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.     

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.     

Interactions between Spathius agrili (Hymenoptera: Braconidae) and Tetrastichus planipennisi (Hymenoptera: Eulophidae), larval parasitoids of Agrilus planipennis (Coleoptera: Buprestidae)

Three hymenopteran parasitoids native to China are being released in the United States as biological control agents for the emerald ash borer (EAB), Agrilus planipennis Fairmaire, an Asian buprestid species responsible for mortality of ash trees (Fraxinus spp.) in North America. Two of these hymenopterans, Spathius agrili Yang (Braconidae), a larval ectoparasitoid, and Tetrastichus planipennisi Yang (Eulophidae), a larval endoparasitoid, prefer late-instar EAB larvae. This overlapping host preference raises concerns that interspecific competition following field releases may compromise establishment of one or both species. In a series of laboratory and field experiments, we found S. agrili and T. planipennisi exhibited similar parasitism rates when presented alone with EAB larvae for 12–14 days. However, S. agrili was more efficient at locating and parasitizing hosts within the first 27 h, possibly explaining why S. agrili excluded T. planipennisi in the laboratory trials and nearly excluded T. planipennisi in field trials when the two species were presented together with EAB larvae. We found that S. agrili parasitized larvae previously parasitized by T. planipennisi but not the reverse. However, S. agrili offspring failed to complete development on hosts that were previously parasitized by T. planipennisi. We recommend releasing these species separately in time or space to avoid the antagonistic interactions observed in this study.     

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.     

Climate variation alters the synchrony of host–parasitoid interactions

Observed changes in mean temperature and increased frequency of extreme climate events have already impacted the distributions and phenologies of various organisms, including insects. Although some research has examined how parasitoids will respond to colder temperatures or experimental warming, we know relatively little about how increased variation in temperature and humidity could affect interactions between parasitoids and their hosts. Using a study system consisting of emerald ash borer (EAB), Agrilus planipennis, and its egg parasitoid Oobius agrili, we conducted environmentally controlled laboratory experiments to investigate how increased seasonal climate variation affected the synchrony of host–parasitoid interactions. We hypothesized that increased climate variation would lead to decreases in host and parasitoid survival, host fecundity, and percent parasitism (independent of host density), while also influencing percent diapause in parasitoids. EAB was reared in environmental chambers under four climate variation treatments (standard deviations in temperature of 1.24, 3.00, 3.60, and 4.79°C), while O. agrili experiments were conducted in the same environmental chambers using a 4 × 3 design (four climate variation treatments × 3 EAB egg densities). We found that EAB fecundity was negatively associated with temperature variation and that temperature variation altered the temporal egg laying distribution of EAB. Additionally, even moderate increases in temperature variation affected parasitoid emergence times, while decreasing percent parasitism and survival. Furthermore, percent diapause in parasitoids was positively associated with humidity variation. Our findings indicate that relatively small changes in the frequency and severity of extreme climate events have the potential to phenologically isolate emerging parasitoids from host eggs, which in the absence of alternative hosts could lead to localized extinctions. More broadly, these results indicate how climate change could affect various life history parameters in insects, and have implications for consumer–resource stability and biological control.     

Ash (Fraxinus spp.) mortality,regeneration, and seed bank dynamics in mixed hardwood forests following invasion by emerald ash borer (Agrilus planipennis)

Emerald ash borer (EAB; Agrilus planipennis) has killed millions of ash trees and threatens ash throughout North America, and long-term persistence of ash will depend on the potential for regeneration. We quantified ash demography, including mortality and regeneration, of Fraxinus americana (white ash), Fraxinus pennsylvanica (green ash), and Fraxinus nigra (black ash) in mixed hardwood forests near the epicenter of the EAB invasion in southeastern Michigan and throughout Ohio. Plots were established across a gradient of ash densities. Ash was the most important species in hydric sites, and ranked second among all species in mesic and xeric sites. In sites nearest the epicenter in Michigan, ash mortality exceeded 99 % by 2009, and few or no newly germinated ash seedlings were observed, leaving only an “orphaned cohort” of established ash seedlings and saplings. As ash mortality increased, the number of viable ash seeds in soil samples decreased sharply, and no viable seeds were collected in 2007 or 2008. In Ohio sites farther from the epicenter, densities of new ash seedlings were much higher in plots with healthy ash trees compared to plots where trees had died. EAB was still present in low densities in Michigan and Ohio stands in 2012 where average mortality of ash was nearly 100 %. The future of ash at these sites will depend on the outcome of the dynamic interaction between the orphaned cohort of previously established ash seedlings and saplings and low density EAB populations.     

Patterns of parasitoid host utilization and development across a range of temperatures: implications for biological control of an invasive forest pest

Although climate change frequently has been linked to observed shifts in the distributions or phenologies of species, little is known about the potential effects of varying temperatures on parasitoids and their relationships with hosts. Using the egg parasitoid Oobius agrili (Hymenoptera: Encyrtidae) we examined host utilization patterns of this species across a range of temperatures (20–35 °C) to explore how changing climate could affect the interaction with its host—the emerald ash borer (EAB) (Coleoptera: Buprestidae), a serious invasive forest pest that has killed tens of millions of ash (Fraxinus spp.) trees in North America. Results from our study showed that the window of host susceptibility to O. agrili parasitism declined significantly from 14.8 to 2.6 days in an inverse second-order relationship with increasing exposure temperatures from 20 to 35 °C. In contrast, parasitoid host attack rate changed in a bell-shaped second-order relationship—i.e., increased with temperatures from 20 to 25 °C, but decreased at about the same rate when temperatures increased from 30 to 35 °C. This range of temperatures also significantly affected the development and mortality of immature parasitoids with 35 °C resulting in 100 % mortality. There was little mortality (0–4.5 %) and no significant differences in the percentage (20.9–34.9 %) of immature O. agrili that entered diapause (as mature larvae) at 20, 25, and 30 °C. However, there were significant differences in the time event of adult wasp emergence within this temperature range. The median time for 50 % of immature O. agrili emerging as adults at 20, 25, and 30 °C were 38, 18, and 17 days after parental wasp oviposition, respectively. Together these findings indicate that the non-linear and unequal temperature effects on these host utilization parameters are likely to result in differential host parasitism rates, and thus could reduce the efficacy of this parasitoid in suppressing host populations due to climate change (global warming and extreme heat).     

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.     

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.     

Quantifying the Impact of Woodpecker Predation on Population Dynamics of the Emerald Ash Borer (Agrilus planipennis)

The emerald ash borer (EAB), Agrilus planipennis, is an invasive beetle that has killed millions of ash trees (Fraxinus spp.) since it was accidentally introduced to North America in the 1990s. Understanding how predators such as woodpeckers (Picidae) affect the population dynamics of EAB should enable us to more effectively manage the spread of this beetle, and toward this end we combined two experimental approaches to elucidate the relative importance of woodpecker predation on EAB populations. First, we examined wild populations of EAB in ash trees in New York, with each tree having a section screened to exclude woodpeckers. Second, we established experimental cohorts of EAB in ash trees in Maryland, and the cohorts on half of these trees were caged to exclude woodpeckers. The following spring these trees were debarked and the fates of the EAB larvae were determined. We found that trees from which woodpeckers were excluded consistently had significantly lower levels of predation, and that woodpecker predation comprised a greater source of mortality at sites with a more established wild infestation of EAB. Additionally, there was a considerable difference between New York and Maryland in the effect that woodpecker predation had on EAB population growth, suggesting that predation alone may not be a substantial factor in controlling EAB. In our experimental cohorts we also observed that trees from which woodpeckers were excluded had a significantly higher level of parasitism. The lower level of parasitism on EAB larvae found when exposed to woodpeckers has implications for EAB biological control, suggesting that it might be prudent to exclude woodpeckers from trees when attempting to establish parasitoid populations. Future studies may include utilizing EAB larval cohorts with a range of densities to explore the functional response of woodpeckers.     

Trichogramma releases in North Carolina cotton: why releases fail to suppress heliothine pests

Field studies were conducted in 1996 and 1997 to determine the fate of naturally oviposited F3 heliothine eggs in cotton plots treated with augmentative releases of Trichogramma exiguum Pinto & Platner and nontreated plots. Four cohorts of newly oviposited eggs (< 24 h old) were followed in 1996 and two cohorts in 1997. In 1996, mean +/- SD percent parasitism, estimated by in-field studies following the fate of naturally oviposited eggs, ranged from 7 +/- 7 to 61 +/- 8% in T. exiguum release plots and 0 +/- 0 to 35 +/- 13% in control plots. The mean +/- SD percent of eggs hatched in T. exiguum release plots ranged from 1 +/- 2 to 11 +/- 4% and 7 +/- 4 to 28 +/- 10% in control plots. In 1997, mean +/- SD percent egg parasitism ranged from 27 +/- 4 to 40 +/- 3% in T. exiguum release plots and 15 +/- 18 to 25 +/- 8% in control plots. The mean +/- SD percent of eggs hatched in T. exiguum release plots ranged from 7 +/- 3 to 12 +/- 2% and 18 +/- 6 to 28 +/- 8% in control plots. Despite increased parasitism and reduced egg hatch in T. exiguum release plots, overall, there was no significant difference in larval density (all instars combined) between T. exiguum release and control plots. Combined analysis of the heliothine larval populations and egg fate data revealed that the additional egg mortality produced by released T. exiguum was offset by lower larval mortality in release plots. Because of the occurrence of compensatory mortality, the egg stage of heliothines is not an appropriate target for biological control using Trichogramma wasp releases.     

Density dependence of Anaphes sordidatus (Hymenoptera: Mymaridae) parasitism on eggs of Listronotus oregonensis (Coleoptera: Curculionidae)

Listronotus oregonensis (LeConte) (Coleoptera: Curculionidae) oviposits in carrot leaves and the larvae feed in carrot roots. Its eggs are parasitized by Anaphes sordidatus (Girault) (Hymenoptera: Mymaridae) throughout its oviposition period. This parasitoid is the major biotic mortality factor for L. oregonensis. Parasitism by A. sordidatus was studied for three years in sequentially sown plots of carrots, Daucus carota var. sativa. Over the scason, significantly fewer L. oregonensis eggs were oviposited in later sown carrots than in earlier sown carrots because oviposition started later in late sown plots of carrots. A positive temporal density-dependent relationship was observed each year between parasitism rates and host densities. This positive density dependence occurred in early and mid-summer for earlysown carrots where host density reached 1–2 host eggs per plant but disappeared in late summer when host density decreased while parasitism remained high. Latesown carrots had low host egg density (0.2 host egg per plant) and contributed little to the total number of eggs. In these late sown plots, parasitism increased rapidly to over 80% but no density dependence was observed. Spatially, few statistically significant regressions were found but all indicated a positive spatial density-dependence. Most non-significant regressions occurred because the range of egg density was too small between plots for a given date.     

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.      

Enhancement of Riptortus clavatus (Thunberg) (Hemiptera: Alydidae) egg parasitism by a component of the bug's aggregation pheromone

Egg parasitism of the bean bug, Riptortus clavatus (Thunberg) (Hemiptera: Alydidae), was surveyed using individual egg bags in a sweet persimmon orchard and at Gyeongsang National University (GNU) campus, Korea in 2006. The effect of (E)-2-hexenyl (Z)-3-hexenoate (E2HZ3H), one component of R. clavatus aggregation pheromone, on the parasitism enhancement was tested at GNU campus and in a soybean field in 2005 and 2006, respectively. In 2006, two egg parasitoid species, Ooencyrtus nezarae Ishii (Hymenoptera: Encyrtidae) and Gryon japonicum (Ashmead) (Hymenoptera: Scelionidae), emerged from R. clavatus eggs. Parasitism by O. nezarae was 9.4–48.3% in mid August to mid September in GNU campus and 6.7–30.0% at the orchard. Total parasitism by G. japonicum (2.5%) at both sites throughout the experimental period was lower than that by O. nezarae (12.5%). This survey revealed nearly no activity of the two species after October at both sites. G. japonicum was solitary and O. nezarae could be either solitary or gregarious. From a single R. clavatus egg, one female or one male G. japonicum emerged. However, an average of 4.3 O. nezarae emerged from one host egg. It took 12.6 d for G. japonicum and 12.0 d for O. nezarae to emerge from R. clavatus eggs in the laboratory. Treatment with E2HZ3H increased parasitism by O. nezarae in both years, but did not increase parasitism by G. japonicum. This suggests that E2HZ3H can be used as a kairomone to reduce the density of R. clavatus in the fields where natural parasitism by O. nezarae is high.     

Measuring the impact of biotic factors on populations of immature emerald ash borers (Coleoptera: Buprestidae)

Cohorts of emerald ash borer larvae, Agrilus planipennis Fairmaire, were experimentally established in July of 2008 on healthy green ash (Fraxinus pennsylvanica) trees in two wooded plots at each of three sites near Lansing, MI, by caging gravid emerald ash borer females or placing laboratory-reared eggs on trunks (0.5-2 m above the ground) of selected trees. One plot at each site was randomly chosen for release of two introduced larval parasitoids, Tetrastichus planipennisi Yang (Hymenoptera: Eulophidae) and Spathius agrili Yang (Hymenoptera: Braconidae), whereas the other served as the control. Stage-specific mortality factors and rates were measured for all experimentally established cohorts and for associated wild (i.e., naturally occurring) emerald ash borer immature stages via destructive sampling of 2.5 m (above the ground) trunk sections of cohort-bearing trees in the spring and fall of 2009. Host tree defense was the most important mortality factor, causing 32.0 to 41.1% mortality in the experimental cohorts and 17.5 to 21.5% in wild emerald ash borer stages by spring 2009, and 16.1 to 29% for the remaining experimental cohorts, and 9.9 to 11.8% for wild immature emerald ash borer stages by fall 2009. Woodpecker predation was the second most important factor, inflicting no mortality in the experimental cohorts but causing 5.0 to 5.6% mortality to associated wild emerald ash borer stages by spring 2009 and 9.2 to 12.8% and 3.2 to 17.7%, respectively, for experimental cohorts and wild emerald ash borer stages by fall 2009. Mortality from disease in both the experimental and wild cohorts was low (<3%) in both the spring and fall sample periods. In the fall 2009 samples, ≈ 1.5% of experimental cohorts and 0.8% of the wild emerald ash borer stages were parasitized by T. planipennisi. While there were no significant differences in mortality rates because of parasitism between parasitoid-release and control plots, T. planipennisi was detected in each of the three release sites by the end of the study but was not detected in the experimental cohorts or associated wild larvae in any of the three control plots.     

Factors influencing the dispersal of a native parasitoid, <Emphasis Type="Italic">Phasgonophora sulcata</Emphasis>, attacking the emerald ash borer: implications for biological control

High parasitism by a native parasitoid, Phasgonophora sulcata Westwood (Hymenoptera: Chalcididae), has been reported on emerald ash borer (hereafter EAB), Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), in North America. Use of this parasitoid in an augmentative biological control program has been proposed to slow the spread of EAB, yet information is lacking on key aspects of this parasitoid’s dispersal. We document the flight capacity and walking activity of P. sulcata, its potential fecundity, and describe how age, body size, temperature, and time of day affect these parameters. Wasp flight capacity, measured using flight mills, increased with temperature and decreased with age. Unexpectedly, age and body size did not affect wasp walking activity, and we saw no relationship between walking activity and flight capacity. Older wasps had lower potential fecundity than younger wasps. These results suggest that P. sulcata should be released as pupae near EAB-infested ash trees to improve efficacy and potential biological control success.     

Effects of combining releases of non-viable host eggs with insecticide application on Riptortus pedestris population and its egg parasitoids

Our previous study demonstrated that the release of refrigerated non-viable eggs of Riptortus pedestris (Fabricius) (Hemiptera: Alydidae) enhanced parasitism rates in soybean fields but did not result in the reduction of R. pedestris populations. This study was further conducted using an open-cage exclusion design in a soybean field in order to evaluate the compatibility of combining releases of non-viable host eggs with a single pre-harvest application of insecticide for the control of R. pedestris. Refrigerated eggs of R. pedestris were released twice in treatment plots, and fresh (< 1 day old) eggs of R. pedestris were deployed in all experimental arenas, every 6 days, for host resource and measurement of field parasitism. The releases of host eggs did not reduce the number of R. pedestris in any life stage except the adult stage on two sampling dates. However, parasitism by Gryon japonicum (Ashmead) (Hymenoptera: Scelionidae) was higher in treated plots (9–25%) than in the control plots (1–9%). Statistical significant reduction was not found in the pest population, but parasitism rates significantly increased. Pesticide application did not reduce the bug population but did affect the parasitoids population. Pest management tactics, using both artificially deployed host eggs and insecticide, are discussed.     

Potential Species Replacements for Black Ash (<Emphasis Type="Italic">Fraxinus nigra</Emphasis>) at the Confluence of Two Threats: Emerald Ash Borer and a Changing Climate

The emerald ash borer (Agrilus planipennis; EAB) is causing widespread mortality of ash (Fraxinus spp.) and climate change is altering habitats of tree species throughout large portions of North America. Black ash (F. nigra), a moist-soil species common in the Northwoods of Minnesota, Wisconsin, and Michigan, USA, is under a double threat of losing habitat from climate change and near annihilation from EAB. Because black ash often occurs in nearly pure stands, planting non-ash species is a management strategy already underway or being planned for thousands of acres. Tools are needed to assist managers in prioritizing sites for early treatment and to select potential species to replace black ash. This study explores the implications of threats to black ash ecosystems using analyses of field data and models to assess both the threats to, and potential replacement species for, black ash in Minnesota. For our analysis we (1) assessed the status of ashes and co-occurring species in forest inventory plots throughout Minnesota; (2) modeled the risk of EAB attack for multiple years in Minnesota; (3) modeled potential impacts of climate change on tree species with current or potential future habitat in Minnesota; (4) evaluated species co-occurring with black ash in plots in Ohio and Michigan, southeast of Minnesota; and (5) synthesized these results to provide a classification for candidate replacement species, both from within Minnesota and from points farther south. Though this process is demonstrated for black ash in Minnesota, the elements to be considered and modeled would be similar for any other location with a pest or pathogen threat for a species which simultaneously faces a changing climate.     

Susceptibility of Nezara viridula (L.) (Hemiptera: Pentatomidae) Egg Masses of Different Sizes to Parasitism by Trissolcus basalis (Woll.) (Hymenoptera: Platygastridae) in the Field

Egg masses of Nezara viridula (L.) are commonly parasitized by Trissolcus basalis (Woll.), and we investigated the role of size of egg masses on parasitization by T. basalis. Sentinel egg masses were exposed to parasitism in the field for 6–7 days, when they were collected for evaluation of parasitoid emergence. We recorded the number of eggs per egg mass, the number of emerged hosts, and the number of empty and parasitized eggs. We calculated the proportion of attacked host egg masses (DE), the proportion of parasitized eggs per attacked egg mass (PE), and total parasitism (PI). The total number of egg masses exposed to parasitism was 330. The minimum, mean, and maximum egg mass sizes were 25, 75.2, and 111, respectively. DE and PE varied widely between different fields, and they were independent of egg mass size. In 14.2% of all parasitized egg masses, we found simultaneous emergence of T. basalis and N. viridula independently of host egg mass size. PE exhibited low variability compared with PI and DE, which were linearly related. PI and DE values from other field studies are consistent with the linear relationship, suggesting that PI is mostly related to the proportion of the DE. This also suggests that total parasitism is independent of egg mass size, of possible differences in plant species, and T. basalis density and strains.