Laboratory Evaluation of the Toxicity of Systemic Insecticides to Emerald Ash Borer Larvae.

Emerald ash borer (Agrilus planipennis Fairmaire) (Coleoptera: Buprestidae), an invasive phloem-feeding insect native to Asia, threatens at least 16 North American ash (Fraxinus) species and has killed hundreds of millions of ash trees in landscapes and forests. We conducted laboratory bioassays to assess the relative efficacy of systemic insecticides to control emerald ash borer larvae in winter 2009 and 2010. Second- and third-instar larvae were reared on artificial diet treated with varying doses of emamectin benzoate (TREE-äge, Arborjet, Inc., Woburn, MA), imidacloprid (Imicide, J. J Mauget Co., Arcadia, CA), dinotefuran (Safari, Valent Professional Products, Walnut Creek, CA), and azadirachtin (TreeAzin, BioForest Technologies, Inc., Sault Ste. Marie, Ontario, and Azasol, Arborjet, Inc., Woburn, MA). All of the insecticides were toxic to emerald ash borer larvae, but lethal concentrations needed to kill 50% of the larvae (LC50), standardized by larval weight, varied with insecticide and time. On the earliest date with a significant fit of the probit model, LC50 values were 0.024 ppm/g at day 29 for TREE-äge, 0.015 ppm/g at day 63 for Imicide, 0.030 ppm/g at day 46 for Safari, 0.025 ppm/g at day 24 for TreeAzin, and 0.027 ppm/g at day 27 for Azasol. The median lethal time to kill 50% (LT50) of the tested larvae also varied with insecticide product and dose, and was longer for Imicide and Safari than for TREE-äge or the azadirachtin products. Insecticide efficacy in the field will depend on adult and larval mortality as well as leaf and phloem insecticide residues.

Enantiomeric composition of α-pinene affects catches of bark and wood boring beetles, and associated species, in ethanol-baited multiple-funnel traps.

In 2009, we determined the effects of the enantiomeric composition of the kairomone, α-pinene, on trap catches of arboreal beetles (Coleoptera) in stands of eastern pine trees with resin dominated by (+)-α-pinene. We hypothesized that the responses of beetles would correlate with the predominant enantiomer of α-pinene found in host pines. Lures of (+)-, racemic (±), and (-)-α-pinene were added separately to ethanol-baited multiple-funnel traps. Species such as Monarthrum mali (Fitch), Dendroctonus terebrans (Olivier), Ips grandicollis (Eichhoff), and Pachylobius picivorus (Germar) (Coleoptera: Curculionidae) showed a preference for traps co-baited with (-)-α-pinene. α-Pinene enhanced attraction of Hylastes salebrosus Eichhoff, Hylastes porculus Erickson and Hylastes tenuis Eichhoff (Coleoptera: Curculionidae) to ethanol-baited traps with no effects from enantiomeric composition of α-pinene. The attraction of the ambrosia beetles, Xyleborinus saxesenii (Ratzeburg) and Dryoxylon onoharaense (Murayama) (Coleoptera: Curculionidae) to ethanol-baited traps was interrupted by the addition of α-pinene, regardless of enantiomeric composition. Species such as Xylosandrus germanus (Blandford), Cnestus mutilatus (Blandford) and Stenoscelis brevis (Boheman) (Coleoptera: Curculionidae) were unaffected by the presence of α-pinene. Trap catches of some species of longhorn beetles and bark beetle predators (Coleoptera: Cerambycidae, Cleridae, Elateridae, Histeridae, and Trogossitidae) were increased by the addition of α-pinene, although results varied by location. Platysoma spp. (Coleoptera: Histeridae) showed a marked preference for traps co-baited with (+)-α-pinene in Florida and Georgia. In summary, we found that the enantiomeric composition of α-pinene in hosts was not a good predictor of enantiomeric preferences by beetles.

Tree species richness and ash density have variable effects on emerald ash borer biological control by woodpeckers and parasitoid wasps in post-invasion white ash stands.

Emerald ash borer (EAB) (Agrilus planipennis Fairmaire) (Coleoptera: Buprestidae) is the most destructive insect to invade North American forests. Identifying habitat features that support EAB natural enemies is necessary to enhance EAB biological control. In many forest ecosystems, tree species diversity has been linked with reduced pest abundance and increases in natural enemy abundance. We assessed the influence of tree species richness, ash density, and proportion of total ash basal area on ash canopy condition, EAB larval densities, and biocontrol by woodpeckers and parasitoids in pairs of healthy and declining overstory (DBH > 10 cm) and recruit-sized ash (DBH 2-10 cm) in 4 post-invasion forests in Michigan, USA. Tree species richness and ash density were not significantly associated with EAB larval densities, ash canopy dieback and transparency, and woodpecker predation of EAB larvae. In declining and healthy overstory ash, woodpeckers killed 38.5 ±â€…3.9% and 13.2 ±â€…3.7% of larvae, respectively, while the native parasitoid Phasgonophora sulcata Westwood killed 15.8 ±â€…3.8% and 8.3 ±â€…3.0% and the introduced parasitoid Spathius galinae Belokobylskij & Strazanac killed 10.8 ±â€…2.5% and 5.0 ±â€…2.6% of EAB larvae. Parasitism by P. sulcata was inversely related to ash density while parasitism by S. galinae was positively associated with ash density. Ash density, but not tree diversity, appears to differentially influence biological control of EAB by parasitoids, but this effect is not associated with reduced EAB densities or improved canopy condition.

Trap lure blend of pine volatiles and bark beetle pheromones for Monochamus spp. (Coleoptera: Cerambycidae) in pine forests of Canada and the United States.

In 2007-2008, we examined the flight responses of Monochamus titillator (F.) complex [M. titillator, Monochamus carolinensis (Olivier), and any possible hybrids], Monochamus scutellatus (Say), Monochamus clamator (LeConte), Monochamus obtusus Casey, and Monochamus mutator LeConte (Coleoptera: Cerambycidae) to multiple-funnel traps baited with and without host volatiles and bark beetle pheromones. Experiments were conducted in mature pine (Pinus) stands in Alberta (Canada), and Arkansas, Arizona, California, Florida, Idaho, Michigan, Montana, New Hampshire, North Carolina, Ohio, Oregon, Tennessee, Utah, and Wisconsin (United States). At each location, traps were deployed in 10 replicate blocks of four traps per block. The trap treatments were: 1) blank control; 2) ipsenol and ipsdienol; 3) ethanol and alpha-pinene; and 4) a quaternary blend of ipsenol, ipsdienol, ethanol, and alpha-pinene. All five species or species complex of Monochamus preferred traps baited with the quaternary blend over all other treatments. The consistency of these results across such a large geographic area suggests that similar selection pressures may be acting on Monochamus spp. in pine forests, regardless of variation in stand composition and climatic conditions. Our results suggest that multiple-funnel traps baited with the quaternary blend ofipsenol, ipsdienol, ethanol, and alpha-pinene may be highly effective for monitoring various Monochamus spp. in pine forests of North America, and may have utility in trapping and detection programs in North America and overseas.

Postrelease assessment of Oobius agrili (Hymenoptera: Encyrtidae) establishment and persistence in Michigan and the Northeastern United States.

The emerald ash borer (EAB), Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), is an invasive woodboring pest of ash trees (Fraxinus sp.) in North America. Among the Asiatic parasitoids being released for the management of EAB in North America, Oobius agrili Zhang and Huang (Hymenoptera: Encyrtidae) is the only EAB egg parasitoid. To date, more than 2.5 million O. agrili have been released in North America; however, few studies have examined its success as a biological control agent of EAB. We conducted studies to assess O. agrili establishment, persistence, spread, and EAB egg parasitism rates in Michigan at the earliest release sites (2007-2010), as well as at more recent release sites (2015-2016) in 3 Northeastern United States (Connecticut, Massachusetts, New York). In both regions, we documented successful O. agrili establishment at all but one release site. In Michigan, O. agrili has persisted at release sites for over a decade and spread to all control sites located 0.6-3.8 km from release sites. Overall, EAB egg parasitism ranged from 1.5% to 51.2% (mean of 21.4%) during 2016-2020 in Michigan and from 2.6% to 29.2% (mean of 16.1%) during 2018-2020 in the Northeastern states. Future research efforts should focus on factors affecting the spatiotemporal variation in EAB egg parasitism rates by O. agrili, as well as its potential range in North America.

Successful establishment, spread, and impact of the introduced parasitoid Spathius galinae (Hymenoptera: Braconidae) on emerald ash borer (Coleoptera: Buprestidae) populations in postinvasion forests in Michigan.

Spathius galinae is a larval parasitoid native to the Russian Far East that was approved for release in the United States in 2015 for biological control of the emerald ash borer (EAB), Agrilus planipennis, an invasive beetle from Asia responsible for widespread mortality of ash trees (Fraxinus spp.) in North America. From 2015 to 2017, 1,340-1,445 females of S. galinae along with males were released into each release plot, paired with a nonrelease control plot (1-12.5 km apart), at 6 postinvasion forested sites containing abundant pole-sized ash trees in Michigan. By 2018, S. galinae had spread to all but one control plot. Based on the first year that S. galinae was found in trees in each control plot and the distances of those trees to the parasitoid release point within each site, we estimated that S. galinae spread at 3.7 (±1.9) km per year after its initial releases in 2015. The proportion of sampled trees with S. galinae broods, brood densities within sampled trees, and parasitism of EAB larvae increased sharply in both control and release plots after the last field releases in 2017, with the highest parasitism rates (42.8-60.3%) in 2020. Life table analysis showed that S. galinae alone reduced EAB's net population growth rate by 35-55% across sites from 2018 to 2020. These results demonstrate that S. galinae has established an increasing population in Michigan and now plays a significant role in reducing EAB populations in the area.

Efficacy of Fluon conditioning for capturing cerambycid beetles in different trap designs and persistence on panel traps over time.

Fluon PTFE is a fluoropolymer dispersion applied as a surface conditioner to cross-vane panel traps to enhance trap efficiency for cerambycid beetles. We describe the results of three experiments to further optimize cerambycid traps of different designs and to test the effect of Fluon over time. We tested Fluon with Lindgren funnel and panel traps fitted with either wet or dry collection cups on catches of cerambycid beetles and how the effect of Fluon on panel traps persisted. Fluon-treated funnel traps with wet collection cups captured approximately 6x more beetles than the untreated funnel traps with wet collection cups. Untreated funnel traps with dry collection cups did not capture any beetles; however, Fluon-treated funnel traps with dry collection cups captured an average of four beetles per trap. Fluon-treated panel traps with wet collection cups captured approximately 9x more beetles than untreated panel traps with wet collection cups. Fluon-treated panel traps with dry collection cups captured approximately 11x more beetles than untreated panel traps with dry collection cups. The effect of Fluon on capturing cerambycid beetles did not decline after use in one or two field seasons. There was no significant difference in the number of beetles captured in freshly treated panel traps compared with traps that had been used for 1 or 2 yr. Fluon-treated traps captured nine species that were not captured in untreated traps. Conditioning both Lindgren funnel and panel traps with Fluon enhances the efficacy and sensitivity of traps deployed to detect exotic cerambycid species, or for monitoring threatened species at low population densities.

A comparison of trap type and height for capturing cerambycid beetles (Coleoptera).

Wood-boring beetles in the family Cerambycidae (Coleoptera) play important roles in many forest ecosystems. However, increasing numbers of invasive cerambycid species are transported to new countries by global commerce and threaten forest health in the United States and worldwide. Our goal was to identify effective detection tools for a broad array of cerambycid species by testing some known cerambycid attractants and a pheromone in different trap designs placed across a range of habitats. We compared numbers and species richness of cerambycid beetles captured with cross-vane panel traps and 12-unit Lindgren multiple-funnel traps, placed either at ground level (1.5 m high) or canopy level (approximately 3-10 m high), at eight sites classified as either residential, industrial, deciduous forest, or conifer forest. We captured 3,723 beetles representing 72 cerambycid species from 10 June to 15 July 2010. Species richness was highest for the subfamilies Cerambycinae and Lamiinae, which accounted for 33 and 46% of all species captured, respectively. Overall, the cross-vane panel traps captured approximately 1.5 times more beetles than funnel traps. Twenty-one species were captured exclusively in traps at one height, either in the canopy or at ground level. More species were captured in hardwood sites (59 species) where a greater diversity of host material was available than in conifer (34 species), residential (41 species), or industrial (49) sites. Low numbers of beetles (n < 5) were recorded for 28 of the beetle species. The number of species captured per week ranged from 49 species on 21 June to 37 species on 12 July. Cross-vane panel traps installed across a vertical gradient should maximize the number of cerambycid species captured.

Differential response in foliar chemistry of three ash species to emerald ash borer adult feeding.

The emerald ash borer (EAB; Agrilus planipennis Fairmaire; Coleoptera: Buprestidae), is an exotic wood-boring beetle that has been threatening North American ash (Fraxinus spp.) resources since its discovery in Michigan and Ontario in 2002. In this study, we investigated the phytochemical responses of the three most common North American ash species (black, green, and white ash) in northeastern USA to EAB adult feeding. Black ash was the least responsive to EAB adult feeding in terms of the induction of volatile compounds, and levels of only two (indole and benzyl cyanide) of the 11 compounds studied increased. In green ash, levels of two [(E)-ß-ocimene and indole] of the 11 volatile compounds studied were elevated, while the levels of two green leaf volatiles [hexanal and (E)-2-hexenal] decreased. White ash showed the greatest response with an increase in levels of seven of the 11 compounds studied. Qualitative differences among ash species were detected. Among the phenolic compounds detected, ligustroside was the only one detected in all three species. Oleuropein aglycone and 2 unidentified compounds were found only in black ash; coumaroylquinic acid and feruloylquinic acid were detected only in green ash; and verbascoside hexoside was detected only in white ash. EAB adult feeding did not elicit or decrease concentrations of any selected individual phenolic compounds. However, although levels of total phenolics from black and green ash foliage were not affected by EAB adult feeding, they decreased significantly in white ash. EAB adult feeding elevated chymotrypsin inhibitors in black ash. The possible ecological implications of these findings are discussed.

Evaluation of double-decker traps for emerald ash borer (Coleoptera: Buprestidae).

Improved detection tools are needed for the emerald ash borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), an invasive forest insect from Asia that has killed millions of ash (Fraxinus spp.) trees in North America since its discovery in Michigan in 2002. We evaluated attraction of adult A. planipennis to artificial traps incorporating visual (e.g., height, color, silhouette) and olfactory cues (e.g., host volatiles) at field sites in Michigan. We developed a double-decker trap consisting of a 3-m-tall polyvinyl pipe with two purple prisms attached near the top. In 2006, we compared A. planipennis attraction to double-decker traps baited with various combinations of manuka oil (containing sesquiterpenes present in ash bark), a blend of four ash leaf volatiles (leaf blend), and a rough texture to simulate bark. Significantly more A. planipennis were captured per trap when traps without the rough texture were baited with the leaf blend and manuka oil lures than on traps with texture and manuka oil but no leaf blend. In 2007, we also tested single prism traps set 1.5 m above ground and tower traps, similar to double-decker traps but 6 m tall. Double-decker traps baited with the leaf blend and manuka oil, with or without the addition of ash leaf and bark extracts, captured significantly more A. planipennis than similarly baited single prism traps, tower traps, or unbaited double-decker traps. A baited double-decker trap captured A. planipennis at a field site that was not previously known to be infested, representing the first detection event using artificial traps and lures. In 2008, we compared purple or green double-decker traps, single prisms suspended 3-5 m above ground in the ash canopy (canopy traps), and large flat purple traps (billboard traps). Significantly more A. planipennis were captured in purple versus green traps, baited traps versus unbaited traps, and double-decker versus canopy traps, whereas billboard traps were intermediate. At sites with very low A. planipennis densities, more A. planipennis were captured on baited double-decker traps than on other traps and a higher percentage of the baited double-decker traps captured beetles than any other trap design. In all 3 yr, peak A. planipennis activity occurred during late June to mid-July, corresponding to 800-1200 growing degree-days base 10 degrees C (DD10). Nearly all (95%) beetles were captured by the end of July at approximately 1400 DD10.

Evaluation of Agrilus planipennis (Coleoptera: Buprestidae) control provided by emamectin benzoate and two neonicotinoid insecticides, one and two seasons after treatment.

Effective methods are needed to protect ash trees (Fraxinus spp.) from emerald ash borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), an invasive buprestid that has killed millions of North American ash (Fraxinus spp.) trees. We randomly assigned 175 ash trees (11.5-48.1 cm in diameter) in 25 blocks located in three study sites in Michigan to one of seven insecticide treatments in May 2007. Treatments included 1) trunk-injected emamectin benzoate; 2) trunk-injected imidacloprid; 3) basal trunk spray of dinotefuran with or 4) without Pentra-Bark, an agricultural surfactant; 5) basal trunk spray of imidacloprid with or 6) without Pentra-Bark; or (7) control. Foliar insecticide residues (enzyme-linked immunosorbent assay) and toxicity of leaves to adult A. planipennis (4-d bioassays) were quantified at 3-4-wk intervals posttreatment. Seven blocks of trees were felled and sampled in fall 2007 to quantify A. planipennis larval density. Half of the remaining blocks were retreated in spring 2008. Bioassays and residue analyses were repeated in summer 2008, and then all trees were sampled to assess larval density in winter. Foliage from emamectin benzoate-treated trees was highly toxic to adult A. planipennis, and larval density was < 1% of that in comparable control trees, even two seasons posttreatment. Larval densities in trees treated with trunk-injected imidacloprid in 2007 + 2008 were similar to control trees. Dinotefuran and imidacloprid were effectively translocated within trees treated with the noninvasive basal trunk sprays; the surfactant did not appreciably enhance A. planipennis control. In 2008, larval densities were 57-68% lower in trees treated with dinotefuran or imidacloprid trunk sprays in 2007 + 2008 than on controls, but densities in trees treated only in 2007 were similar to controls. Highly effective control provided by emamectin benzoate for > or = 2 yr may reduce costs or logistical issues associated with treatment.

Comparison of male and female emerald ash borer (Coleoptera: Buprestidae) responses to phoebe oil and (Z)-3-hexenol lures in light green prism traps.

We conducted trapping experiments for the emerald ash borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae) in Michigan, U.S.A., and Ontario, Canada, to compare unbaited light green sticky prism traps with traps baited with phoebe oil, (Z)-3-hexenol (Z3-6:OH), or blends of other green leaf volatiles (GLVs) with Z3-6:OH. Traps were placed in the lower canopy of ash trees (Fraxinus spp.). Catches with Z3-6:OH-baited traps showed a significant male bias and these traps caught significantly more males than the unbaited controls at both sites. They were also superior to phoebe oil-baited traps and those baited with GLV blends. Catches with phoebe oil showed a significant female bias but there was no difference in the number of females captured between traps baited with phoebe oil or Z3-6:OH lures. Catches were analyzed at regular time intervals to examine the response of A. planipennis to the lures over the course of the flight season. Z3-6:OH-baited traps consistently caught more males than the controls at each interval throughout the flight season. Catches of females with Z3-6:OH and phoebe oil were significantly better than the controls early in the flight season but declined to control levels by midseason. Our results suggest that Z3-6:OH-baited green traps placed in the ash canopy would be a superior lure for detecting and monitoring A. planipennis throughout the flight season.

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.

A Phenology Model for Simulating Oobius agrili (Hymenoptera: Encyrtidae) Seasonal Voltinism and Synchrony With Emerald Ash Borer Oviposition.

In North America, the emerald ash borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), continues to spread, and its egg parasitoid, Oobius agrili Zhang and Huang (Hymenoptera: Encyrtidae), is being released for emerald ash borer biocontrol well beyond their endemic climatic ranges in China. We developed a multiple cohort rate summation model to simulate O. agrili F0, F1, and F2 generations, and emerald ash borer oviposition for examining host-parasitoid synchrony across a north-south gradient from Duluth, MN (latitude 46.8369, longitude -92.1833) to Shreveport, LA (latitude 32.4469, longitude -93.8242). Temporal occurrences of critical day length for O. agrili diapause induction were integrated into the model. We used O. agrili and emerald ash borer trapping data from south central and northwestern Lower Michigan for model validation. Simulations demonstrated that 1) F0 adult emergence consistently occurred 2-5 d before emerald ash borer oviposition began; 2) F1 adult emergence was most synchronized with peak emerald ash borer oviposition compared with other generations; and 3) emerald ash borer oviposition was complete, or near so, when F2 adult emergence was predicted across the north-south gradient. Comparison of O. agrili trap captures with model simulations demonstrated that primarily two adult O. agrili generations (F0 and F1) emerged per year in Michigan and almost all F2 larvae entered diapause despite day lengths longer than critical day length in south central Michigan. Critical day length varied temporally across the north-south gradient during emergence of O. agrili generations. Determining day lengths perceived by O. agrili larvae in the field should improve model realism for examining spatiotemporal variation in O. agrili population dynamics.

Evaluation of Trapping Schemes to Detect Emerald Ash Borer (Coleoptera: Buprestidae).

Management responses to invasive forest insects are facilitated by the use of detection traps ideally baited with species-specific semiochemicals. Emerald ash borer, Agrilus planipennis Fairmaire, is currently invading North American forests, and since its detection in 2002, development of monitoring tools has been a primary research objective. We compared six trapping schemes for A. planipennis over 2 yr at sites in four U.S. states and one Canadian province that represented a range of background A. planipennis densities, canopy coverage, and ash basal area. We also developed a region-wide phenology model. Across all sites and both years, the 10th, 50th, and 90th percentile of adult flight occurred at 428, 587, and 837 accumulated degree-days, respectively, using a base temperature threshold of 10°C and a start date of 1 January. Most trapping schemes captured comparable numbers of beetles with the exception of purple prism traps (USDA APHIS PPQ), which captured significantly fewer adults. Trapping schemes varied in their trap catch across the gradient of ash basal area, although when considering trap catch as a binary response variable, trapping schemes were more likely to detect A. planipennis in areas with a higher ash component. Results could assist managers in optimizing trap selection, placement, and timing of deployment given local weather conditions, forest composition, and A. planipennis density.

Interactive influence of leaf age, light intensity, and girdling on green ash foliar chemistry and emerald ash borer development.

Biotic and abiotic environmental factors affect plant nutritional quality and defensive compounds that confer plant resistance to herbivory. Influence of leaf age, light availability, and girdling on foliar nutrition and defense of green ash (Fraxinus pennsylvanica Marsh) was examined in this study. Longevity of the emerald ash borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), adults reared on green ash foliage subjected to these factors was assayed. Mature leaves generally were more nutritious with greater amino acids and a greater ratio of protein to non-structural carbohydrate (P:C) than young leaves, in particular when trees were grown in shade. On the other hand, mature leaves had lower amounts of trypsin and chymotrypsin inhibitors, and total phenolics compared to young leaves. Lower defense of mature leaves alone, or along with higher nutritional quality may lead to increased survival and longevity of emerald ash borer feeding on mature leaves. Sunlight reduced amino acids and P:C ratio, irrespective of leaf age and girdling, and elevated total protein of young foliage, but not protein of mature leaves. Sunlight also dramatically increased all investigated defensive compounds of young, but not mature leaves. Girdling reduced green ash foliar nutrition, especially, of young leaves grown in shade and of mature leaves grown in sun. However emerald ash borer performance did not differ when fed leaves from trees grown in sun or shade, or from girdled or control trees. One explanation is that emerald ash borer reared on lower nutritional quality food may compensate for nutrient deficiency by increasing its consumption rate. The strong interactions among leaf age, light intensity, and girdling on nutrition and defense highlight the need for caution when interpreting data without considering possible interactions.

Host selection and feeding preference of Agrilus planipennis (Coleoptera: Buprestidae) on ash (Fraxinus spp.).

We studied the host selection behavior and feeding preference of the emerald ash borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae). A. planipennis is an exotic forest insect pest native to Asia that was discovered in North America in 2002 and is causing widespread mortality of ash trees (Fraxinus spp.) in southeast Michigan and surrounding states. We compared host selection and feeding behavior on different species of ash including Manchurian ash (F. mandshurica Rupr.), green ash (F. pennsylvanica Marsh), white ash (F. americana L.), black ash (F. nigra Marsh), blue ash (F. quadrangulata Michx.), and European ash (F. excelsior L). Manchurian ash is native to Asia, whereas the other species (native to North America or Europe) represent novel hosts for A. planipennis. Beetles distributed themselves more frequently and fed to a greater extent on green, black, and white ash compared with blue, European, and Manchurian ash. Although beetles consumed every ash species offered to them, Manchurian ash and blue ash were least preferred in feeding bioassays. When we analyzed the volatile content of intact and girdled ash for quantitative variation in 11 compounds that elicited antennal activity, we found that the overall volatile profiles of the six ash species differed significantly in their relative amounts of antennally active compounds. Green ash has lower relative amounts of volatiles compared with Manchurian ash, which might render it more attractive and less resistant to A. planipennis. Lower tolerance and resistance of green ash might make it more susceptible to mortality compared with Manchurian ash, which coevolved with the beetle in its native range. Repellent odors, potential antifeedants, and genes for resistance in Manchurian ash could be explored for methods to manage A. planipennis populations.

Emerald Ash Borer (Coleoptera: Buprestidae) Densities Over a 6-yr Period on Untreated Trees and Trees Treated With Systemic Insecticides at 1-, 2-, and 3-yr Intervals in a Central Michigan Forest.

We assessed density of emerald ash borer (Agrilus planipennis Fairmaire) (Coleoptera: Buprestidae) larvae over a 6-yr period by felling and sampling a total of 315 green ash (Fraxinus pennsylvanica Marsh.) trees that were left untreated or treated with imidacloprid, dinotefuran, or emamectin benzoate products at 1-yr, 2-yr, or 3-yr intervals. Our study, conducted across a 32-ha forested area, began soon after emerald ash borer became established and continued through the peak and eventual decline of the emerald ash borer population. Less than half of the 96 trees in the pretreatment sample were infested and larval densities were very low. Densities of emerald ash borer remained low for 3 yr, then increased exponentially, eventually resulting in mortality of most untreated overstory ash. Trees treated with either low or moderate rates of emamectin benzoate applied via trunk injection had few or no emerald ash borer galleries, even 3 yr post-treatment. Basal trunk sprays of dinotefuran applied annually were also effective at preventing larval densities from reaching damaging levels. Average larval densities on trees treated with a trunk injection of imidacloprid were lower but did not differ from untreated trees, regardless of treatment frequency. Larval parasitism was rare, while woodpecker predation was common and accounted for nearly all natural larval mortality, even on trees with very low densities of larvae.

Host range of the emerald ash borer (Agrilus planipennis Fairmaire) (Coleoptera: Buprestidae) in North America: results of multiple-choice field experiments.

Emerald ash borer (Agrilus planipennis Fairmaire) (Coleoptera: Buprestidae), an invasive phloem-feeding pest, was identified as the cause of widespread ash (Fraxinus) mortality in southeast Michigan and Windsor, Ontario, Canada, in 2002. A. planipennis reportedly colonizes other genera in its native range in Asia, including Ulmus L., Juglans L., and Pterocarya Kunth. Attacks on nonash species have not been observed in North America to date, but there is concern that other genera could be colonized. From 2003 to 2005, we assessed adult A. planipennis landing rates, oviposition, and larval development on North American ash species and congeners of its reported hosts in Asia in multiple-choice field studies conducted at several southeast Michigan sites. Nonash species evaluated included American elm (U. americana L.), hackberry (Celtis occidentalis L.), black walnut (J. nigra L.), shagbark hickory [Carya ovata (Mill.) K.Koch], and Japanese tree lilac (Syringa reticulata Bl.). In studies with freshly cut logs, adult beetles occasionally landed on nonash logs but generally laid fewer eggs than on ash logs. Larvae fed and developed normally on ash logs, which were often heavily infested. No larvae were able to survive, grow, or develop on any nonash logs, although failed first-instar galleries occurred on some walnut logs. High densities of larvae developed on live green ash and white ash nursery trees, but there was no evidence of larval survival or development on Japanese tree lilac and black walnut trees in the same plantation. We felled, debarked, and intensively examined >28 m2 of phloem area on nine American elm trees growing in contact with or adjacent to heavily infested ash trees. We found no sign of A. planipennis feeding on any elm.

Development of a host-based semiochemical lure for trapping emerald ash borer Agrilus planipennis (Coleoptera: Buprestidae).

Bark volatiles from green ash Fraxinus pennsylvanica were tested for electrophysiological activity by Agrilus planipennis using gas chromatographic-electroantennographic detection (GC-EAD) and for behavioral activity using baited purple traps in Michigan. GC-EAD analysis of the headspace volatiles of bark tissue samples from 0- and 24-h-old fully girdled (stressed) ash trees showed that the latter had elevated sesquiterpene levels. Six of the elevated compounds consistently elicited antennal responses by both male and female A. planipennis. Five of the antennally active compounds were identified as alpha-cubebene, alpha-copaene, 7-epi-sesquithujene, trans-beta-caryophyllene, and alpha-humulene (alpha-caryophyllene). The sixth EAD-active compound remains unidentified. We monitored capture of adult A. planipennis on traps baited with several combinations of ash tree volatiles. Treatments included two natural oil distillates (Manuka and Phoebe oil) that were found to contain, respectively, high concentrations of four and five of the six antennally active ash bark volatiles. A four-component leaf lure developed by the USDA Forest Service and Canadian Forest Service was also tested. In three separate field studies, Manuka oil-baited traps caught significantly more adult beetles than unbaited traps. Lures designed to release 5, 50, and 500 mg of Manuka oil per day all caught more insects than unbaited traps. In a field test comparing and combining Phoebe oil with Manuka oil, Phoebe oil-baited traps caught significantly more beetles than either Manuka oil-baited traps or unbaited traps. We hypothesize that the improved attractancy of Phoebe oil to A. planipennis over Manuka oil is caused by the presence of the antennally active sesquiterpene, 7-epi-sesquithujene.