Parasitism of orangestriped oakworm (Lepidoptera: Saturniidae) eggs in the urban landscape

Orangestriped oakworm, Anisota senatoria (J. E. Smith), has caused widespread defoliation of oak trees in the urban landscape of southeastern Virginia since 1985. Egg masses were collected from 1988 to 1990 to determine the impact of native egg parasites on A. senatoria populations. The most abundant egg parasite was Aprostocetus new sp. and mean egg mass parasitism was 24.6%. The eupelmid Anastatus hirtus (Ashmead), a new host record, parasitized a mean of 11.7% of A. senatoria egg masses. The encyrtid Ooencytrus sp., a new host record, had a mean egg mass parasitism of 0.09%. Inundative releases of Trichogramma minutum (Riley) in 1989 and 1990 did not increase parasitism rates and mean egg mass parasitism was 2.3%. Parasitism of first generation A. senatoria egg masses was higher compared with second generation. The four egg parasites collected in this study parasitized 30% of A. senatoria egg masses and within egg mass parasitism was 7.9%. These relatively low parasitism rates may partially explain the presence of consistently high A. senatoria populations in southeastern Virginia.     

Parasitism of cotton leafworm Alabama argillacea eggs by Trichogramma pretiosum in commercial cotton fields

Release of natural enemies in commercial fields is challenging and has been inconsistent in the results achieved. This work discusses the augmentative releases of Trichogramma pretiosum to control the cotton leafworm (CLW) Alabama argillacea and also examines the parasitoid–host interaction under grower field conditions. The treatments consisted of fields with and without releases of T. pretiosum set up in Primavera do Leste and Campo Verde Counties, MT, Brazil, during three different seasons (2003 and 2004 dry and 2004 regular summer seasons). Trichogramma wasps were weekly released in the treated fields throughout the entire sampling period (14–15 week period) at a rate of 100 000 wasps per hectare. One‐way repeated measures analysis of variance was performed for the number of parasitized eggs, followed by a meta‐analysis procedure to determine the contribution of T. pretiosum release on overall parasitism. In addition, regression analysis was conducted with each season’s data sets to study the relationship of the host density and parasitism response by T. pretiosum. The overall results of Trichogramma augmentative releases did not result in significant increase of CLW egg parasitism beyond the natural parasitism in the areas studied. However, based on Cohen’s d effect sizes from the meta‐analysis, the parasitism rate was greater in fields under T. pretiosum releases during four out of 15 weeks surveyed. The parasitism of CLW eggs by T. pretiosum exhibited host density‐dependence only in one out of three seasons surveyed. These findings are encouraging as they are evidence that T. pretiosum is able to maintain a considerable level of parasitism under commercial field conditions, highlighting their potential value in large‐scale commercial areas of cotton as previously found at the small and diverse farming scale. Future studies should address the potential of early‐season, low density releases of the parasitoid.     

Factors influencing egg parasitism in sub‐social insects: insights from the treehopper Alchisme grossa (Hemiptera,Auchenorrhyncha, Membracidae)

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Dispersal of Trichogramma ostriniae in field corn

Trichogramma ostriniae has shown success as a biological control agent for European corn borer (Ostrinia nubilalis) in sweet corn and the species offers potential for suppression of lepidopteran pests of field corn. Field corn is typically planted at higher densities, is taller, and has greater leaf area than sweet corn, presenting a possible restriction on T. ostriniae dispersal and efficacy. Therefore, parasitoid dispersal in field corn from the centre of a 6.25 ha square grid was determined using sticky cards to capture adult T. ostriniae and sentinel eggs of O. nubilalis to monitor parasitism after releases of ~1 million of T. ostriniae each into four fields of corn. Dispersal was rapid and extensive, achieving distances of ~175 m within 4–7 days after release. The pattern of movement fit well with a diffusion model of dispersal, with the greatest level of dispersal occurring from 7 to 10 days post-release. Parasitism of O. nubilalis sentinel egg masses declined linearly with distance from the release foci, and was also greatest 7–10 days post-release. However, measurement of association showed no significant differences between the spatial distributions of sticky trap captures and percentage parasitism of O. nubilalis egg masses. The distances from the release point that encompassed 98% of re-captured T. ostriniae increased over time and were estimated to range from a low of 100 m at 4 days post-release to 365 m at 14 days post-release. The results of this research suggest that T. ostriniae relies initially on random movement to locate host patches, and that a single release locus per hectare would be sufficient in field corn.     

Does mass rearing of field collected Trichogramma brassicae wasps influence acceptance of European corn borer eggs?

The purpose of our study was to identify a trait that changes quickly during Trichogramma mass rearing, and that could therefore be used to monitor stock deterioration. Quality deterioration in mass reared Trichogramma wasps was evaluated by examining host acceptance behaviour of Trichogramma brassicae Bezdenko (Hymenoptera: Trichogrammatidae) on the target host Ostrinia nubilalis Hübner (Lepidoptera: Crambidae). We compared three replicate lines (designated ‘E’) reared in the laboratory on the factitious host Ephestia kuehniella Zeller (Lepidoptera: Pyralidae) for 27 generations, a line ‘O’ reared in the laboratory on O. nubilalis for 24 generations, and a line ‘F’ reared in the laboratory on O. nubilalis for only two generations. All lines were initiated from field collections of O. nubilalis egg masses. We also evaluated natal host effects by rearing each line on E. kuehniella as well as on O. nubilalis for the last generation prior to testing. The percentage of wasps accepting the O. nubilalis egg mass was significantly higher for the E lines (69.6%) than for the F line (46.5%), while wasps of the O line showed intermediate (57.4%) acceptance. Thus, wasps laboratory reared on E. kuehniella performed better than wasps which had recently been collected in the field. Wasps of the O line showed extended probing behaviour compared to the other lines. Lines did not differ in the duration from the first host contact to the beginning of the drilling, probing or trembling behaviour. Natal host (E. kuehniella or O. nubilalis) did not affect acceptance of the target host O. nubilalis. Even though there is some evidence of adaptation to laboratory rearing conditions, we found no indication for quality deterioration in terms of acceptance behaviour of the target host O. nubilalis when T. brassicae was mass reared on the factitious host E. kuehniella.     

Forest composition, host-population density, and parasitism of spruce budworm Choristoneura fumiferana eggs by Trichogramma minutum

The frequency of egg parasitism in the spruce budworm, Choristoneura fumiferana (Clem.) (Lepidoptera: Tortricidae) by Trichogramma minutum Riley (Hymenoptera: Trichogrammatidae) was recorded in several host populations during the rising phase of an outbreak in the Gatineau Valley (Quebec, Canada) in 1999 and 2000. Some of the observed spruce budworm populations were subjected to experimental aerial applications of Bacillus thuringiensis var. kurstaki during the course of the study. Untreated spruce budworm population densities at the onset of the study were directly related to stand composition (the proportion of non‐host basal area) but unrelated to plant species diversity. Much of the variation in budworm egg mass parasitism rates was accounted for (77.1% of total variance) by three variables: year, spruce budworm egg population density, and proportion of non‐host basal area in the stand. The exploitation of eggs within parasitised egg masses was related to year, B. thuringiensis treatment, egg mass size, and the proportion of egg masses parasitised in the host population (76.9% of variance). The yearly variation in parasitism rate was considerable, and was consistent with climatic conditions, namely heat accumulation and precipitation during the spruce budworm oviposition period. Parasitism by T. minutum on eggs of C. fumiferana was recorded in 16 additional sites in 2000, to confirm the general form of the inverse density dependence of parasitism. Observations recorded by Neilson in the Green River area of New Brunswick (Canada) between 1948 and 1957 are discussed in the context of the present analysis. These results indicate that parasitism by T. minutum may be more predictable than previously believed and may be an important and consistent source of mortality in the several years preceding an outbreak of C. fumiferana, especially in mixed‐wood stands and in warmer, drier portions of the spruce budworm's range.     

Dispersal and field progeny production of Trichogramma species released in an olive orchard in Egypt

Dispersal ability and field progeny production of augmentative released biological control agents depend on ecological adaptations of the particular species or strains used. Four species of the egg parasitoid genus Trichogramma were compared aiming to select suitable candidates for control of lepidopteran olive pests. Three of them (T. bourarachae Pintureau and Babault, T. cordubensis Vargas and Cabello, T. euproctidis Girault) had been previously collected from olive groves, whereas the commercially available strain used (T. evanescens Westwood) was originally isolated from sugarcane fields. During five consecutive field releases in an olive orchard near Cairo, dispersal and/or progeny production of these species was monitored using sentinel eggs placed at different heights in the release tree canopies as well as in neighboring trees (“distance effect”). The cardinal direction of dispersal was random for T. euproctidis and T. evanescens. Significant higher parasitism occurred on sentinel eggs placed on the middle part of tree canopy and highest parasitism was observed in trees where wasps had been released. Field progeny production was highest for T. bourarachae, followed by T. euproctidis and T. cordubensis. T. evanescens propagated less under field conditions. Inter-tree dispersal of all species except T. bourarachae was limited and, for biological control, releasing material should therefore be distributed on each olive tree, preferably also at different levels of the canopy.     

Assessment ofTrichogramma species for biological control of forest lepidopteran defoliators

In a laboratory study, we determined the potential of threeTrichogramma (Hymenoptera: Trichogrammatidae) species,T. brassicae Bezdenko,T. minutum Riley andT. nr.sibiricum Sorokina, for biological control against six species of forest lepidopteran pests, black army cutworm, hemlock looper, eastern spruce budworm, western spruce budworm, white-marked tussock moth, and gypsy moth. Females of each parasitoid species were offered eggs from each of the six host species. Parasitization and the effect of the host species on the emerging progeny were examined and recorded.Trichogramma minutum had the broadest host range and successfully parasitized four host species out of the six offered.Trichogramma nr.sibiricum had the narrowest host range and parasitized only two species of hosts. Of the six host species, black army cutworm was the most preferred by all threeTrichogramma species; white-marked tussock moth and gypsy moth were not parasitized by any parasitoids. There was a positive correlation between the size of female offspring and their corresponding egg complement in all three parasitoid species. The developmental time of parasitoids from egg to adult was influenced by both the parasitoid and host species. Our results suggest thatT. minutum has the greatest potential for biological control against various forest lepidopteran pests and that the black army cutworm may be the best target candidate for further study.     

Parasitism in diversified agroecosystems: Phenology ofTrichogramma minutum [Hymenoptera: Trichogrammatidae]

Parasitism ofOstrinia nubilalis egg masses byTrichogramma minutum was observed in maize-bare ground monocultures and polycultures of maize/bean/squash and maize/clover. Parasitism rates were 1.9 times higher in monocultures than in polycultures; seasonal phenology of parasitism, however, was similar in both. Parasitism first occurred during the late whorl/early tassel stage of maize; peak parasitism occurred during the mid to late tassel stage, and then dropped off rapidly. We speculate that direct predation of egg masses, and ammensal preemptive competition for egg masses byColeomegilla maculata may have contributed to the decline in parasitism rates in both monocultures and polycultures during the latter part of the season.      

Host examination walk and oviposition site selection ofTrichogramma minutum: Studies on spherical hosts

The host examination behavior of the parasitoid wasp Trichogramma minutumon spherical Manduca sextahosts is described. The mean walking speed during host examination was 0.64 ± 0.03 mm s ^–1 and is independent of wasp size. The paths taken by the wasps are not evenly distributed over the host surface; the majority of time is spent below 45 ° latitude. The distribution of oviposition sites is also nonrandom with respect to latitude and is not influenced by phototaxic and geotaxic responses. The initial transit made by the wasps over the host surface is a straight path which frequently passes across the highest point on the host. The length and duration of the initial transit are independent of wasp body length. However, stride length and stepping rates are proportional to wasp body length, and small wasps take more steps to complete their initial transit. The roles of the examination walk in host recognition and host volume measurement by Trichogrammaare also discussed.     

Inoculative releases of Trichogramma ostriniae for suppression of Ostrinia nubilalis (European corn borer) in sweet corn: field biology and population dynamics

The effectiveness of inoculative releases of Trichogramma ostriniae Pang and Chen for suppression of Ostrinia nubilalis (Hübner) in sweet corn was assessed. Early-season, low-density (75,000 females ha⁻¹) releases were made, and establishment, levels of parasitism and sex ratios of emerging T. ostriniae quantified. T. ostriniae established effectively for each season that they were released, but appeared to be unable to overwinter. Parasitism levels tracked egg mass numbers closely, and T. ostriniae persisted in fields even where insecticides were applied. Parasitism by indigenous Trichogramma species was 3%. Field populations of T. ostriniae were distinctly female biased (78%), with males produced in the majority of broods. Numbers of males did not increase linearly with number of O. nubilalis eggs parasitized, but appeared to remain constant above an egg mass size of about 20 eggs. A Type-I functional response to increasing egg and egg mass density was found under field conditions, where the proportion of egg masses parasitized remained constant with increasing egg mass density. A relatively consistent percentage of eggs per egg mass was parasitized, with a linear increase in number of eggs parasitized with increasing number of eggs per egg mass. These results show that T. ostriniae established viable reproductive populations in sweet corn following inoculative release, with the potential to contribute to reduced dependence on insecticides for the control of O. nubilalis in an integrated pest management program.     

Variation in biological parameters of Trichogramma spp. purchased from commercial suppliers in the United States

The efficacy of biological controlobtained through releases of commerciallysupplied Trichogramma is influenced by bothabiotic conditions in the field during andfollowing release and an array of biologicalattributes of the released Trichogramma. Thisstudy was undertaken to assess the degree ofvariation that existed among commerciallysupplied Trichogramma in an array of biologicalparameters that potentially may influence thelevel of control that can be expected in aTrichogramma release program. Shipments ofTrichogramma were obtained from 12 commercialsuppliers of beneficial insects over a 2-yearperiod and the following parameters weremeasured: percentage of emergence duringshipping, total percentage of emergence,percentage of females, percentage ofbrachypterous females, percentage ofbrachypterous males, adult female longevity,and species composition. In addition, weestimated the percentage of macropterousfemales that had not emerged prior to the timethe shipment was received, based on the firstfour parameters. Results indicate a high levelof variation in each of these parameters amongsuppliers of Trichogramma and among shipmentsby individual suppliers. The proportion ofnon-emerged macropterous female wasps was lowfor all suppliers (range 2.5–27.7%). Thiswas primarily attributable to the highincidence of brachyptery among female wasps. However, variation among suppliers in theincidence of emergence during shipping and theproportion of female wasps in a shipment alsocontributed to the low proportion ofnon-emerged macropterous female wasps. Inaddition, in 11 of 24 shipments received duringthe 2-year study, the Trichogramma speciesdiffered from that specified by the shipper orthe shipment contained a mixture ofTrichogramma species.     

Spatio‐temporal variation in Helicoverpa egg parasitism by Trichogramma in a tropical Bt‐transgenic cotton landscape

• 1 Understanding the spatio‐temporal dynamics of insects in agroecosystems is crucial when developing effective management strategies that emphasize the biological control of pests.
• 2 Wild populations of Trichogramma Westwood egg parasitoids are utilized for the biological suppression of the potentially resistant pest species Helicoverpa armigera (Hübner) in Bt‐transgenic cotton Gossypium hirsutum L. crops in the Ord River Irrigation Area (ORIA), Western Australia, Australia.
• 3 Extensive, spatially‐stratified sampling during a season of relatively high Trichogramma abundance found that spatial patterns of pest egg parasitism in the ORIA tend toward heterogeneity, and do not necessarily coincide with host spatio‐temporal dynamics. Both patterns of host egg density and mean rates of parasitism are not good indicators of parasitoid spatio‐temporal dynamics in ORIA cotton crops.
• 4 Parasitism rates can be significantly higher within the middle strata of the cotton plant canopy before complete canopy closure, despite a similar number of host eggs being available elsewhere in the plant.
• 5 Spatial variation in egg parasitism by Trichogramma in Bt‐transgenic cotton is evident at the between‐field, within‐field and within‐plant scale, and is not solely driven by host spatial dynamics. These factors should be considered when estimating Trichogramma impact on pest species during biological control and spatio‐temporal studies of host‐parasitoid interactions in general.

     

Efficacy of inoculative releases of Trichogramma ostriniae (Hymenoptera: Trichogrammatidae) against European corn borer Ostrinia nubilalis (Lepidoptera: Crambidae) in field corn

We evaluated the egg parasitoid Trichogramma ostriniae (Hymenoptera: Trichogrammatidae) to control European corn borer [Lepidoptera: Crambidae: Ostrinia nubilalis (Hübner)] in field corn in 2001 and 2002. Inoculative releases of 75,000 T. ostriniae/ha occurred in New York and Virginia in 5–10 cornfields per state when corn was at mid-whorl. Incidence of egg mass parasitism, number of stalk tunnels, incidence of ear damage, and whole-plant yield were evaluated. Parasitism of European corn borer egg masses ranged from 0 to 75% in release plots and was greater in release plots than in control plots. Individual comparisons between paired release and control plots showed no reductions in either stalk or ear damage. However, when data were combined across both years and fields, stalk and ear damage were significantly reduced in New York. In Virginia, no significant differences were detected using data obtained from one year. There were no differences in yield between release and control plots. Low densities of European corn borer, drought conditions in 1 year, and a larger plant canopy in field corn are possible reasons why T. ostriniae releases provided less control than has been observed in previous trials in sweet corn. Additional research focused on improved timing and frequency and number of releases is warranted.     

Release density,efficiency and disappearance ofTrichogramma nubilale for control of European corn borer

We evaluated how manyTrichogramma nubilale should be released at a single location to controlOstrinia nubilalis in sweet corn. Six 8.6×16 m plots received 18.4 to 2 090 ΦΦT. nubilale/SAI when plants were in the mid to late whorl stage, where SAI, surface area index, is the plant surface area/m². To evaluate the potential control by our releases, we exposed laboratory-rearedO. nubilalis egg masses to the released parasitoids at 4 times after the release. When an egg mass was parasitized byT. nubilale, 75.7% of the eggs in the egg mass were parasitized. We developed an equation to estimate the percent of egg masses that a single female was expected to parasitize in a day (efficiency of parasitism) and female disappearance (death and dispersal) rates, if both were constant during our experiment. The exponential disappearance rate was −0.52±0.03 day⁻¹, which implied that 40% of the remaining ΦΦ disappeared per day. The efficiency of parasitism was 0.050% parasitism/Φ/SAI/day, which implied that at least 351,000 ΦΦ/ha would be needed to achieve 90% parasitism. Clearly, forT. nubilale to be a successful biological control agent, efficiency of parasitism must be increased and disappearance rates must be reduced.      

Importance of plant size, distribution of egg masses, and weather conditions on egg parasitism of the European corn borer, Ostrinia nubilalis by Trichogramma ostriniae in sweet corn

Trichogramma ostriniae (Hym: Trichogrammatidae), an egg parasitoid of the European corn borer, Ostrinia nubilalis (Lep: Pyralidae), were released into sweet corn (Zea mays L.) fields to study the effects of weather, plant size and distribution of egg masses on egg parasitism by the wasp. Sentinel European corn borer eggs were stapled onto leaves located in the upper, middle and lower third of sweet corn plants 5 to 35 meters away from the wasp release point in either a radial or grid manner. Weather conditions and plant architecture were monitored during the experiments. Logistic regression was used to analyze the data. The results indicated that percentage of eggs parasitized was negatively related to an increase in leaf area as well as an increase in distance eggs were located from the point of release of wasps. Eggs distributed on plants at different directions from the release point received different levels of parasitism. Eggs that were stapled onto leaves in the upper third of a corn plant received much less parasitism than those on the middle and lower third of the plant. Higher mean temperature adversely affected the level of parasitism during hotter times of the season and conversely, lower temperatures (<17 °C ) reduced the egg parasitism during cooler times of the season. The longer the exposure of eggs to wasps, the higher the level of egg parasitism. However, the levels of egg parasitism for 2 day's exposure were almost the same as that for 3 day's exposure due to the limited longevity and egg-laying behavior of the wasp. These results suggest that inundative releases of T. ostriniae should be made every two to three days, with multiple release points per hectare. In addition, weather conditions and plant architecture, especially temperature, plant height and leaf area must be taken into consideration to optimize levels of parasitism.     

Side effect of some Neem products on natural enemies of Helicoverpa (Trichogramma spp.) and Chrysoperla carnea

Abstract

The present work was conducted to evaluate the side effect of Neem products as natural insecticide on some of natural enemies of Helicoverpa armigera Hüb. (Trichogramma spp. and Chrysoperla sp.). The work is divided into laboratory and greenhouse experiments. The laboratory experiments dealt with the side effects of Neem products: NeemAzal-T/S, NeemAzal Blank, NeemAzal PC 05 and NeemAzal PC 05 Blank on Trichogramma spp. and C. carnea (Stephen). Neem concentrations: 2, 1, 0.5 and 0.25% were arranged in a completely randomized block design and each treatment was replicated at least six times. Parasitism and emergence rates of T. pretiosum (Riley) and T. minutum (Riley), and predation rates of C. carnea and its longevity were investigated. The greenhouse experiments dealt with the side effects of Neem products on parasitism rates of T. pretiosum and T. minutum on three different cotton cultivars. There were no serious side effects on parasitism and emergence rates of Trichogramma spp. and on efficiency of Chrysoperla. Similarly, Neem products achieved a good control of H. armigera in laboratory and greenhouse. Therefore, Neem products are recommended for controlling Helicoverpa, and are compatible with mass release of Trichogramma and Chrysoperla for organic cotton production. Future field experiments are suggested to further elucidate the effects of Neem on H. armigera and its natural enemies.     

Potential for the biological control ofLeptinotarsa decemlineata [Col.: Chrysomelidae] by the egg parasite,Edovum puttleri [Hym.: Eulophidae] in Maryland, 1981–84

The biological control potential ofEdovum puttleri Grissell, an exotic egg parasite ofLeptinotarsa decemlineata (Say), was examined in experimental potato plots. The parasite does not overwinter in Maryland, but through annual inoculative releases it consistently parasitized Colorado potato beetle (CPB) egg masses throughout the season. Approximately 50% of all egg masses collected were parasitized and maximum parasitism byE. puttleri occurred in egg masses produced by first generation adults.Edovum puttleri reproduced and maintained itself in potato plots for 15 weeks. The random sampling of CPB egg masses to evaluate the parasite in the field is described.      

Aerodynamic separation of parasitised from unparasitised insect host eggs

Aerodynamic sorting of host eggs exposed to Trichogramma egg parasitoids improved the percentage parasitism of retained product. Losses of parasitised eggs declined as percentage parasitism of the unsorted eggs increased. Trichogramma eclosion from parasitised eggs was similar for the unsorted and retained fractions; emergence from the parasitised culls was significantly lower.     

Phoresy in the field: natural occurrence of Trichogramma egg parasitoids on butterflies and moths

Phoretic insects utilize other animals to disperse to new environments. We recently discovered how egg parasitoids use an exciting phoretic strategy to reach egg-laying sites of their butterfly hosts. In the laboratory, female Trichogramma wasps detect and mount mated female cabbage white butterflies that emit an anti-aphrodisiac pheromone. Hardly any information exists about the natural occurrence of phoresy in wasps of this genus. Therefore, we monitored the presence of phoretic Trichogramma wasps on lepidopteran hosts in the field. Only female wasps were found at low prevalence on six lepidopteran species. Wasps were mostly found on female hosts and mainly on abundant solitary host species. This is the first report of phoretic Trichogramma wasps on butterflies in nature. We suggest that phoresy is only one of several strategies used by these polyphagous egg parasitoids. The evolution of phoresy is discussed in relation to the nutritional ecology of egg parasitoids.