The effect of bark thickness on host partitioning between Tetrastichus planipennisi (Hymen: Eulophidae) and Atanycolus spp. (Hymen: Braconidae), two parasitoids of emerald ash borer (Coleop: Buprestidae)

Parasitoids have recently been introduced from Asia to aid in biological control in the United States of the invasive, highly damaging, emerald ash borer, Agrilus planipennis. Three introduced parasitoids have established and field biological studies are underway to improve our understanding of niche partitioning among them. Here we report one such investigation, a field experiment conducted to determine how outer bark thickness of ash trees might affect parasitism by one introduced (Tetrastichus planipennisi) and one native parasitoid (Atanycolus spp.). We found that T. planipennisi was unable to parasitize EAB larvae in trees with outer bark thicker than 3.2 mm (>11.2-cm DBH) whereas Atanycolus spp. parasitized EAB larvae in ash trees with outer bark up to 8.8 mm thick (>57.4-cm DBH). These results suggest that establishment of, and control by T. planipennisi at release sites with only large diameter trees is less likely, and that T. planipennisi will be more effective in stands with younger trees (<12-cm DBH). Releasing T. planipennisi near the leading edge of EAB invasion may have little impact on EAB populations if many ash trees are too large. We recommend releasing T. planipennisi in stands dominated by small, early successional or regenerating ash trees. This may maximize the establishment and effectiveness of this species. This limitation of T. planipennisi for biological control of emerald ash borer suggests that other EAB parasitoids from its native range with longer ovipositors, such as Spathius galinae (Hymenoptera: Braconidae), should be sought and evaluated for possible use as EAB biocontrol agents in the US. The results of this study also suggest the importance of parasitoid guild introduction for biological control in general, and hint at possible broader implications relating to resource partitioning among native and introduced parasitoids.     

Host-seeking behavior and parasitism by Spathius agrili Yang (Hymenoptera: Braconidae), a parasitoid of the emerald ash borer

Spathius agrili Yang (Hymenoptera: Braconidae) is a newly described and important idiobiont ectoparasitoid of the emerald ash borer (EAB) that has excellent potential as a biological control agent against EAB populations in the USA. In order to understand the ecological factors involved in the search and discovery of concealed hosts by S. agrili, we investigated the behavioral responses of adult female wasps to potential semiochemicals from host plants, hosts, and host frass as well as to vibration signals from host feeding and movement. Using a bioassay, we showed that S. agrili first finds the host’s habitats by detecting the volatile compounds emitted by ash. In the second phase of host location and acceptance, the parasitoids detect the mechanical vibrations produced by host feeding and movement under the surface of the bark and then probe to find the EAB larvae. Contact chemicals seem to play little or no role in short-range host finding. Female wasps avoided laying eggs on EAB larvae already parasitized and thus paralyzed. We hypothesized that female wasps were not attracted to these larvae due to their lack of feeding or movement. While an induced paralysis in the host is instrumental in avoiding superparasitism, we cannot rule out that S. agrili females also use an oviposition pheromone to deter conspecific females. Together, these results suggest that vibration and olfactory cues play significant roles in distinct phases of S. agrili host habitat and host location behaviors.     

Population responses of hymenopteran parasitoids to the emerald ash borer (Coleoptera: Buprestidae) in recently invaded areas in north central United States

Populations of hymenopteran parasitoids associated with larval stages of the invasive emerald ash borer (EAB) Agrilus planipennis Fairmaire (Coleoptera: Buprestidae) were surveyed in 2009 and 2010 in the recently invaded areas in north central United States (Michigan), where two introduced EAB larval parasitoids, Tetrastichus planipennisi Yang and Spathius agrili Yang were released for classical biological control. Results from two years of field surveys showed that several hymenopteran parasitoids have become associated with EAB in Michigan. Among these parasitoids, the gregarious species T. planipennisi was the most abundant, accounting for 93% of all parasitoid individuals collected in 2009 (immediately after field release) and 58% in 2010 (a year later after field releases). Low levels (1–5%) of parasitism of EAB larvae by T. planipennisi were consistently detected at survey sites in both years. Separately, the abundance of the native parasitoid, Atanycolus spp., increased sharply, resulting in an average parasitism rate of EAB larvae from <0.5% in 2009 to 19% in 2010. Other parasitoids such as Phasgonophora sulcata Westwood, Spathius spp., Balcha indica Mani & Kaul, Eupelmus sp., and Eurytomus sp. were much less abundant than T. planipennisi and Atanycolus spp., and each caused <1% parasitism. Besides hymenopteran parasitoids, woodpeckers consumed 32–42% of the immature EAB stages present at our study sites, while undetermined biotic factors (such as microbial disease and host tree resistance) caused 10–22% mortality of observed EAB larvae. Relevance of these findings to the potential for biological control of EAB in the invaded areas of North America is discussed.     

Host suitability and fitness-related parameters of Campoletis sonorensis (Hymenoptera: Ichneumonidae) as a parasitoid of the cabbage looper,Trichoplusia ni (Lepidoptera: Noctuidae)

The seven age-classes of Trichoplusia ni (Hübner) larvae evaluated in this study as hosts of Campoletis sonorensis indicates that early 2nd larval instar (3–5 day-old larvae) of T. ni represents the most suitable host stage for the development of the larval endoparasitoid C. sonorensis. The higher suitability of early 2nd larval instar of T. ni resulted in more parasitised larvae, a higher rate of successful parasitoid emergence, a higher rate of female progeny, and a lower rate of immature parasitoid mortality. The fitness gain of C. sonorensis on late 1st larval instar (2 day-old larvae) and late 2nd larval instar –early 3rd instars (6–8 day-old larvae) stages of T. ni is negatively affected by the trade-offs between the different physiological and behavioral characteristics influencing their suitability as hosts of C. sonorensis.     

Abundance of volatile organic compounds in white ash phloem and emerald ash borer larval frass does not attract Tetrastichus planipennisi in a Y‐tube olfactometer

Many natural enemies employ plant‐ and/or herbivore‐derived signals for host/prey location. The larval parasitoid Tetrastichus planipennisi Yang (Hymenoptera: Eulophidae) is 1 of 3 biocontrol agents currently being released in an effort to control the emerald ash borer (EAB), Agrilus planipennis Fairmaire (Coloeptera: Burprestidae) in North America. To enhance its efficiency, allelochemicals that attract it need to be assessed. In this study, ash phloem volatile organic compounds (VOCs) of black, green, and white ash, and EAB larval frass were compared. Foraging behavior of T. planipennisi females in response to VOCs of white ash or frass from EAB larvae feeding on white ash phloem was tested using a Y‐tube olfactometer. Results indicated that the 3 ash species had similar VOC profiles. EAB larval frass generally contained greater levels of VOCs than phloem. Factor analysis indicated that the 11 VOCs could be broadly divided into 2 groups, with α‐bisabolol, β‐caryophyllene, (E)‐2‐hexenal, (Z)‐3‐hexenal, limonene, methyl benzoate, methyl indole‐3‐acetic acid, methyl jasmonate, methyl salicylate as the first group and the rest (i.e., methyl linoleate and methyl linolenate) as a second. Abundance of VOCs in white ash phloem tissue and frass, nevertheless, did not attract T. planipennisi females. The concealed feeding of EAB larvae might explain the selection for detectable and reliable virbrational signals, instead of undetectable and relatively unreliable VOC cues from phloem and frass, in short‐range foraging by T. planipennisi. Alternatively, it is possible that T. planipennisi is not amenable to the Y‐tube olfactometer assay employed.     

An improved method for monitoring parasitism and establishment of Oobius agrili (Hymenoptera: Encyrtidae), an egg parasitoid introduced for biological control of the emerald ash borer (Coleoptera: Buprestidae) in North America

Oobius agrili Zhang and Huang (Hymenoptera: Encyrtidae) is a solitary egg parasitoid that has been released in the United States since 2007 for biocontrol of the invasive emerald ash borer (EAB), Agrilus planipennis Fairmaire (Coleoptera: Buprestidae). Field and laboratory trials with ash logs infested with EAB eggs were conducted in Michigan between 2009 and 2010 to improve methods for monitoring the establishment of O. agrili. Naturally occurring EAB eggs were collected in both parasitoid-release and control (non-parasitoid-release) plots to compare with the EAB egg-sentinel log (ESL) technique. In three parasitoid-release plots, >50% of ESLs had O. agrili-parasitized eggs ranging from 3.9% to 48.2% egg parasitism after one week of field exposure. No EAB eggs were attacked by O. agrili on the ESLs deployed in control plots. In the laboratory, 100% of ESLs exposed to O. agrili inside rearing jars for one week had parasitized-eggs (68.5% egg parasitism). Deployment of ESLs detected low levels of parasitism by O. agrili in all three ash stands where O. agrili was released in previous years. In contrast, collection of naturally occurring EAB eggs detected the parasitism in only one of these three parasitoid-release ash stands. No parasitism was detected in control ash stands with either method. These findings indicate that populations of O. agrili released in previous years had successfully overwintered and established in the released ash stands by 2010, but had not yet dispersed to the control stands.     

Performance of Microplitis tuberculifer (Hymenoptera: Braconidae) parasitizing Mythimna separata (Lepidoptera: Noctuidae) in different larval instars

The solitary parasitoid Microplitis tuberculifer (Wesmael) is an important biological control agent of various lepidopteran pests in Asia. We examined the preference of M. tuberculifer for different instars of its common host, Mythimna separata (Walker), host instar effects on parasitoid development, and the consequences of parasitism in different stages for growth and consumption of host larvae. The wasp successfully parasitized the first four larval instars of M. separata, but not the fifth, which appeared to be behaviorally resistant. First and second instars were parasitized at higher rates compared to thirds and fourths in no-choice situations, ostensibly due to longer handling times for the latter, but second instars were most preferred in a choice test that presented all stages simultaneously. Although later instar hosts yielded heavier cocoons, the fastest parasitoid development was obtained in second instars. Lower sex ratios were obtained from first instars as females appeared to lay a smaller proportion of fertilized eggs in small hosts. Both weight gain and food consumption of parasitized larvae were reduced significantly within 24 h of parasitism, regardless of the stage parasitized, and final body weights were less than 10% those of unparasitized larvae. Thus, M. tuberculifer has good potential as a biological control agent of M. separata, successfully parasitizing the first four larval instars and dramatically reducing plant consumption by the host in all cases.     

Long-term monitoring of the introduced emerald ash borer (Coleoptera: Buprestidae) egg parasitoid,Oobius agrili (Hymenoptera: Encyrtidae), in Michigan,USA and evaluation of a newly developed monitoring technique

Emerald ash borer (EAB), Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), is a serious invasive pest of ash trees (Fraxinus spp.) in North America from China. The egg parasitoid Oobius agrili Zhang and Huang (Hymenoptera: Encyrtidae) was introduced from China as a biological control agent for this pest in Michigan and throughout the infested area of the United States. A critical component of any biological control program is post-release monitoring and evaluation; however, because of the small size and cryptic nature of O. agrili, evaluation of its impact is difficult. We compared two methods for measuring parasitism of emerald ash borer eggs: (1) timed visual searches of bark on standing ash trees and (2) bark collection, sifting, and sorting. Both methods were carried out in paired parasitoid-release and control plots, the visual search method over a six-year period (2008–2013) and the more recently developed bark-collection and sifting method for 2 years (2012–2013). The visual search method found parasitism in release plots remained low (0.7–4.2%) in samples taken from 2008 to 2012 and reached 10.6% in 2013. In comparison, the bark-sifting method found that rates of egg parasitism were considerably higher in release plots, 21.8% and 18.9% for samples taken in 2012 and 2013, respectively. These findings indicate that the population-level impact of O. agrili is increasing and may be an important source of mortality for EAB populations. We recommend the bark-collection and sifting method as the more effective method to recover parasitoids and estimate parasitism rates of O. agrili.     

Recent advances in biological control of important native and invasive forest pests in China

In recent decades, China has suffered severe attacks by both native and invasive forest pests. We have carried out a series of research projects on biological control of these pests. The fall webworm, Hyphantria cunea (Drury) (Lepidoptera: Arctiidae) has been sustainably controlled by an effective gregarious pupal endoparasitoid, Chouioia cunea Yang (Chalcidoidea: Eulophidae), which is native to China, and spraying HcNPV virus against the pest’s larval stage. Pine wilt disease, caused by the pine wood nematode (Bursaphelenchus xylophilus (Steiner et Buhrer) Nickle) (Aphelenchida: Aphelenchidae), is currently the number one pest in China. The strategy for controlling the disease is to manipulate the nematode’s vector, Monochamus alternatus Hope (Coleoptera: Cerambycidae). We discovered that Dastarcus helophoroides (Fairmaire) (Coleoptera: Bothrideridae) is the most important natural enemy in China pine forests. Mass rearing and release techniques were studied and developed. By releasing the parasitoid, 92.6% of the M. alternatus were parasitized in the first year. Meanwhile, three elaterid beetle species were found to prey on the larva of M. alternatus. The red turpentine beetle, Dendroctonus valens (LeConte) (Coleoptera:Scolytidae) was suppressed by a predator, Rhizophagus grandis Gyllenhal (Coleoptera: Rhizophagidae) introduced from Belgium and a total 3334 ha. of pine forests were protected. The oak longhorned beetle, Massicus raddei (Blessig) (Coleoptera: Cerambycidae) is the number one pest in the northeast forests of China, where it damages trunk of oaks, mainly Quercus liaotungensis and Q. mongolicus. An integrated management technique was developed for controlling the longhorned beetle: a special black light was invented for trapping the adults; the parasitoid Sclerodermus pupariae Yang et Yao (Hymenoptera: Bethylidae) was released against young larvae; and the parasitoid Dastarcus helophoroides eggs and/or adults were released when the hosts were mature larvae and/or pupae. By applying the technique for five years in northeastern China oak forests, the oak longhorned beetle has been controlled to a large extent. The emerald ash borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), is native to eastern Asia, including China, and feeding by larvae damages ash trees. Natural enemies of the emerald ash borer were investigated and seven species were found in China, of which Spathius agrili Yang (Hymenoptera: Braconidae), Tetrastichus planipennisi Yang (Hymenoptera: Eulophiae), Sclerodermus pupariae Yang et Yao and Oobius agrili Zhang et Huang (Hymenoptera: Encyrtidae) are predominant and have high potential for biocontrol of the pest. The biology, behavior, ecology and mass rearing techniques of the parasitoids were studied.     

Development of methods for the field evaluation of Oobius agrili (Hymenoptera: Encyrtidae) in North America,a newly introduced egg parasitoid of the emerald ash borer (Coleoptera: Buprestidae)

A field study was conducted in forested plots near Lansing, Michigan in 2008 and 2009 to evaluate the newly introduced egg parasitoid Oobius agrili Zhang and Huang (Hymenoptera: Encyrtidae) for control of the invasive emerald ash borer (EAB), Agrilus planipennis Fairmaire (Coleoptera: Buprestidae). To measure parasitism by O. agrili, laboratory-reared “sentinel EAB eggs” were deployed under bark flaps on trunks of selected ash trees in both parasitoid-release and non-release control plots. In addition, naturally occurring EAB eggs were collected in both parasitoid-release and control plots to measure parasitism. While no parasitism was detected with either sentinel or naturally occurring EAB eggs in control plots in either 2008 or 2009, a low level of parasitism by O. agrili was detected in the parasitoid-release plots in both artificially deployed sentinel eggs (?1%) and field-collected, naturally occurring eggs (1.1–4.2%) in both years. In addition to losses due to parasitism by O. agrili, a large proportion (37–52%) of the field-deployed sentinel eggs disappeared, possibly due to predators such as ants, in both parasitoid-release and control plots. While no statistical differences in parasitism by O. agrili were detected between parasitoid release and control plots, other sources of egg mortality such as disappearance due to predation on eggs, varied significantly across study sites in both 2008 and 2009. The relevance of these findings to future release and evaluation strategies for O. agrili for biological control of the invasive emerald ash borer in the US is discussed.     

Effects of the endoparasitoid Cotesia chilonis (Hymenoptera: Braconidae) parasitism,venom, and calyx fluid on cellular and humoral immunity of its host Chilo suppressalis (Lepidoptera: Crambidae) larvae

The larval endoparasitoid Cotesia chilonis injects venom and bracoviruses into its host Chilo suppressalis during oviposition. Here we study the effects of the polydnavirus (PDV)-carrying endoparasitoid C. chilonis (Hymenoptera: Braconidae) parasitism, venom and calyx fluid on host cellular and humoral immunity, specifically hemocyte composition, cellular spreading, encapsulation and melanization. Total hemocyte counts (THCs) were higher in parasitized larvae than in unparasitized larvae in the late stages following parasitization. While both plasmatocyte and granulocyte fractions and hemocyte mortality did not differ between parasitized and unparasitized hosts, in vitro spreading behavior of hemocytes was inhibited significantly by parasitism throughout the course of parasitoid development. C. chilonis parasitism suppressed the encapsulation response and melanization in the early stages. Venom alone did not alter cellular immune responses, including effects on THCs, mortality, hemocyte composition, cell spreading and encapsulation, but venom did inhibit humoral immunity by reducing melanization within 6 h after injection. In contrast to venom, calyx fluid had a significant effect on cell spreading, encapsulation and melanization from 6 h after injection. Dose–response injection studies indicated the effects of venom and calyx fluid synergized, showing a stronger and more persistent reduction in immune system responses than the effect of either injected alone.     

Host age preference of Microplitis mediator (Hymenoptera: Braconidae), an endoparasitoid of Mythimna separata (Lepidoptera: Noctuidae)

Microplitis mediator (Haliday) is a solitary endoparasitoid of larvae of the cotton bollworm, Helicoverpa armigera (Hübner) and the oriental armyworm, Mythimna = Leucania separata (Walker). The preference and suitability of different instars of M. separata for M. mediator were determined under laboratory conditions at a constant temperature of 26 ± 1 °C, 65 ± 5% RH and L14:D10 photoperiod. The selection coefficient revealed that M. mediator parasitized 1st to 4th instars, but preferred 2nd and 3rd instars. Seventy-one percent of parasitism was achieved within 24 h when the 2nd instars were used as hosts at a density of one parasitoid per 20 Larvae. Parasitoid egression and pupation were dependent on the host instar parasitized and occurred from the 1st through the 4th instar. The mean developmental time from egg to prepupae of M. mediator within 1st to 4th instars of the host was 8.27, 8.30, 8.30 and 9.20 days, respectively. Cocoon weights were lower when 1st and 2nd instars served as hosts rather than 3rd and 4th instars. The percentage of host larva that died before parasitoid egression declined as the age of the host increased, ranging from 26% to 2% for 1st–5th instars, respectively. The results of this study suggest that 2nd and 3rd instars of M. separata would be the best host stages for mass production of M. mediator in the laboratory and the best host instars to target for effective control in field releases.     

Assessment of frozen larvae of Callosobruchus maculatus as hosts for rearing Pteromalus cerealellae (Ashmead) (Hymenoptera: Pteromalidae)

The suitability of frozen host larvae for rearing Pteromalus cerealellae (Ashmead) (Hymenoptera: Pteromalidae), an ectoparasitoid of Callosobruchus maculatus (F.) (Coleoptera: Chrysomelidae) and other stored-product insects was investigated. The reproductive potential (number and sex ratio of progeny) of female P. cerealellae was compared on live (fresh) C. maculatus larvae (concealed within cowpea seeds) versus frozen larvae (obtained by freezing infested cowpea seeds at ?20 °C for 48 h) which were subsequently thawed and held at ambient conditions (～25 ± 1 °C, 50 ± 5% RH) for 4, 24, 48, 72, 96, and 120 h before exposure to female parasitoids. No significant differences were recorded in the numbers and sex ratios of the progeny produced by female P. cerealellae on live larvae compared to frozen host larvae that were thawed and held at ambient conditions for up to 96 h, suggesting that live and frozen larvae of C. maculatus are equally suitable for rearing P. cerealellae. However, the data showed that progeny production on frozen hosts gradually declined with thawing duration and was significantly reduced at the thawing duration of 120 h. When live and frozen host larvae were simultaneously presented together to female P. cerealellae at different exposure periods, relatively greater progeny production was recorded on live hosts than on frozen hosts at 12, 24, and 48 h of exposure. This may suggest preference of female P. cerealellae for live versus frozen host larvae. These results are discussed in relation to the life history strategy and host location behavior of P. cerealellae, and may have practical implications in the development of efficient mass rearing systems for the parasitoid.     

Bionomics of Orasema simplex (Hymenoptera: Eucharitidae), a parasitoid of Solenopsis fire ants (Hymenoptera: Formicidae) in Argentina

Biological characteristics of the parasitoid Orasema simplex Heraty (Hymenoptera: Eucharitidae), a potential candidate for the biological control of fire ants in the United States were investigated. Female survivorship, fertility and oviposition preferences were studied in the laboratory. Naturally parasitized colonies were examined to determine offspring sex ratio, development success and time, and to artificially parasitize healthy ant colonies. In addition, field studies were carried out to establish natural oviposition substrates and adult activity patterns. Orasema simplex female survivorship was 3.6 ± 1.5 days. Newly emerged females contained 613.5 ± 114.0 mature eggs. The adult development success in natural parasitized colonies was 22.2% with a female-biased sex ratio (4:1). The time required from planidia to adult was 29.5 ± 5.4 days. In the field, adults were mostly found around the ant nests at midday. A broad range of plant species was observed as oviposition substrates. The transfer of planidia to healthy ant colonies was achieved but the development success was very low. Orasema simplex appears to have a limited potential as a fire ant biocontrol agent because of cosmetic damage to a wide variety of plants used for oviposition. However, further studies are necessary to evaluate the real damage exerted by oviposition punctures.     

Pre-release evaluation of Semielacher petiolatus (Hymenoptera: Eulophidae) in quarantine for the control of citrus leafminer: Host discrimination,relative humidity tolerance,and alternative hosts

To evaluate the potential benefits and risks associated with releasing Semielacher petiolatus Girault (Hymenoptera: Eulophidae) in a classical biological control project directed against the citrus leafminer Phyllocnistis citrella Stainton (Lepidoptera: Gracillariidae) in Florida, we evaluated the ability of S. petiolatus females to discriminate between hosts previously parasitized by Ageniaspis citricola Logvinovskaya (Hymenoptera: Encyrtidae) and unparasitized hosts. In laboratory trials, S. petiolatus females did not discriminate between hosts previously parasitized and oviposited and fed on each host category equally. Hatch rate of S. petiolatus eggs on hosts previously parasitized by A. citricola was normal and development time was not different. However, mortality of immatures was significantly higher on previously parasitized hosts when compared to hosts that were not parasitized by A. citricola, and size of adult females reared on hosts previously parasitized was reduced. The relative survivorship of S. petiolatus adults compared with adults of P. citrella and another parasitoid, A. citricola, under three relative humidities (RHs) (55, 76, and 95% at 28 °C) indicated that S. petiolatus survives longer than A. citricola at all RHs tested, but did not survive as long as the citrus leafminer. Finally, the leafminer Liriomyza trifolii Burgess (Diptera: Agromyzidae) was evaluated as a possible host for S. petiolatus but no progeny were produced in choice and no-choice tests. The lack of discrimination raises the concern that S. petiolatus could disrupt the efficacy of A. citricola, which is already established in Florida, without providing substantial reduction of citrus leafminer populations during early spring.     

Biology of Apanteles myeloenta (Hymenoptera: Braconidae), a larval parasitoid of carob moth Ectomyelais ceratoniae (Lepidoptera: Pyralidae)

The carob moth, Ectomyelois ceratoniae (Lepidoptera: Pyralidae), is the most important pest of pomegranate orchards (in terms of economic damage) within Iran, and hence, several control procedures, including biological methods of control, have been attempted as a means of controlling populations of this insect. This research was carried out in order to study the biology of Apanteles myeloenta (Hymenoptera: Braconidae), a larval parasitoid of the carob moth. Laboratory studies were conducted to determine larval developmental time, adult longevity, sex ratio, parasite progeny production, and host stage preference of A. myeloenta. At 25 ± 1 °C, immature developmental time (egg to pupa; mean ± SE) was 28.33 ± 0.85 days and 27.46 ± 0.37 days for male and females, respectively. Adult females survived on average 17.5 ± 0.14, 11.7 ± 0.22, 3.4 ± 0.18, and 2.8 ± 0.12 days at 25 C when provided with honey and water, honey only, water only or no food source, respectively. The sex ratio (females to males) of A. myeloenta was 1:3.5 from hosts parasitized in the first instar, 1:3 for second instars and 1:2 for third instar carob moth larvae. Female A. myeloenta typically preferred to parasitize second instar over third or first instar. The oviposition activity peaked on the 7th and 8th days following emergence, when provided with honey, and 10% sucrose solution, respectively.     

Oviposition and reproductive performance of a generalist parasitoid (Trichogramma pretiosum) exposed to host species that differ in their physical characteristics

The response of generalist egg parasitoids to alternative natural hosts that are present simultaneously is not well known. We investigated the behavior of Trichogramma pretiosum Riley (Hymenoptera: Trichogrammatidae) in relation to two field hosts Helicoverpa armigera Hübner and Spodoptera litura Fabricius, in choice and no choice tests. We quantified the effects of natal host species and post-emergence adult age on the oviposition preference of the parasitoids. H. armigera eggs were consistently preferred over S. litura eggs, regardless of the natal host and adult age. When only S. litura eggs were available as hosts, they were parasitized at statistically similar rates to H. armigera eggs (average of 17 ± 2.7 vs. 13 ± 3.0, H. armigera to S. litura). The adult lifespan and lifetime fecundity of T. pretiosum were variable but were affected by natal host species and/or host species to which they were exposed. Mean lifespan and fecundity of parasitoids that had developed in H. armigera eggs and were exposed to H. armigera eggs for oviposition were 13.9 ± 1.8 days and 98.7 ± 11.0 adult offspring. By contrast, those that developed in S. litura eggs and were exposed to S. litura eggs for oviposition lived for 7 ± 0.9 days and produced 53.8 ± 8.0 adult offspring. The ovigeny index (OI) was significantly lower in the parasitoids exposed to H. armigera eggs than in those exposed to S. litura eggs, regardless of the natal host, indicating that H. armigera eggs sustain the adult parasitoids better than S. litura eggs. These results are used to predict parasitoid behavior in the field when both hosts are available.     

Native parasitoids and recovery of Spathius agrili from areas of release against emerald ash borer in eastern Tennessee,USA

The parasitoid Spathius agrili Yang, introduced in the USA to suppress populations of the emerald ash borer (EAB), Agrilus planipennis Fairmaire, has been recovered at a release site for the first time in eastern Tennessee after a single year of releases. Other native parasitoids, including Spathius floridanus Ashmead, undetermined species of Spathius (possibly Spathius elegans Matthews and Spathius parvulus Matthews) and Atanycolus cappaerti Marsh & Strazanac, also known to be associated with EAB, were recovered. These recoveries represent the first documentation of these four species, including the introduced S. agrili, associated with EAB in the southern USA. Implications for biological control efforts against EAB are discussed.     

Role of emerald ash borer (Coleoptera: Buprestidae) larval vibrations in host-quality assessment by Tetrastichus planipennisi (Hymenoptera: Eulophidae)

The biological control agent Tetrastichus planipennisi Yang (Hymenoptera: Eulophidae) is a gregarious larval endoparasitoid of the emerald ash borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), an invasive cambium-feeding species responsible for recent, widespread mortality of ash (Fraxinus spp.) in North America. T. planipennisi is known to prefer late-instar emerald ash borer, but the cues used to assess host size by this species and most other parasitoids of concealed hosts remain unknown. We sought to test whether vibrations produced by feeding emerald ash borer vary with larval size and whether there are any correlations between these cues and T. planipennisi progeny number (i.e., brood size) and sex ratio. The amplitudes and rates of 3-30-ms vibrational impulses produced by emerald ash borer larvae of various sizes were measured in the laboratory before presenting the larvae to T. planipennisi. Impulse-rate did not vary with emerald ash borer size, but vibration amplitude was significantly higher for large larvae than for small larvae. T. planipennisi produced a significantly higher proportion of female offspring from large hosts than small hosts and was shown in previous work to produce more offspring overall from large hosts. There were no significant correlations, however, between the T. planipennisi progeny data and the emerald ash borer sound data. Because vibration amplitude varied significantly with host size, however, we are unable to entirely reject the hypothesis that T. planipennisi and possibly other parasitoids of concealed hosts use vibrational cues to assess host quality, particularly given the low explanatory potential of other external cues. Internal chemical cues also may be important.