulophid Oomyzus sokolowskii,a biological control agent of diamondback moth,Plutella xylostella (Lepidoptera: Plutellidae)

Oomyzus sokolowskii (Kurdjumov) is a gregarious larval-pupal parasitoid of the diamondback moth, Plutella xylostella L. Under laboratory conditions, we explored the reproductive attributes including the functional response of O. sokolowskii and the impact of mutual interference among female parasitoids on progeny production and progeny sex ratio. In the functional response experiment, groups of 1–50 late instar larvae were exposed to groups of five female parasitoids for 48 h. Both Holling's disc equation (type II response) and Williams and Martinez's model (type III response) fit significantly better than the null-model assuming density-independent parasitation efficiency. Williams and Martinez's model was only marginally better than Holling's disc equation which explained 82% of the variation in the number of parasitized hosts. According to Holling's disc equation, the estimated maximum number of hosts parasitised during 48 h was 17 per group of five parasitoids, the estimated attack efficiency of a single female parasitoid (a) was 0.0024 larvae per hour, and the estimated handling time of a single female (T _h) was 14.4 h per host. In the mutual interference experiment 50 host larvae were exposed to different numbers of female parasitoids (P=5, 10, 20, 30, 40 and 50) for 48 h. With increasing numbers of conspecifics, the proportion of male progeny increased and total number of progeny per parasitoid decreased, yielding an estimated mutual interference constant (m) (±SE) of 0.95±0.14. The total number of wasps emerging from individual host pupae increased with increasing number of conspecifics. The age-specific fecundity of O. sokolowskii was investigated by providing 10 host larvae daily to individual female parasitoids. After 3 days of oviposition, 90% of the females were still alive and had achieved 76.4% of their lifetime fecundity. The proportion of female progeny decreased with female age. No relationship was found between female longevity and lifetime fecundity. The results suggest that parasitoid and host densities influence the progeny production and sex allocation strategy of O. sokolowskii. The information obtained from this study would help to develop a mass-rearing protocol for O. sokolowskii.     

Costs and benefits of host feeding in the parasitoid wasp Trichogramma turkestanica

Females of the parasitoid wasp Trichogramma turkestanica Meyer (Hymenoptera: Trichogrammatidae) generally host feed after ovipositing on the first egg of Ephestia kuehniella Zeller (Lepidoptera: Pyralidae) they encounter. We measured the impact of host feeding on the fecundity and longevity of females, in absence of host or food, and on the fitness of their progeny. We also determined if the frequency of host feeding is influenced by the humidity level at which T. turkestanica females developed. Host feeding increased egg production by 70% but decreased female longevity. This impact of host feeding on the longevity of females is probably due to the allocation of carbohydrates to egg production at the expense of somatic maintenance. Humidity did not influence the occurrence or duration of host feeding. The size of individuals developing in eggs on which females host fed was smaller, indicating that their fitness was affected.     

Reproductive biology of the microtype tachinid fly Zenillia dolosa (Meigen) (Diptera: Tachinidae)

Reproductive biology including mating, adult longevity, fecundity and development of the tachinid fly Zenillia dolosa was investigated for optimizing rearing procedures using Mythimna separata as a host in the laboratory. Females lay microtype eggs containing a first instar larva on food plants of the host and then the eggs must be ingested by the host for parasitization. Mating success was 58.5% with mating duration of 80.7 min. Mating was most successful when day 0–1 females were kept with day 2–4 male flies. Female body size was positively correlated with its fecundity but not with longevity. However, females that survived longer produced more eggs during their lifetime. Parasitoids successfully developed in 4th to 6th instar host larvae. Host instars at the time of parasitoid egg ingestion significantly influenced development time of the immature parasitoid, but did not affect body size of the emerging parasitoid. We suggest that pairing newly emerged females with day 2–4 males should result in higher mating success and using the last instar hosts for parasitization should minimize development time of the parasitoid for rearing.     

Body size as an indicator of parasitoid quality in male and female Anagyrus kamali (Hymenoptera: Encyrtidae)

The parasitoid Anagyrus kamali Moursi was recently introduced into the Caribbean as a biological control agent against the hibiscus mealybug, Maconellicoccus hirsutus Green. In the laboratory, parasitoid size, as measured by left hind tibia length, was positively correlated with several indicators of the parasitoid's fitness: longevity, mating preference, fecundity, reproductive longevity, progeny emergence and sex-ratio. When fed ad libidum with honey drops, large male parasitoids lived significantly longer (29.1 +/- 6.5 days) than small ones (18.4 +/- 5.7 days). Large females also lived significantly longer (35.4 +/- 10 days) than small females (27.9 +/- 9.6 days). Females showed no significant mating preference between large and small males. Lifetime fecundity was positively correlated with the size of adult females and ranged from 37 +/- 21 eggs for small females to 96 +/- 43 eggs for large ones. The reproductive longevity, daily oviposition rate, and number of progeny were also higher among large parasitoids. The sex ratio of progeny from small female parasitoids was higher (0.76 +/- 0.24) than that of large individuals (0.47 +/- 0.18).     

Effect of mating status and age of Microplitis rufiventris (Hym., Braconidae) females on the growth pattern and number of their teratocytes

The effect of mating status and age of the female of Microplitis rufiventris parasitoid on the growth pattern of its teratocytes during the larval development is described as well as the changes in the number of these cells. The growth pattern of teratocytes derived from haploid eggs significantly differed from that of eggs deposited by mated females. Both cell diameter and the number of maturing M. rufiventris teratocytes was fertilization-dependent. Following the parasitization of Spodoptera littoralis larvae with virgin parasitoid females, the host larvae hosted a large number of cells of smaller size at the end of parasitoid development. The opposite effect was seen when the host larvae contained female parasitoid larvae. The age of the female at parasitization had an effect on the maximum cell size attained prior to parasitoid emergence. It seems that there are two factors other than mating which affect the number of teratocytes: degeneration inside the host haemolymph and to some degree accidental ingestion by parasitoid larva.     

Effects of rearing conditions on reproduction of Spathius agrili (Hymenoptera: Braconidae), a parasitoid of the emerald ash borer (Coleoptera: Buprestidae)

Spathius agrili Yang (Hymenoptera: Braconidae) can be successfully reared on emerald ash borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), larvae feeding in chambers drilled in small ash twigs that are wrapped with floral tape. Females maintained in groups with males for one week can receive enough sperm for production of female progeny throughout their lives. Volatiles released by emerald ash borer adults feeding on ash foliage increased parasitoid fecundity over ash foliage alone or no stimulus. The temperature at which the parasitoids were reared ranged from 20 to 25 degrees C in a daily cycle; however, raising the daily maximum temperature to 28 degrees C did not affect parasitoid longevity or fecundity. Adult females lived between 12 and 127 d, with an average of 60.8 +/- 4.5 d. Males lived slightly longer, with an average of 66 +/- 4.5 d. The first clutch of eggs was laid when the female was between 2 and 42 d old, with the average preoviposition period lasting 11.4 +/- 1.4 or 19.5 +/- 2.0 d in 2007 and 2009 trials, respectively. A higher proportion of the emerald ash borer larvae were feeding and thus attractive to parasitoids in the 2009 trial, and female S. agrili laid an average of 9.5 +/- 1.0 clutches containing 5.4 +/- 0.2 eggs, for an average of 51.2 eggs per female. Approximately three quarters of the progeny were female. The number of eggs per clutch was significantly greater when deposited on larger emerald ash borer larvae, further highlighting the need for quality larvae in rearing. Chilling S. agrili pupae at 10 degrees C to stockpile them for summer release was not successful; chilling resulted in lower survival and lower fecundity of emerging progeny. Female S. agrili proved capable of attacking emerald ash borer larvae through even the thickest bark of an ash tree that was 30-cm diameter at breast height. Even emerald ash borer larvae that were creating overwintering chambers in the outer sapwood of the tree were successfully attacked, suggesting that S. agrili could be reared on field collected logs infested with emerald ash borer.     

Sex ratio regulation in Coccygomimus turionella Linnaeus (Hymenoptera: Ichneumonidae) and its ecological implications

Abstract. 1. Sex regulation and its relevance to the ecology of a k-selected pupal parasitoid was investigated in Coccygomimus turionellae.
2. The ratio varied with host size, females predominating in large hosts.
3. Egg transfer experiments and comparative mortality rates among the progeny of virgin and inseminated females demonstrated that the phenomenon was due to parental behaviour.
4. The study of the mechanism confirmed Aubert's hypothesis of host size estimation with the addition that the host had to be exposed from the poles and had to contain the proper host kairomones.
5. Host size was highly correlated to parasitoid fecundity via influences on size and longevity but not the number of ovarioles per female or daily egg production.
6. Host encounters are more limiting than egg production and host size had no effect on host acceptance.
7. Sex regulation allows a maximal host encounter rate at the same time that it maximizes the reproductive potential of female progeny.
8. Low host density increased the production of female progeny resulting in more offspring searching for the limited resource. Concealed hosts and a high host density resulted in a shift towards a Fisher 1:1 sex ratio.     

Biology of a thelyotokous biotype of Eretmocerus mundus (Hymenoptera: Aphelinidae) on Bemisia tabaci (Homoptera: Aleyrodidae)

Abstract Thelyotokous biotype of Eretmocerus mundus Mercet (Hymenoptera: Aphelinidae), a parasitoid of Bemisia tabaci (Gennadius) (Homoptera: Aleyrodidae), was recently recorded in northern Iran. Reproductive biology of this biotype was studied as part of an evaluation of its potential for biological control of B. tabaci. The parasitoid deposited more eggs under 2nd and 3rd nymphal instars than 1st or 4th instars. Adult females fed honey, with no access to whitefly nymphs, lived significantly longer (13.6 ± 4.7 d) than those given access to nymphs, but not fed honey (7.6 ± 2.21 d). Lifetime fecundity averaged 81.7 ± 26.9 female progeny per female parasitoid, ranging from 11–132. Daily fecundity, measured as the number of whitefly nymphs parasitized by per female each day for 10 d, averaged 18.06 ± 3.95 for the first 6 d of life, and then declined to < 11. Developmental time from oviposition to parasitoid emergence was significantly shorter in the 3rd instar of the host (15.9 ± 1.06 d) than in the 1st instar (18.7 ± 2.3 d), but not in the 2nd instar (16.4 ± 1.3 d).     

Factors affecting the bionomics of the eastern African egg parasitoid Trichogramma bournieri Pintureau & Babault (Hymenoptera: Trichogrammatidae)

The trichogrammatid Trichogramma bournieri Pintureau & Babault is a polyphagous parasitoid of eggs of several cereal stemborer species in eastern Africa. The effects of host species, host age and duration of host deprivation on the performance of the parasitoid were studied in the laboratory. Host acceptance and suitability were tested using five stemborer species. The noctuids Sesamia calamistis Hampson, Sesamia nonagrioides (Lefebvre), Busseola fusca (Fuller) the crambid Chilo partellus (Swinhoe) and the pyralid Eldana saccharina Walker were successfully parasitized by T. bournieri. Parasitism, number of progeny and developmental time varied significantly with host species. The eggs of S. calamistis and B. fusca were the most suitable, whereas those of E. saccharina were the least suitable. While parasitism and number of progeny tended to decrease with age of hosts, there were no significant differences in sex ratio. Longevity of the parasitoid increased with increase in deprivation of hosts from 0 to 12 days. Average lifetime fecundity per female decreased, indicating resorption of eggs.     

Intraspecific and Interspecific Competition Between Cotesia flavipes and Cotesia sesamiae (Hymenoptera: Braconidae), Gregarious Larval Endoparasitoids of Lepidopteran Stemborers

Super and multiple parasitism of Chilo partellus (Lepidoptera: Pyralidae) and Sesamia calamistis (Lepidoptera: Noctuidae) by Cotesia flavipes Cameron and Cotesia sesamiae (Cameron) (Hymenoptera: Braconidae) were investigated in the laboratory. Progeny production of Co. flavipes increased as a result of increasing the number of ovipositions, from one to three per one Ch. partellus host larva, then decreased as a result of four and five ovipositions per larva. Cocoon weight, sex ratio and emergence of the parasitoid progeny were not affected by superparasitism. Low progeny production of Co. sesamiae and poor survival of Ch. partellus host larvae were found as a result of superparasitism. When S. calamistis was the host, the duration of immature stages of Co. flavipes , parasitoid emergence, progeny production and sex ratio were not affected by superparasitism, but cocoon weight, adult longevity and the potential fecundity of adult females decreased. Superparasitism of S. calamistis by Co. sesamiae did not affect emergence, longevity or sex ratio of adult progeny of the parasitoid, but prolonged immature development, lowered cocoon weight and decreased potential fecundity of adult female progeny. Co. flavipes out-competed Co. sesamiae when Ch. partellus was parasitized by both species. The potential for local displacement of Co. sesamiae by Co. flavipes in areas dominated by Ch. partellus in East Africa is discussed.     

Life history parameters of the microtype parasitoid fly Pales pavida (Diptera: Tachinidae), and effects of host age and number of eggs ingested by host on parasitism success

With the aim of developing better procedures for rearing the microtype tachinid fly Pales pavida (Meigen), we performed ecological studies in the laboratory using the natural host Mythimna separata (Walker), investigating larval development, mating behaviour, individual oviposition patterns and relationships between parasitisation and number of eggs ingested (NEI) per host. The host instar at the time of parasitoid egg ingestion significantly affected the development time of the immature parasitoid: development took longer when the hosts ingested eggs when at the fifth instar than at the sixth (last) instar. There was no difficulty obtaining mated females in the laboratory when day 0–1 female flies were kept with day 2–4 males. Mean lifetime fecundity was 5805?±?568 eggs per female. Daily rates of oviposition by individual females varied widely; the greatest number of eggs laid in a day was exactly 1700. When the NEI by day 1 fifth instars or day 0 or 3 last instars was 1, 3, 6 or 10, the parasitisation percentage tended to increase with increasing NEI, although it did not differ significantly between NEI 6 and 10. Therefore, the percentage adult emergence per egg ingested decreased from NEI 6 to 10, particularly in the case of last instars. Using day 0 last instars, with six eggs ingested per host, should increase parasitisation rates and shorten the development time of the parasitoid for rearing.