Inoculative Releases and Natural Spread of the Fungal Pathogen Entomophaga maimaiga (Entomophthorales: Entomophthoraceae) into U.S. Populations of Gypsy Moth, Lymantria dispar (Lepidoptera: Erebidae).

While emphasis with entomopathogens has often been on inundative releases, we describe here historic widespread inoculative releases of a fungal entomopathogen. Several U.S. states and municipalities conducted inoculative releases of the gypsy moth, Lymantria dispar (L.) (Lepidoptera: Erebidae), pathogen Entomophaga maimaiga Humber, Shimazu et Soper (Entomophthorales: Entomophthoraceae) after 1993, as gypsy moth populations spread into the Midwest and North Carolina. This Japanese pathogen first caused epizootics in northeastern North America in 1989 and methods for its inoculative release were tested and proven to be effective from 1991 to 1993. After 1993, spores in soil or in late instar cadavers were collected during or after epizootics and were released inoculatively into newly established populations of this spreading invasive; the goal was that spores would overwinter and germinate the next spring to infect larvae, thus speeding pathogen spread and hastening the development of epizootics in newly established populations. The fungus was released in gypsy moth populations that were separated from areas where the fungus was already established. In particular, extensive releases by natural resource managers in Wisconsin and Michigan aided the spread of E. maimaiga throughout these states. Where it has become established, this acute pathogen has become the dominant natural enemy and has exerted considerable influence in reducing gypsy moth damage. While this pathogen most likely would have invaded these new regions eventually, releases accelerated the spread of E. maimaiga and helped to reduce impacts of initial outbreaks, while further outbreaks were reduced by the pathogen's subsequent persistence and activity in those areas.

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.

Optimizing Use of Girdled Ash Trees for Management of Low-Density Emerald Ash Borer (Coleoptera: Buprestidae) Populations.

Effective survey methods to detect and monitor recently established, low-density infestations of emerald ash borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), remain a high priority because they provide land managers and property owners with time to implement tactics to slow emerald ash borer population growth and the progression of ash mortality. We evaluated options for using girdled ash (Fraxinus spp.) trees for emerald ash borer detection and management in a low-density infestation in a forested area with abundant green ash (F. pennsylvanica). Across replicated 4-ha plots, we compared detection efficiency of 4 versus 16 evenly distributed girdled ash trees and between clusters of 3 versus 12 girdled trees. We also examined within-tree larval distribution in 208 girdled and nongirdled trees and assessed adult emerald ash borer emergence from detection trees felled 11 mo after girdling and left on site. Overall, current-year larvae were present in 85-97% of girdled trees and 57-72% of nongirdled trees, and larval density was 2-5 times greater on girdled than nongirdled trees. Low-density emerald ash borer infestations were readily detected with four girdled trees per 4-ha, and 3-tree clusters were as effective as 12-tree clusters. Larval densities were greatest 0.5 ± 0.4 m below the base of the canopy in girdled trees and 1.3 ± 0.7 m above the canopy base in nongirdled trees. Relatively few adult emerald ash borer emerged from trees felled 11 mo after girdling and left on site through the following summer, suggesting removal or destruction of girdled ash trees may be unnecessary. This could potentially reduce survey costs, particularly in forested areas with poor accessibility.

Colonization of Three Maple Species by Asian Longhorned Beetle, Anoplophora glabripennis, in Two Mixed-Hardwood Forest Stands.

Asian longhorned beetle (ALB), Anoplophora glabripennis (Motschulsky), is an invasive insect that has successfully established multiple times in North America. To investigate host colonization and reproductive success (exit holes/eggs), two ALB infested forest stands were sampled in central Massachusetts, USA. Infested Acer platanoides L., Acer rubrum L., and Acer saccharum Marsh. were felled, bucked into 1 m sections and dissected to determine indications of ALB infestations, such as presence of life stages or signs of damage on trees. ALB damage was also aged on a subset of trees to determine the earliest attacks on the three Acer species. In one stand, ALB oviposition was significantly higher on the native A. rubrum and A. saccharum than the exotic A. platanoides. In the second stand, ALB oviposition was significantly higher and cumulative reproductive success was higher on A. rubrum than A. platanoides or A. saccharum. An A. saccharum had the earliest signs of attack that occurred in 2006. Acer rubrum (2007) and A. platanoides (2010) were colonized shortly thereafter. Overall, ALB was more successful in A. rubrum, where adults emerged from 53% and 64% of trees in each stand, compared to A. platanoides (11% and 18%) or A. saccharum (14% and 9%).

Evaluation of potential versus realized primary infection of gypsy moth (Lepidoptera: Lymantriidae) by Entomophaga maimaiga (Zygomycetes: Entomophthorales).

The fungal entomopathogen Entomophaga maimaiga has provided important biological control of gypsy moth, Lymantria dispar (L.), since the first epizootics occurred in the northeastern United States in 1989. Epizootics are initiated by germination of soil-borne resting spores, which are highly sensitive to spring temperature and moisture. We compared gypsy moth infection by E. maimaiga in 33 oak stands in Michigan with infection under optimal laboratory conditions from 1999 to 2001 to assess differences between potential and realized efficacy of E. maimaiga. Field bioassays were conducted by exposing laboratory-reared, fourth-instar gypsy moth to soil at the base of oak trees for 4 d. Additional larvae were similarly exposed to soil collected from the field plots in laboratory bioassays with temperature, humidity, and moisture levels optimal for fungal germination. Overall E. maimaiga infection ranged from means of 3.2-29.8% in the field compared with 20.9-59.7% in the laboratory during three field seasons. Resting spore density in soil and gypsy moth egg mass density were significant predictors of field infections in two of the 3 yr, whereas resting spore density was a significant predictor of laboratory infections each year. Other variables that significantly predicted laboratory infections in one of the 3 yr included egg mass density, canopy cover, and soil pH. In laboratory bioassays, soil pH and E. maimaiga resting spore density were positively associated with increasing E. maimaiga infection rates of gypsy moth larvae.

The influence of satellite populations of emerald ash borer on projected economic costs in U.S. communities, 2010-2020.

The invasion spread of the emerald ash borer (Agrilus planipennis Fairmaire) (Coleoptera: Buprestidae) is characterized by the formation of satellite populations that expand and coalesce with the continuously invading population front. As of January 2010, satellite infestations have been detected in 13 states and two Canadian provinces. Understanding how newly established satellite populations may affect economic costs can help program managers to justify and design prevention and control strategies. We estimate the economic costs caused by EAB for the 10-yr period from 2010 to 2020 for scenarios of fewer EAB satellite populations than those found from 2005 to 2010 and slower expansion of satellite populations found in 2009. We measure the projected discounted cost of treatment, removal, and replacement of ash trees (Fraxinus spp.) growing in managed landscapes in U.S. communities. Estimated costs for the base scenario with the full complement of satellites in 2005-2010 and no program to mitigate spread is $12.5 billion. Fewer EAB satellites from 2005 to 2010 delay economic costs of $1.0 to 7.4 billion. Slower expansion of 2009 satellite populations delays economic costs of $0.1 to 0.7 billion. Satellite populations that are both distant from the core EAB infestation and close to large urban areas caused more economic costs in our simulations than did other satellites. Our estimates of delayed economic costs suggest that spending on activities that prevent establishment of new satellite EAB populations or slow expansion of existing populations can be cost-effective and that continued research on the cost and effectiveness of prevention and control activities is warranted.

Effects of trap type, placement and ash distribution on emerald ash borer captures in a low density site.

Effective methods for early detection of newly established, low density emerald ash borer (Agrilus planipennis Fairmaire) infestations are critically needed in North America. We assessed adult A. planipennis captures on four types of traps in a 16-ha site in central Michigan. The site was divided into 16 blocks, each comprised of four 50- by 50-m cells. Green ash trees (Fraxinus pennsylvanica Marshall) were inventoried by diameter class and ash phloem area was estimated for each cell. One trap type was randomly assigned to each cell in each block. Because initial sampling showed that A. planipennis density was extremely low, infested ash logs were introduced into the center of the site. In total, 87 beetles were captured during the summer. Purple double-decker traps baited with a blend of ash leaf volatiles, Manuka oil, and ethanol captured 65% of all A. planipennis beetles. Similarly baited, green double-decker traps captured 18% of the beetles, whereas sticky bands on girdled trees captured 11% of the beetles. Purple traps baited with Manuka oil and suspended in the canopies of live ash trees captured only 5% of the beetles. At least one beetle was captured on 81% of the purple double-decker traps, 56% of the green double-decker traps, 42% of sticky bands, and 25% of the canopy traps. Abundance of ash phloem near traps had no effect on captures and trap location and sun exposure had only weak effects on captures. Twelve girdled and 29 nongirdled trees were felled and sampled in winter. Current-year larvae were present in 100% of the girdled trees and 72% of the nongirdled trees, but larval density was five times higher on girdled than nongirdled trees.

Estimating potential emerald ash borer (Coleoptera: Buprestidae) populations using ash inventory data.

Emerald ash borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), a phloem-feeding pest native to Asia, was identified in June 2002 as the cause of widespread ash (Fraxinus spp.), mortality in southeastern Michigan and Windsor, Ontario, Canada. Localized populations of A. planipennis have since been found across lower Michigan and in areas of Ohio, Indiana, Illinois, Maryland, and Ontario. Officials working to contain A. planipennis and managers of forestlands near A. planipennis infestations must be able to compare alternative strategies to allocate limited funds efficiently and effectively. Empirical data from a total of 148 green ash, Fraxinus pennsylvanica Marsh., and white ash, Fraxinus americana L., trees were used to develop models to estimate surface area of the trunk and branches by using tree diameter at breast height (dbh). Data collected from 71 additional F. pennsylvanica and F. americana trees killed by A. planipennis showed that on average, 88.9 +/- 4.6 beetles developed and emerged per m2 of surface area. Models were applied to ash inventory data collected at two outlier sites to estimate potential production of A. planipennis beetles at each site. Large trees of merchantable size (dbh > or = 26 cm) accounted for roughly 6% of all ash trees at the two sites, but they could have contributed 55-65% of the total A. planipennis production at both sites. In contrast, 75- 80% of the ash trees at the outlier sites were < or =13 cm dbh, but these small trees could have contributed only < or =12% of the potential A. planipennis production at both sites. Our results, in combination with inventory data, can be used by regulatory officials and resource managers to estimate potential A. planipennis production and to compare options for reducing A. planipennis density and slowing the rate of spread for any area of interest.