Deborah G. McCullough

Emerald Ash Borer Invasion of North America: History, Biology, Ecology, Impacts, and Management

Since its accidental introduction from Asia, emerald ash borer (EAB), Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), has killed millions of ash trees in North America. As it continues to spread, it could functionally extirpate ash with devastating economic and ecological impacts. Little was known about EAB when it was first discovered in North America in 2002, but substantial advances in understanding of EAB biology, ecology, and management have occurred since. Ash species indigenous to China are generally resistant to EAB and may eventually provide resistance genes for introgression into North American species. EAB is characterized by stratified dispersal resulting from natural and human-assisted spread, and substantial effort has been devoted to the development of survey methods. Early eradication efforts were abandoned largely because of the difficulty of detecting and delineating infestations. Current management is focused on biological control, insecticide protection of high-value trees, and integrated efforts to slow ash mortality.

Economic Impacts of Non-Native Forest Insects in the Continental United States

Reliable estimates of the impacts and costs of biological invasions are critical to developing credible management, trade and regulatory policies. Worldwide, forests and urban trees provide important ecosystem services as well as economic and social benefits, but are threatened by non-native insects. More than 450 non-native forest insects are established in the United States but estimates of broad-scale economic impacts associated with these species are largely unavailable. We developed a novel modeling approach that maximizes the use of available data, accounts for multiple sources of uncertainty, and provides cost estimates for three major feeding guilds of non-native forest insects. For each guild, we calculated the economic damages for five cost categories and we estimated the probability of future introductions of damaging pests. We found that costs are largely borne by homeowners and municipal governments. Wood- and phloem-boring insects are anticipated to cause the largest economic impacts by annually inducing nearly

Historical Accumulation of Nonindigenous Forest Pests in the Continental United States

1.7 billion in local government expenditures and approximately

Arrival rate of nonindigenous insect species into the United States through foreign trade

830 million in lost residential property values. Given observations of new species, there is a 32% chance that another highly destructive borer species will invade the U.S. in the next 10 years. Our damage estimates provide a crucial but previously missing component of cost-benefit analyses to evaluate policies and management options intended to reduce species introductions. The modeling approach we developed is highly flexible and could be similarly employed to estimate damages in other countries or natural resource sectors.

Interceptions of nonindigenous plant pests at US ports of entry and border crossings over a 17-year period

Nonindigenous forest insects and pathogens affect a range of ecosystems, industries, and property owners in the United States. Evaluating temporal patterns in the accumulation of these nonindigenous forest pests can inform regulatory and policy decisions. We compiled a comprehensive species list to assess the accumulation rates of nonindigenous forest insects and pathogens established in the United States. More than 450 nonindigenous insects and at least 16 pathogens have colonized forest and urban trees since European settlement. Approximately 2.5 established nonindigenous forest insects per year were detected in the United States between 1860 and 2006. At least 14% of these insects and all 16 pathogens have caused notable damage to trees. Although sap feeders and foliage feeders dominated the comprehensive list, phloem- and wood-boring insects and foliage feeders were often more damaging than expected. Detections of insects that feed on phloem or wood have increased markedly in recent years.

Host Range of the Emerald Ash Borer (Agrilus planipennis Fairmaire) (Coleoptera: Buprestidae) in North America: Results of Multiple-Choice Field Experiments

Introductions of invasive nonindigenous species, and the ensuing negative ecological and economic consequences, have increased with expanding global trade. Quantifying the influx of nonindigenous plant pest species through foreign trade is required for national and international risk assessments, monitoring and conservation efforts, and evaluation of ecological factors that affect invasion success. Here we use statistically robust data collected at US ports of entry and border crossings to estimate arrival rates of nonindigenous insect species via four cargo pathways and to evaluate the effectiveness of current efforts to monitor arrival of nonindigenous insect species. Interception rates were highest in refrigerated maritime cargo where a new insect species was intercepted on average every 54 inspections. Projected estimates of insect species richness stabilized only for non-refrigerated maritime cargo and US–Mexico border cargo, where inspectors likely detected 19–2% and 30–50% of the species being transported through these respective pathways. Conservative estimates of establishment suggest that 42 insect species may have become established through these four pathways between 1997 and 2001.

Dispersal of Agrilus planipennis (Coleoptera: Buprestidae) From Discrete Epicenters in Two Outlier Sites

Despite the substantial impacts of nonindigenous plant pests and weeds, relatively little is known about the pathways by which these organisms arrive in the U.S. One source of such information is the Port Information Network (PIN) database, maintained by the U.S. Department of Agriculture, Animal and Plant Health Inspection Service (APHIS) since 1984. The PIN database is comprised of records of pests intercepted by APHIS personnel during inspections of travelers’ baggage, cargo, conveyances and related items arriving at U.S. ports of entry and border crossings. Each record typically includes the taxonomic identify of the pest, its country of origin, and information related to the commodity and interception site. We summarized more than 725,000 pest interceptions recorded in PIN from 1984 to 2000 to examine origins, interception sites and modes of transport for nonindigenous insects, mites, mollusks, nematodes, plant pathogens and weeds. Roughly 62% of intercepted pests were associated with baggage, 30% were associated with cargo and 7% were associated with plant propagative material. Pest interceptions occurred most commonly at airports (73%), U.S.-Mexico land border crossings (13%) and marine ports (9%). Insects dominated the database, comprising 73 to 84% of the records annually, with the orders Homoptera, Lepidoptera and Diptera collectively accounting for over 75% of the insect records. Plant pathogens, weeds and mollusks accounted for 13, 7 and 1.5% of all pest records, respectively, while mites and nematodes comprised less than 1% of the records. Pests were intercepted from at least 259 different locations. Common origins included Mexico, Central and South American countries, the Caribbean and Asia. Within specific commodity pathways, richness of the pest taxa generally increased linearly with the number of interceptions. Application of PIN data for statistically robust predictions is limited by nonrandom sampling protocols, but the data provide a valuable historical record of the array of nonindigenous organisms transported to the U.S. through international trade and travel.

Nonnative forest insects and pathogens in the United States: Impacts and policy options

Abstract 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.

Estimating Potential Emerald Ash Borer (Coleoptera: Buprestidae) Populations Using Ash Inventory Data

ABSTRACT Emerald ash borer, Agrilus planipennis (Fairmaire) (Coleoptera: Buprestidae), a phloem-feeding beetle native to Asia, has become one of the most destructive forest pests in North America. Since it was first identified in 2002 in southeast Michigan and Windsor, Ontario, dozens of isolated A. planipennis populations have been discovered throughout Michigan and Ontario, and in 12 other states and the province of Quebec. We assessed realized A. planipennis dispersal at two discrete outlier sites that originated 1 yr and 3 yr earlier from infested nursery trees. We systematically sampled ash trees within an 800 m radius of the origin of each infestation to locate galleries constructed by the progeny of dispersing A. planipennis adults. Our sampling identified eight trees at the 1 yr site infested with a mean ± SE of 11.6 ± 8.4 A. planipennis larvae and 12 trees at the 3 yr site with 25.8 ± 11.1 larvae per meter squared. Dendroentomological analysis indicated that A. planipennis populations were predominantly undergoing a 2 yr (semivoltine) life cycle at both sites. Colonized trees were found out to 638 and 540 m from the epicenters at the 1 yr and 3 yr sites, respectively. Logistic regression was used to determine whether the likelihood of A. planipennis colonization was affected by wind direction, ash phloem abundance, distance from the epicenter, or land-use type (i.e., wooded, residential, agricultural, or urban). Results show that the probability of A. planipennis colonization was significantly affected by ash phloem abundance and decreased with distance from the epicenter.

Emerald ash borer (Coleoptera: Buprestidae) attraction to stressed or baited ash trees

Abstract We review and synthesize information on invasions of nonnative forest insects and diseases in the United States, including their ecological and economic impacts, pathways of arrival, distribution within the United States, and policy options for reducing future invasions. Nonnative insects have accumulated in United States forests at a rate of ~2.5 per yr over the last 150 yr. Currently the two major pathways of introduction are importation of live plants and wood packing material such as pallets and crates. Introduced insects and diseases occur in forests and cities throughout the United States, and the problem is particularly severe in the Northeast and Upper Midwest. Nonnative forest pests are the only disturbance agent that has effectively eliminated entire tree species or genera from United States forests within decades. The resulting shift in forest structure and species composition alters ecosystem functions such as productivity, nutrient cycling, and wildlife habitat. In urban and suburban areas, loss of trees from streets, yards, and parks affects aesthetics, property values, shading, stormwater runoff, and human health. The economic damage from nonnative pests is not yet fully known, but is likely in the billions of dollars per year, with the majority of this economic burden borne by municipalities and residential property owners. Current policies for preventing introductions are having positive effects but are insufficient to reduce the influx of pests in the face of burgeoning global trade. Options are available to strengthen the defenses against pest arrival and establishment, including measures taken in the exporting country prior to shipment, measures to ensure clean shipments of plants and wood products, inspections at ports of entry, and post‐entry measures such as quarantines, surveillance, and eradication programs. Improved data collection procedures for inspections, greater data accessibility, and better reporting would support better evaluation of policy effectiveness. Lack of additional action places the nation, local municipalities, and property owners at high risk of further damaging and costly invasions. Adopting stronger policies to reduce establishments of new forest insects and diseases would shift the major costs of control to the source and alleviate the economic burden now borne by homeowners and municipalities.