describe the case study of the asian longhorned beetle

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Asian longhorned beetle

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• The Nature Conservancy - Asian Longhorned Beetle

Asian longhorned beetle , ( Anoplophora glabripennis ), species of beetle (order Coleoptera , family Cerambycidae), originally native to eastern China and Korea, that became a serious pest of hardwood trees in North America and parts of Eurasia.

The glossy black adults are large, 17–40 mm (0.7–1.6 inches) in length, and have 10–20 white or yellow-orange irregular spots on their smooth elytra (wing covers). The long antennae each have 11 segments and are 1.5 (female) to 2 (male) times as long as the body. The base of each segment is pale blue-white, grading distally (away from the centre of the body) to black.

The beetle’s life cycle lasts one to two years. Adults are active from April or May through October. New adults feed on twigs or leaf veins and petioles (leaf stalks) for approximately two weeks before mating. Adults locate host trees by using visual or chemical cues and detect mates by using both short-range and contact pheromones . Adult females have a life span of about 66 days, during which they can lay between 50 and 125 individual eggs, depending on their geographic strain, the host trees available, and exposure to pathogens in the environment . Mated females chew a pit in the bark on the upper trunk or main branches of a host tree and deposit a single 6-mm- (0.2-inch-) long white egg under the bark. The egg hatches in 7 to 14 days. The legless larva creates a feeding tunnel by chewing through the cambium (layer of actively dividing cells) into sapwood and heartwood . A feeding larva excretes sawdustlike waste, which is pushed out through the tunnel opening. When fully developed, the larva is 30–50 mm (1.2–2 inches) long and pupates at the end of the feeding tunnel. The adult emerges by chewing to the surface and exiting through a 10–15-mm- (0.4–0.6-inch-) diameter hole.

In its native environment on the Korean peninsula, the Asian longhorned beetle occurs at low densities at the edge of mixed forest habitats . Given their low numbers and the limited availability of host trees at the forest edge, the beetles do not significantly damage trees in their native environment.

In China the beetle’s native range was in the eastern portion of the country until the mid-1980s, when it was first reported in high numbers in western China and began killing significant numbers of trees. That population increase and westward movement followed the large-scale planting of poplar trees that began in the 1960s and culminated as part of China’s Three-North Shelterbelt Programme, initiated in 1978. The program’s goal was to plant a 4,506-km- (2,800-mile-) long shelterbelt of trees in the northwest regions by 2050 to prevent soil erosion , slow desertification , enhance urban beautification, and increase pulpwood production. The dramatic increase in host trees allowed the Asian longhorned beetle to become a serious pest in China.

In 1992 the Asian longhorned beetle was first detected at ports of entry on the east coasts of both the United States and Canada , but it was exterminated before it could escape into the surrounding habitats. The first established population outside Asia was found in New York City in 1996. Subsequent populations have been reported in New Jersey , Illinois , Ohio , and Massachusetts , though the infestations in both Illinois and New Jersey have since been eradicated . Established populations have also been reported in Japan , Austria , France , Germany , and Italy . Transport of the Asian longhorned beetle to North America, Europe , and Japan occurred primarily in solid-wood packing materials (e.g., pallets and packing crates) containing developing larvae or pupae . In rare instances the beetle has been found in shipments of live plants.

The Asian longhorned beetle can develop in at least 15 tree genera, its preferred hosts being species of poplar , maple , willow , and elm . Larval feeding is the primary cause of tree damage, as tunneling in the cambium disrupts vascular flow. In trees that have been repeatedly infested, limbs or trunks often break under high winds or heavy snow, as numerous feeding tunnels weaken the tree.

Because the Asian longhorned beetle can inflict significant tree damage, all countries in which it has been accidentally introduced have established eradication protocols . Around an infestation an eradication zone is created, in which all infested trees are removed and all uninfested host trees are treated with a systemic insecticide . In areas beyond the eradication zone, potential host trees are regularly inspected for signs of infestation. Infestations are considered eradicated only after no new infested trees are found following four to six years of inspections.

Eradication efforts in the United States between 1997 and 2010 cost more than $373 million. More than 110,000 trees have been removed since 1996 as part of the eradication programs in New York , Illinois, New Jersey, Massachusetts, and Ohio, and pesticides were applied on a large scale to susceptible species. In an attempt to reduce the spread of the pest, citizens in many areas are urged to report infestations or sightings of the beetle and are warned not to transport firewood or other potentially contaminated wood items.

Other management strategies being examined for the Asian longhorned beetle include studies to locate natural enemies that can manage the pest as a form of biological control . The natural enemies include fungi and nematodes as well as insect predators and parasitoids . Given the inherent risks involved in introducing yet another foreign species, the natural enemies are still being researched to determine their suitability for biological control.

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• Terrestrial Invasives
• Terrestrial Invertebrates

Asian Long-Horned Beetle

Anoplophora glabripennis (Motschulsky, 1853) ( ITIS )

Asian long-horned beetle (ALB), starry sky beetle

Asia ( Hu et al. 2009 )

First breeding populations discovered in New York in 1996 ( Haack et al. 2010 )

Arrived accidentally in cargo from Asia ( Hu et al. 2009 )

Destructive wood-boring pest of maple and other hardwoods ( Haack et al. 2010 )

Asian Longhorned Beetle, adult

Photo by Donald Duerr; USDA, Forest Service

Find more images

• Google Images - Asian Longhorned Beetle
• Invasive.org - Asian Longhorned Beetle

USDA APHIS Asks for Help Looking for Asian Longhorned Beetle by Checking Trees

• Jul 29, 2024

USDA . Animal and Plant Health Inspection Service.

The U.S. Department of Agriculture’s Animal and Plant Health Inspection Service (APHIS) is asking the public to look for and report the Asian longhorned beetle (ALB). APHIS declares every August Tree Check Month and is asking you to look for this destructive, invasive beetle by checking trees on your property and in your community for damage. Left unchecked, the ALB can cause infested trees to die. August is an ideal time of year to look for the beetle and the damage it causes.

State Asks Public to Check Trees for Invasive Pests this August

• Aug 6, 2024

Washington Invasive Species Council.

Washington state agencies are asking for your help to check your trees for signs and symptoms of invasive insects. Damaging invasive insect species, such as spotted lanternfly, longhorned beetles, and emerald ash borer, are emerging in their adult form. ate summer is often the peak time for these invasive insects to emerge from trees in their adult stage. If you see or suspect you see an invasive insect, report a sighting .

USDA APHIS Makes Gains Removing Asian Longhorned Beetle in New York

• Jan 31, 2024

The U.S. Department of Agriculture’s (USDA) Animal and Plant Health Inspection Service (APHIS), together with the New York State Department of Agriculture and Markets, is announcing that the Asian longhorned beetle (ALB) quarantine on Long Island is now smaller. New York is now closer to being ALB -free.

Plant Protection Today - PPQ's Ohio ALB Eradication Staff Continue to Win the Beetle Battle

• May 31, 2022

USDA . APHIS . Plant Protection Today.

Last month PPQ's Asian Longhorned Beetle (ALB) Eradication Program in Ohio celebrated another victory—the ALB quarantine is officially 7.5 square miles smaller! This invasive beetle from Asia is a destructive wood-boring pest that feeds on maple and other hardwoods, eventually killing them. After completing their final round of tree inspection surveys, the ALB staff reported no sign of the beetle in a portion of East Fork State Park in Clermont County, Ohio.

"Inspecting ALB host trees is painstaking work, and the staff meticulously survey for the pest," said ALB National Policy Manager Kathryn Bronsky. "It’s been three years since the last time we've lifted ALB quarantine restrictions in Ohio, and this is the first removal of the initial area placed under quarantine. That makes this success especially gratifying."

Distribution / Maps / Survey Status

Alien forest pest explorer (afpe).

USDA . FS . Northern Research Station.

The Alien Forest Pest Explorer (AFPE) is an interactive web tool which provides detailed spatial data describing pest distributions and host inventory estimates for damaging, non-indigenous forest insect and disease pathogens currently established in the U.S. The database is maintained as a joint effort of Purdue University, the U.S. Forest Service Northern Research Station, and the U.S. Forest Service Forest Health Protection.

Early Detection & Distribution Mapping System (EDDMapS) - Asian Longhorned Beetle

University of Georgia. Center for Invasive Species and Ecosystem Health.

Provides state, county, point and GIS data. Maps can be downloaded and shared.

Pest Tracker - Survey Status of Asian Longhorned Beetle

USDA . APHIS . Cooperative Agricultural Pest Survey. National Agricultural Pest Information System.

Federally Regulated

Domestic quarantine notices (title 7: agriculture, part 301) - asian longhorned beetle.

U.S. Government Printing Office. Electronic Code of Federal Regulations.

Federal Asian Longhorned Beetle Quarantine Boundary Viewer

See related resource: Data Visualization Tools to explore plant and animal health management data and interactive story maps

Import Federal Orders

USDA . APHIS . Plant Protection and Quarantine.

A Federal Order is a legal document issued in response to an emergency when the Administrator of APHIS considers it necessary to take regulatory action to protect agriculture or prevent the entry and establishment into the United States of a pest or disease. Federal Orders are effective immediately and contain the specific regulatory requirements.

YouTube - USDA Asian Longhorned Beetle PSA

Google. YouTube; USDA . Animal and Plant Health Inspection Service.

YouTube - USDA Asian Longhorned Beetle Videos

Google. YouTube; United States Department of Agriculture.

YouTube - Outsmart Invasive Species Project: Asian Longhorned Beetle

Google. YouTube; University of Massachusetts - Amherst. 

All Resources

Selected resources.

The section below contains highly relevant resources for this species, organized by source.

Priority Species: Citrus, Asian, and Red-Necked Longhorned Beetles

Washington State Recreation and Conservation Office. Washington Invasive Species Council.

Asian Longhorned Cooperative Eradication Program

Massachusetts Introduced Pests Outreach Project.

EPPO Global Database - Anoplophora glabripennis

European and Mediterranean Plant Protection Organization.

Gallery of Pests - Asian Longhorned Beetle

Nature Conservancy. Don't Move Firewood.

Global Invasive Species Database - Anoplophora glabripennis (insect)

IUCN . Species Survival Commission. Invasive Species Specialist Group.

Invaders Factsheet: Asian Long-horned Beetle

Ontario's Invading Species Awareness Program (Canada).

Invasive Species Compendium - Anoplophora glabripennis

CAB International.

New York Invasive Species Information - Asian Longhorned Beetle

New York Invasive Species Clearinghouse.

NHBugs - Asian Longhorned Beetle

University of New Hampshire. Cooperative Extension; New Hampshire Department of Agriculture, Markets & Food.

The Asian Longhorned Beetle (ALB) was found in Worcester, MA in August 2008 and in Boston in July 2010. This insect pest poses a serious risk to trees and forests. ALB has not yet been found in New Hampshire. Help us by looking at the debris from your swimming pools. Whenever you clean your pool, look at the debris you collect in your filter and skimmers. Use this fact sheet [PDF, 1.22 MB] to compare collected insects to common insects. Upload pictures of any insect you think is a longhorned beetle.

Plantwise Technical Factsheet - Asian Longhorned Beetle ( Anoplophora glabripennis )

CABI . PlantwisePlus Knowledge Bank.

Regional Pest Alert: Asian Longhorned Beetle

North Central Integrated Pest Management Center.

See also: Pest Alerts for more resources

Southern Forest Health - Asian Longhorned Beetle

USDA . Forest Service; Southern Regional Extension Forestry. Forest Health Program.

Includes species related publications, webinars and other resources.

APHIS Campaign: Check Trees for Asian Longhorned Beetle

Public outreach and educational site (former Asianlonghornedbeetle.com site).

Asian Longhorned Beetle

Provides comprehensive Asian longhorned beetle (ALB) information including: what to look for, how to prevent this pest and how it is treated. Also provides image gallery and information how to report signs of pest. And provides information for ALB cooperative eradication including: current status and quarantine information, APHIS ' response, regulatory information, federal orders, information for cooperators and producers, ALB research, and reports and assessments.

Forest Invasive Alien Species - Asian Longhorned Beetle

Natural Resources Canada. Canadian Forest Service.

Invasive Species - Asian Longhorned Beetle

Canadian Food Inspection Agency.

Forest Pests: Invasive Plants and Insects of Maryland - Asian Longhorned Beetle [PDF, 279 KB]

Maryland Department of Natural Resources. Forest Service.

See also: Invasive Plants and Insects Fact Sheets for additional species to help control invasive species in Maryland.

Fact Sheet: Asian Longhorned Beetle [PDF, 108 KB]

Connecticut Agricultural Experiment Station.

See also: Insect Factsheets for more resources

Field Guide: Invasive - Asian Longhorned Beetle

Missouri Department of Conservation.

See also: For more information about Invasive Tree Pests (insects and diseases) that are not native to Missouri

Identify and Report - Asian Longhorned Beetle

Michigan.gov. Michigan Invasive Species Program.

Insects & Pests - Asian Longhorned Beetle

Minnesota Department of Agriculture.

Pest Profile - Asian Longhorned Beetle

California Department of Food and Agriculture. Plant Health Division. Pest Detection/Emergency Projects Branch.

Plant Pests - Asian Longhorned Beetle

Ohio Department of Agriculture. Plant Health.

Utah Pests Fact Sheet - Asian Longhorned Beetle [PDF, 2.59 MB]

Utah State University Extension; Utah Plant Pest Diagnostic Laboratory.

See also: Invasive Species for more fact sheets

Asian Long-Horned Beetle, Anoplophora glabripennis

University of California - Riverside. Center for Invasive Species Research.

Emerging Threats: Asian Longhorned Beetle

Kansas State University. Kansas Forest Service.

Insects and Arthropods - Asian Longhorned Beetle

Oklahoma State University. Extension.

Introduced Species Summary Project - Asian Longhorned Beetle

Columbia University. Center for Environmental Research and Conservation.

Solve a Problem: Asian Longhorn Beetle

Morton Arboretum (Illinois).

Haack, R.A., F. Herard, J. Sun, and J.J. Turgeon. 2010. Managing invasive populations of Asian longhorned beetle and citrus longhorned beetle: a worldwide perspective . Annual Review of Entomology 55:521-546.

Hu, J., S. Angeli, S. Schuetz, Y. Luo, and A.E. Hajek. 2009. Ecology and management of exotic and endemic Asian longhorned beetle Anoplophora glabripennis . Agricultural and Forest Entomology 11(4):359-375.

Integrated Taxonomic Information System. Anoplophora glabripennis . [Accessed Mar 12, 2023].

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• Research Highlights
• Inside the Forest Service

Asian longhorned beetle has broad climate adaptability and invasion potential

• Partnerships

Asian longhorned beetle infestations have been found in urban and suburban trees or forested areas as far north as Helsinki, Finland, and as far south as Bethel, Ohio. This beetle can adjust the number of years the larvae take to complete development, allowing it to adapt to a broad range of local climate conditions. For instance, development can be completed in less than 2 years in warm locations, or it can take as long as a decade in colder climates. Forest Service scientists developed a new climate-driven phenology model that predicts the number of years required for the beetle to complete development and the timing of adult emergence for any location so that management actions may be adapted as needed. This model also has been used in conjunction with host availability to predict where in the U.S. this beetle can develop and how long it will take to complete development. This new information highlights how potentially invasive this insect can be, and the great urgency for finding and eradicating new populations, particularly in warmer areas where the potential for population increase is higher due to more rapid larval development.

External Partners

• Clark University, Worcester, Mass.

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Mapping of the Asian longhorned beetle’s time to maturity and risk to invasion at contiguous United States extent

• Original Paper
• Published: 30 June 2017
• Volume 19 , pages 1999–2013, ( 2017 )

Cite this article

• Alexander P. Kappel   ORCID: orcid.org/0000-0002-8777-0512 1 ,
• R. Talbot Trotter 2 ,
• Melody A. Keena 2 ,
• John Rogan 1 &
• Christopher A. Williams 1  

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Anoplophora glabripennis , the Asian Longhorned Beetle (ALB), is an invasive species of high economic and ecological relevance given the potential it has to cause tree damage, and sometimes mortality, in the United States. Because this pest is introduced by transport in wood-packing products from Asia, ongoing trade activities pose continuous risk of transport and opportunities for introduction. Therefore, a geographic understanding of the spatial distribution of risk factors associated with ALB invasions is needed. Chief among the multiple risk factors are (a) the potential for infestation based on host tree species presence/absence, and (b) the temperature regime as a determinant of ALB’s growth time to maturity. This study uses an empirical model of ALB’s time to maturity as a function of temperature, along with a model of heat transfer in the wood of the host and spatial data describing host species presence/absence data, to produce a map of risk factors across the conterminous United States to define potential for ALB infestation and relative threat of impact. Results show that the region with greatest risk of ALB infestation is the eastern half of the country, with lower risk across most of the western half due to low abundance of host species, less urban area, and prevalence of cold, high elevations. Risk is high in southeastern states primarily because of temperature, while risk is high in northeastern and northern central states because of high abundance of host species.

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Acknowledgements

The authors thank Peter Meng for access to pre-publication host lists. Support for this research was provided by the Graduate School of Geography at Clark University, and the US Forest Service, Northern Research Station. We also thank the editor and 2 anonymous reviewers for comments on previous versions of the manuscript.

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Kappel, A.P., Trotter, R.T., Keena, M.A. et al. Mapping of the Asian longhorned beetle’s time to maturity and risk to invasion at contiguous United States extent. Biol Invasions 19 , 1999–2013 (2017). https://doi.org/10.1007/s10530-017-1398-0

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DOI : https://doi.org/10.1007/s10530-017-1398-0

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Research Article

Description of an establishment event by the invasive Asian longhorned beetle ( Anoplophora glabripennis ) in a suburban landscape in the northeastern United States

Roles Conceptualization, Formal analysis, Investigation, Methodology, Project administration, Writing – original draft, Writing – review & editing

* E-mail: [email protected]

Current address: Environmental Protection Agency, USEPA Headquarters, William Jefferson Clintom Building, 1200 Pennsylvania Avenue, N.W., Mail Code 7505P, Washington, D.C. 20460 United States of America

Affiliation United States Department of Agriculture, Center for Plant Health Science and Technology, Animal and Plant Health Inspection Service, Plant Protection and Quarantine, Otis Lab, United States of America

Roles Methodology, Software

Affiliation United States Department of Agriculture, Animal and Plant Health Inspection Service, Asian Longhorned Beetle Cooperative Eradication Program, 151 West Boylston Dr., Worcester, United States of America

Roles Methodology, Resources

Roles Conceptualization, Data curation, Formal analysis, Funding acquisition, Investigation, Methodology, Resources, Supervision, Writing – review & editing

Affiliation United States Department of Agriculture Forest Service, Northern Research Station, Hamden, United States of America

• Helen Hull-Sanders, 
• Eugene Pepper, 
• Kevin Davis, 
• Robert Talbot Trotter III

• Published: July 20, 2017
• https://doi.org/10.1371/journal.pone.0181655
• Reader Comments

The establishment of non-native species is commonly described as occurring in three phases: arrival, establishment, and dispersal. Both arrival and dispersal by the Asian longhorned beetle ( Anoplophora glabripennis Motschulsky), a xylophagous Cerambycid native to China and the Korean peninsula, has been documented for multiple locations in both North America and Europe, however the transitional phase, establishment, is not well understood for this species due to the need to rapidly remove populations to prevent dispersal and assist eradication, and the evident variation in the behavior of populations. Here we describe the dynamics of an establishment event for the Asian longhorned beetle in a small, isolated population within the regulated quarantine zone near Worcester, Massachusetts, USA. These data were collected during an opportunity afforded by logistical limits on the Cooperative Asian Longhorned Beetle Eradication Program administered by state, federal, and local government partners. Seventy-one infested red maple ( Acer rubrum ) trees and 456 interspersed un-infested trees were surveyed in an isolated, recently established population within a ~0.29 ha stand in a suburban wetland conservation area in which nearly 90% of the trees were host species, and nearly 80% were Acer rubrum . Tree-ring analyses show that within this establishing population, Asian longhorned beetles initially infested one or two A . rubrum , before moving through the stand to infest additional A . rubrum based not on distance or direction, but on tree size, with infestation biased towards trees with larger trunk diameters. Survey data from the larger landscape suggest this population may have generated long-distance dispersers (~1400 m), and that these dispersal events occurred before the originally infested host trees were fully exploited by the beetle. The distribution and intensity of damage documented in this population suggest dispersal here may have been spatially more rapid and diffuse than in other documented infestations. Dispersal at these larger spatial scales also implies that when beetles move beyond the closed canopy of the stand, the direction of dispersal may be linked to prevailing winds.

Citation: Hull-Sanders H, Pepper E, Davis K, Trotter RT III (2017) Description of an establishment event by the invasive Asian longhorned beetle ( Anoplophora glabripennis ) in a suburban landscape in the northeastern United States. PLoS ONE 12(7): e0181655. https://doi.org/10.1371/journal.pone.0181655

Editor: Daniel Doucet, Natural Resources Canada, CANADA

Received: February 3, 2017; Accepted: July 5, 2017; Published: July 20, 2017

This is an open access article, free of all copyright, and may be freely reproduced, distributed, transmitted, modified, built upon, or otherwise used by anyone for any lawful purpose. The work is made available under the Creative Commons CC0 public domain dedication.

Data Availability: All relevant data are within the paper and its Supporting Information files.

Funding: This work supported by the USDA Animal Plant Health Inspection Service and the USDA Forest Service. The funder had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.

Competing interests: The authors have declared that no competing interests exist.

Introduction

The movement of large quantities of commodities and people in the global economy has created opportunities for non-native species to transcend ecological barriers and be introduced to new environments. Many of these introductions do not lead to the establishment of new populations, and even when populations are established, many of these species may remain innocuous [ 1 – 2 ]. However, a minority of non-native species successfully colonize new habitats, and their presence leads to considerable ecological, economic and/or evolutionary impacts in the introduced range [ 2 – 5 ]. The Asian longhorned beetle, Anoplophora glabripennis (Coleoptera: Cerambycidae) is an example of one of these high-impact invaders. A polyphagous wood borer, Asian longhorned beetle larvae feed on a broad range of deciduous trees [ 6 , 7 ], resulting in extensive damage to both the cambium and xylem. Female feeding and oviposition has been documented on 43 species of trees [ 7 – 9 ] in 15 families, with larvae (the life stage responsible for much of the damage of concern) feeding most extensively on the preferred hosts Acer sp., Populus sp., Salix sp., and Ulmus sp. [ 10 – 13 ]. Within the beetle’s native range, which includes portions of China [ 14 ] and the Korean peninsula [ 11 , 12 , 15 ], outbreak populations of Asian longhorned beetles are most common in managed monocultures, windbreaks, and urban trees, which are often made up of non-native host species (frequently Populus sp.) [ 16 ], and the beetle is less common in stands of mixed native species.

Asian longhorned beetles are presumed to have been introduced to North American and European landscapes through international trade utilizing solid wood packing material [ 17 – 19 ] produced from infested wood. In many countries in Europe (including Austria (first reported 2001), Finland (2015), France (2003), Germany (2004), Italy (2007), Montenegro (2015), the Netherlands (2010), Switzerland (2011), and the United Kingdom (2011)), significant non-contiguous beetle infestations have been reported, and the potential for new detections continues. In North America, Asian longhorned beetles have been discovered in five states (Illinois (1998), Massachusetts (2008), New Jersey (2002), New York (1996), and Ohio (2011)) and Ontario, Canada (2003). North American infestations were first associated with cities with significant international trading centers, and their surrounding landscapes [ 20 , 21 ]. However, more recent infestations in Worcester, Massachusetts (2008) and Bethel, Ohio (2011) may be the result of the movement of international commodities beyond their port of entry to their consumer destination [ 20 ].

The process of invasion by non-native species can be classified into three stages: arrival (in which individuals are moved to regions outside of their native range), establishment (the process by which reproducing populations establish and stabilize at levels such that population decline and dissolution is unlikely), and spread (the range expansion of an invading species into novel areas) [ 5 ]. While arrival of Asian longhorned beetles can be inferred by the location of established populations, opportunities to understand the mechanisms driving establishment are more limited due to the high priority placed on the eradication of this species, which results in the rapid removal and destruction of infested trees following detection. This typically leaves little time to carry out intensive surveys and sampling to study infested landscapes in the early stages of establishment. As a result of these limits, past studies on beetle establishment and dispersal have either used mark-recapture methods in the beetles’ native range [ 22 , 23 ], statistical and graphical inference [ 24 , 25 ], or intensive reconstructions of beetle behavior based on damage within individual trees [ 26 , 27 ] in localized populations to identify patterns of beetle movement and dispersal.

Both the mark-recapture studies and the statistical inference models have provided new information regarding the distribution of risk on the landscape following the detection of the beetle. As is the case for many invasive species, prior to being identified as an environmental concern, basic ecological data describing beetle behavior and patterns of population dispersal were unavailable to guide Asian longhorned beetle eradication programs. In an effort to fill these knowledge gaps, Smith et al. [ 22 ] and Smith et al. [ 23 ] conducted large-scale mark-recapture studies on native beetle populations infesting non-native trees in Gansu Province, China. These studies found that 98% of the beetles that were recaptured were found within 920 m of the location where they were released, with a maximum observed dispersal distance of <2600 m. These early studies were critical in providing much needed information on dispersal; however, these studies were limited to assessing beetle behavior in native landscapes, and represented natural within-population dispersal rather than the patterns associated with range expansion. Due to the eradication status of the beetle in most introduced areas, mark-recapture studies are not an option as live beetles cannot be released in areas under eradication.

Statistical and graphical methods are applicable in invaded landscapes, and can often take advantage of data collected by eradication programs [ 24 , 25 ]. However, these studies often require extensive datasets, and the models may provide an assessment of beetle movements at large spatial scales. When isolated pockets of infestation are found, survey and eradication efforts are typically focused at the stand-scale. Filling this knowledge gap requires reconstructions of beetle population development within and between trees.

Two studies have sought to fill this stand-scale knowledge gap [ 26 , 27 ]. These studies have taken advantage dendrochronological methods to date the damage caused by ovipositing females and emerging adults within a stand. The work by Turgeon et al. [ 26 ] provided a reconstruction of beetle population growth and movement in an isolated infestation near Toronto, Canada, and revealed the infestation had been in place for 10 years during which the beetles had spread to infest 50 trees. However, within the infestation, the vast majority (more than 800) of the adult beetles had emerged from a single tree, while 1 tree had more than 30 exit holes, 2 trees had 10–30 exit holes, and 10 trees had fewer than 10 exit holes. This concentration of exit holes on a focal tree suggests the population had been slow to spread, and that many of the adult beetles remained on their natal host. A second reconstruction of beetle establishment and spread was carried out at Paddock Wood in the United Kingdom by Straw et al. [ 27 ]. This study found similar patterns; a 10-year old population which had spread to include 66 infested trees. Like the population in Canada, the beetles had also remained primarily on a focal tree from which the majority of the adult beetles (nearly 500) had emerged. Only one additional tree in the stand had more than 10 exit holes. These studies suggest several common patterns including a relatively slow spread of infestation of new trees and a tendency for the adults to remain on their natal tree.

While these studies have expanded our understanding of the local dynamics of Asian longhorned beetle population growth and movement, both of these studies represent single sample points, and patterns may be highly dependent on local conditions. Here, we seek to expand our understanding of beetle behavior and assess some of the general patterns previously observed by assessing population establishment and movement in an isolated pocket of infested trees identified by the United States Department of Agriculture (USDA) Asian Longhorn Beetle Eradication program in 2013 near Worcester, Massachusetts, USA. The identification of this isolated infestation, and logistical delays in its removal provided a brief window of time in which the site could be surveyed in detail before the removal and destruction of infested material. Within the isolated plot, all host and non-host trees were surveyed and beetle damage was dated using basic tree ring methods [ 28 ] to reconstruct the pattern of establishment within that stand. The identification of a nearby, isolated pocket of infested trees also provided an opportunity to evaluate landscape factors such as wind that may drive larger-scale patterns of beetle dispersal. By generating a spatial analysis of host tree usage, the information on early within-stand beetle movement enables managers to assess the potential for beetle dispersal in a novel landscape, and in turn provides additional biological information on which to base risk assessments and management strategies.

Data sources

Infested trees were identified by surveys carried out by the Massachusetts division of the Asian Longhorned Beetle Eradication Program. This program, which is jointly managed by the USDA Animal and Plant Health Inspection Service (APHIS) and the Massachusetts Department of Conservation and Recreation (Mass DCR), conducts surveys using ground-based teams using binoculars and aerial teams of tree climbers to identify infested trees for removal and disposal. When infested trees are found, personnel with the eradication program assign each tree a unique program identification number and document the tree’s genus and, (when possible) species, as well as the diameter at breast height (dbh), and location of the trees using handheld GPS units.

Site description

The stand of infested trees surveyed for this analysis occupied an overall area of ~ 2,860 m 2 in a wetland conservation area ( Fig 1 ) owned by the Town of Shrewsbury, Massachusetts, U.S.A. (Town of Shrewsbury, Taxplate 29, Block 05100). Work performed under Massachusetts general law chapter 132 section 8 and 11. Inspectors with the Massachusetts Asian Longhorned Beetle Eradication Program found the study stand imbedded within a suburban environment in Shrewsbury, MA, within the Asian longhorned beetle regulated area. The current regulated area includes ~ 285 km 2 of urban, suburban, and wooded landscapes centered on Worcester, MA, an urban environment which grades rapidly into the surrounding forested landscape. The first infested tree to be detected in the Worcester, MA area was found in early in the winter of 2008 and since then, ~25,000 infested trees have been identified and (along with ~11,000 trees considered to be at high risk for infestation) destroyed by the eradication program [ 29 ]. Among the documented infested trees, nearly 97% have been in the genus Acer .

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A) After tree removal April 2014 and B) before tree removal August 2013 with red circles indicating the location of Asian longhorned beetle infested trees.

https://doi.org/10.1371/journal.pone.0181655.g001

Stem-mapping

The stand described above included 526 individual tree stems with inter-tree distances ranging from 0.1 m– 18.5 m. While the canopy of most of the stand was closed and stem density was high, the understory was open and relatively unobstructed, with visibility to ~ 70 meters. The effect of wind was evident in the upper canopy as indicated by the movement of limbs and leaves, however, within the stand, little air movement was noted.

The dense, closed canopy, and close proximity of the stems made the mapping of the stem locations by GPS unfeasible, so locations were mapped using a standard bearing and distance method [ 30 ]. The method consisted of establishing 1 primary control point, and 4 sub-control points. The distance and bearing for each of the sub-points from the control point was determined using a level transit (Topcon AR-F6 Auto Level). The distance and bearing from one of these points was then determined for each stem, using the same equipment. Bearings were taken to the center of the stem to the nearest 1/10 th of a degree, and distances were determined to the nearest 0.1 m. From these data, the locations were converted to Cartesian coordinates relative to the primary control point. These Cartesian coordinates were then converted to UTM coordinates using a single GPS coordinate collected at the primary control point. The primary control point location was determined using a Trimble Geo Explorer 3000 Series operating with Windows Mobile 6 and ArcPad ver. 10 [ 31 ]. The coordinate was determined in UTM Zone-19, and was estimated by spatially averaging 1,000 points (maximum Positional Dilution of Precision allowed was 7). Converted tree coordinates were then used to create a shapefile using ArcMap V10.2 [ 32 ], and overlaid on aerial photographs to produce Fig 1 . Measurements for tree diameters at breast height (dbh) were recorded for all trees; comparisons between distributions of infested and un-infested tree dbh values were made using the Wilcoxon signed rank statistic using R statistical package [ 33 ].

Identification of infested trees

Two distinctive indicators of beetle presence were used to diagnose infestation. Initially, female beetles will chew an elliptical oviposition pit through the bark into which she will typically oviposit a single egg. After development within the tree, adult Asian longhorned beetles will emerge, creating a nearly perfectly round exit hole ~1 cm diameter. Aerial inspectors climbed, surveyed, and verified all infested and non-infested trees within 25 m radius from any tree with an exit hole. Tree sections with oviposition pits and exit holes were isolated after the tree was felled and placed into quarantine for determination of aging characteristics ( see Dating Beetle Damage below).

Beetle infestation levels

The severity of the infestation within individual trees was defined by identifying the type and abundance of beetle damage. Trees which have oviposition pits but no emergence holes were identified with infestation level of A, trees with 1–10 emergence holes and trees with 11–100 emergence holes were identified as B and C level infestations, respectively. Although not explicitly documented, the locations of the observed exit holes and oviposition pits were limited to the main stem and lower canopies of trees. Exit holes and oviposition pits were not found in the upper canopy.

Wind analysis

Daily records for the direction of the maximum 5-s wind for the assumed beetle flight seasons (June through October, inclusive) from 1998 through 2014 (estimated duration of infestation in Worcester, MA regulated area) were obtained from the National Oceanic and Atmospheric Administration (NOAA) National Climatic Data Center (NCDC, available at https://www.ncdc.noaa.gov ), these records provide the directionality of daily wind gusts. Data used for analysis were obtained from the NOAA station at the Worcester Regional Airport (Station No.: GHCND: USW00094746) located to the southwest of the center of the Asian longhorned beetle infestation in Worcester County, MA, and ~14.5 km west-southwest of the study stand. This is the closest meteorological station with complete records within the Asian longhorned beetle regulated area. Daily values for the full record set (records were complete) were tabulated to provide a probability distribution for wind directionality.

Wind source frequency was calculated based on 22.5° bins to generate a “wind” direction bins. Destination Frequency (the direction the beetle would be expected to disperse) was calculated as the inverse of the Wind Source Frequency ( Fig 2 ). The 16 direction bins were then ranked by frequency (with two bins tied for rank 10). Dominant wind direction was plotted in ArcGIS Ver. 10.2 [ 32 ].

Wind relative frequency distnace vectors overlaid onto a map of the infested area, Shrewsbury, MA USA. Red circles indicate location of Asian longhorned beetle infested trees, the callout box indicates the area in Fig 1 .

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Dating beetle damage

Asian longhorned beetle damage was dated using standard dendrochronological techniques as described in Turgeon et al. [ 26 ] and Straw et al. [ 27 ]. Sections of trees with visible exit holes, both healed and unhealed, were removed from all B and C level trees in the field October 2013. Trees were also sampled to identify the oldest oviposition pits (based on weathering and wound response). Sections were placed in a kiln the same day they were removed at the USDA APHIS Plant Protection and Quarantine (PPQ) quarantine facility, Barnstable County, Massachusetts. The kiln was held at 80° C for 15 h to ensure all beetles had been destroyed. The wood was then allowed to continue to dry at room temperature for an additional 72 h. Using a band saw, wood sections were cross-cut through the Asian longhorned beetle exit holes or oviposition pits, and the surfaces were sanded to reveal growth rings. Rings beyond the edge of the emergence holes were counted to determine the year of beetle emergence. Holes with no evidence of healing were dated as the current year. Rings formed over healed or partially healed exit holes were counted to determine the year of emergence.

Establishment distances

A heavily infested tree (42 visible exit holes and 30 oviposition pits) assumed to be the source of the subsequent infestation was confirmed by dendrochronological methods to be the earliest infested tree (2008 oviposition, 2010 exit). The Kolmogorov-Smirnov test for distribution differences was used to determine if beetles moved randomly throughout the stand using R statistical package [ 33 ]. Expected dispersal distances were calculated using the source tree and finding the distance to all other trees in the stand, both infested and un-infested. Observed dispersal distances were calculated by determining distances between the source tree and the infested trees only.

Adjacency rules

The movement of female beetles from the natal tree to a newly infested tree is used here to define connections, or adjacencies among trees. The movement of beetles is inferred based on two rules established by Trotter and Hull-Sanders [ 24 ], those rules are: 1) host trees receive beetles from the nearest tree with a higher level of infestation (which is assumed to be older) and 2) trees can serve as the source of beetles for multiple trees, but each tree is assumed to have received beetles once.

The lengths and directions of these inferred dispersal events (adjacency vectors), and the dispersal distance percentiles were calculated using a custom built MatLab R2013b [ 34 ] script (TreeAdjacency) as described in Trotter and Hull-Sanders [ 24 ]. Distance vectors were tabulated to produce models relating vector distance to frequency.

The infested stand in Shrewsbury, MA included 526 trees within a ~ 2,860 m 2 area ( Fig 1 ), and included trees in the genera Acer , Betula , Carya , Cornus , Fagus , Fraxinus , Ilex , Malus , Pinus , Populus , Quercus , Salix , Ulmus , and Viburnum . Host genera (as listed by Meng et al. 2015), including Acer , Betula , Fagus , Fraxinus , Malus , Populus , Salix , and Ulmus accounted for 94% of the stems. Red maple ( A . rubrum) alone accounted for 81% of the stems while Malus spp. accounted for 8%. Acer platanoides , Populus tremuloides , and Salix spp. were represented by a single stem each. The earliest exit hole dated to 2010 and corresponded with the earliest oviposition pit dated to 2008, suggesting a ~2 year period for development. A total of 71 trees had signs of infestation (oviposition pits and/or exit holes) at the Shrewsbury location, all of these were A . rubrum . In total, the stand included 2 trees with 42 and 40 exit holes, 4 trees with 2–4 exit holes, and 65 trees with oviposition pits only. Within the Asian Longhorned Beetle Eradication Program, these are referred to as C, B, and A level infestations, respectively. The remaining host trees did not show indications of infestation.

Infested A . rubrum host trees were, as a group, larger (median = 19.5 cm dbh, IQR = 15.2 cm dbh) than un-infested A . rubrum (median = 6.0 cm dbh, IQR = 8 cm dbh) (Wilcoxon rank sum W = 23348, P < 0.0001) ( Fig 3 ). The distribution of distances that beetles were inferred to have moved differed significantly from the distances that might be expected if movements were to occur at random (Kolmogorov-Smirnov D = 0.6847, P < 0.0001) ( Fig 4 ), with beetles moving longer distances than might be expected if beetles were to disperse randomly through the stand (median inferred movement distance 5.74 m, IQR = 5.84 m, median possible inter-tree movement distance 18.89 m, IQR = 13.13 m). Movement among trees by adult females was inferred by using assumptions described in Trotter and Hull-Sanders [ 24 ], in which beetles on A trees are assumed to have originated from the nearest B or C tree, and beetles on B trees are assumed to have originated on the nearest C tree. These assumptions of tree adjacency (connectivity) are shown graphically on the Fig 5B by the lines, which denote the movement of beetles to B and A trees.

Infested (A) and un-infested (B) trees based on the diameter at breast height (dbh) in centimeters for 527 trees in Shrewsbury, MA USA.

https://doi.org/10.1371/journal.pone.0181655.g003

Inferred (A) and Potential (B) below canopy dispersal movements away from source trees (>10 exit holes) to surrounding host trees.

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Within the stand, open circles represent un-infested trees and filled circles represent infested trees. (A) Large square represents the assumed source for the infestation (initially infested tree based on exit hole data). (B) Lines connecting trees indicate inferred patterns of Asian longhorned beetle movement through the stand.

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Wind patterns on the landscape during the emergence months were generally SSW to NNE ( Table 1 ; Fig 2 ); this probability distribution provides an expected distribution of dispersal directions from a given point source ( Table 1 ). When the resultant polygon was applied at the county level to the initially infested tree within the study stand, more distal infested trees identified by the eradication program generally fell within four of the most common wind direction bins, between absolute bearings 45° and 112.5° ( Fig 2 ).

https://doi.org/10.1371/journal.pone.0181655.t001

Frequencies were determined by counting the number of fastest 5- second wind intervals. Sixteen directional bins ranked by wind frequency ranked the lowest (1) to highest (16) wind destination likelihood.

At smaller spatial scales, if the establishment of new pockets of infestation are the result of stratified dispersal from the source, then it would be expected that these pockets of infestation would be younger than the source population, as indicated by the number of infested trees and exit holes. Similarly, if adult dispersal is influenced by wind direction then new infestations may be expected to be associated with wind patterns. An assessment of the survey records for the Asian Longhorned Beetle Eradication Program shows that two trees which were ~1400 m from the surveyed population were found to be infested, with both trees having only had small numbers of oviposition pits. Trees closer to the infestation, but outside the surveyed stand had more severe damage, including a greater number of oviposition pits, feeding galleries, sapwood entrance holes, and a few exit holes. All infested trees were located within the top 4 wind destination categories, including the two outliers shown in Fig 2 , suggesting outside of the stand, wind direction may play a role in structuring dispersal.

Past research has shown the dispersal behavior of Cerambycid beetles to be variable, often dispersing when host quality goes down, adult population densities increase, or inter-specific competition with other large cerambycids occurs [ 35 – 39 ]. Recent studies by Turgeon et al. [ 26 ] and Straw et al. [ 27 ] have provided reconstructions of the spread of the Asian longhorned beetle within infested stands, and these data have also suggested beetles may remain on the natal tree, leading to high beetle density in focal trees within an isolated infestation. However, the data from the Worcester, MA stand seems to indicate that this may not be a rule in Asian longhorned beetle population dynamics. Here, the beetles seemed to move rapidly to infest numerous trees within the stand, and perhaps some trees well beyond the stand, long before populations in the originally infested tree grew to the sizes observed in Canada or the United Kingdom. Weather, stand structure, tree density, and the nature of the landscape may play a role in structuring these patterns. Although each of these three infestations was found in urban environments, the structures of the landscapes varied. The infestation in Canada was found in an area described as “light industrial” [ 26 ], while the infestation in the United Kingdom was found in an area with “light industry” as well as agricultural lands and woodlots [ 27 ]. The localized infestation described in this study was found in a heavily wooded residential area, which lacked industry. In all of these locations, there exists the potential for beetles to disperse under their own power; to be moved by transport of infested materials or by hitch-hiking on vehicles; however, limited replication (3 locations) and lack of direct observations mean that the role these variations in the landscape play in the active and passive dispersal of the beetle remain unknown, but clarifying this pattern may be important in providing guidance to management and eradication programs, as the propensity of the beetle to disperse is a key issue in determining the needed extent and intensity for effective surveys.

The study by Trotter and Hull-Sanders [ 24 ] highlights the potential importance of this information. At the time the study was conducted, limited information on the propensity of adult female Asian longhorned beetles to dispersal from natal trees in the introduced range was available. To accommodate this knowledge gap, the potential patterns of beetle distribution were inferred based on two sets of assumptions. Under the strict assumptions of dispersal, it was assumed that the beetle would be reluctant to disperse from trees with low levels of infestation, a pattern consistent with that observed by both Turgeon et al. [ 26 ] and Straw et al. [ 27 ]. This set of assumptions, by the nature of the distribution of the infested trees, means that while fewer beetles disperse, the dispersal events that do occur are longer. Under the alternative, relaxed dispersal scenario, it is assumed that beetles may disperse from trees with low levels of infestation. This pattern results in more trees acting as sources for dispersing beetles, but implies that when beetles do disperse, they typically do so over shorter distances. This relaxed pattern of dispersal assumptions is more in alignment with the patterns observed within this study. Ultimately, these patterns may alter the spatial distribution of beetle risk on the landscape around infested areas.

Other studies conducted in the beetles’ native range may shed some light on this issue. In the introduced range, under strict dispersal rules (i.e. beetles do not disperse from the tree from which they emerged until the tree has been heavily infested) dispersal distances were estimated to be greater than 2.3 km based on infested tree data [ 24 ]. In trees within an agricultural wind-break in China, Smith et al. [ 22 ] observed 98% of marked Asian longhorned beetles dispersed within 920 m of their release point; however, a minority could be found as far as 2.6 km [ 23 ] and the direction of dispersal was associated with wind direction [ 40 ]. The outlying population of recently infested trees (oviposition pits only, no exit holes), which are aligned with the direction of prevailing daytime winds supports both the direction of movement, and the indication of long-distance dispersal events. The association with wind direction suggests long-range dispersers may have left the stand, perhaps by flying above the canopy where they would have been subjected to stronger winds. Although it is possible that the ENE infestation of two trees >1400 m from the surveyed area was due to human mediated movement, the two trees lie within the dominant wind vectors and are within dispersal distances inferred by Trotter and Hull-Sanders [ 24 ], suggesting spread may have been driven by natural dispersal.

The limited distribution of infested trees observed in both Canada and the UK do not necessarily suggest the absence of long distance dispersal. In fact, Turgeon et al. [ 26 ] discuss the likelihood that the isolated infestation studied was the result of long-distance dispersal by beetles from the center of a larger infestation in Toronto, Canada, some 10 km away. This process, by which larger infestations produce satellite infestations is also likely to have produced the pocket of Asian longhorned beetle described in this study. What is not known however, is why these two satellite infestations, with similar climatic conditions, seem to have resulted in different patterns of beetle movement.

While most non-indigenous species fail to establish after arrival, problematic alien species become invasive after both establishing and dispersing following introduction [ 5 ]. Short-distance dispersal may be part of the establishment phase. The establishment phase serves as the decisive interval during which expanding populations stabilize and increase their distribution such that population decline and dissolution is unlikely [ 5 ]. Unfortunately, management and subsequent eradication programs often only detect the problem during the spread phase and have little opportunity to determine, control, or understand the establishment phase. Our survey of all standing trees within the conservation area before the Asian longhorned beetle population was expected to substantially increase (within the first five years after arrival [ 41 ]) allowed us to characterize the establishment phase.

Three major patterns of Asian longhorned beetle movement following establishment within a stand have emerged. The first, is that the pattern of strict beetle movement, in which dispersal from the natal tree is unlikely until the tree has been heavily utilized has been supported by some studies, however, the current study suggests this pattern may not be consistent among all locations. The second issue of interest is that, once a beetle does disperse from its natal tree, it does not appear to randomly select a new host, but may be selective based on characteristics such as size and species, with an emphasis on A . rubrum . This finding agrees with previous studies such as Dodds et al. [ 42 ], which found Asian longhorned beetles in mixed stands with A . rubrum , Acer saccharum , and Acer platanoides , tended to infest A . rubrum trees with diameters similar to infested trees found at our site (mean dbh Delaval 17.0 ± 1.2 cm, Boylston 19.2 ± 2.3 cm, this study 21.8 ± 1.3 cm). Asian longhorned beetles may bypass smaller host trees that may be located closer in proximity to the natal tree, continuing to move through the understory until “appropriate” hosts are found. If this behavior is driven by the suitability of the host for the development of the larvae, this pattern would suggest a potential preference-performance link [ 43 ], though more research on this is needed. While female Asian longhorned beetles will create significantly more oviposition pits and deposit more viable eggs on A . saccharum in the lab (Smith et al. 2002) [ 44 ] and in the field [ 45 ] than on A . rubrum ; in the field, similar sized A . rubrum trees had a greater number of exit holes [ 45 ]. Dodds and Orwig also [ 45 ] found that within two infested stands in Worcester, MA, only Acer tree species were infested. All of the trees at one site and more than half of the trees at a second site >30 cm dbh were infested. However, these infestations were larger and presumably older than the infestation described in this paper. The potential interaction between host size and host species in determining host quality may help structure the dynamics of invasions where the beetle and hosts may lack an evolutionary history such as in the invaded stands in Canada, the United States, and throughout Europe, and merits further study.

These results may have application to the management of Asian longhorned beetles, and provide additional information on how beetle populations may behave when establishing in novel environments. However, these data suggest inconsistencies among invaded stands, and highlight the potential for as-yet undocumented factors to play a large role in how beetle move through the landscape. This information in turn is likely to be of value for basic biogeographic theory, as it may help explain variations in dispersal patterns, and some of the factors (and evolutionary history, such as host associations) which may drive that variation. This information is also likely to be critical for management and eradication efforts, where the effective use of surveys depends on a sound understanding of the probabilities associated with the directions, distances, and frequencies of propagule movement, and survey efforts may find new efficiencies from increased focus on tree size and local wind patterns in identifying the boundaries of infestations, though additional work at multiple spatial scales is needed to parameterize survey priorities.

Acknowledgments

The authors would like to thank the USDA PPQ–Otis Laboratory, Buzzards Bay, Massachusetts and USDA PPQ Asian Longhorned Beetle Eradication Program, Worcester, Massachusetts for all their cooperation and support on this project. We would also like to acknowledge the contributions of Christopher Perry, J. Chris Fagan, and David Mikus for their invaluable technical help in the field and processing of wood samples. Any reference to commercial software is made for the convenience of the reader, and does not constitute an endorsement of any software or company.

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• 41. Sawyer A. Expected dispersal of Asian longhorned beetles from preferred host trees as a function of infestation level and date of removal during the flight season. United States Department of Agriculture. 2009. Report from the USDA APHIS PPQ Otis Laboratory to the ALB Technical Working Group.

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Successful Eradication of the Asian Longhorn Beetle, Anoplophora glabripennis , from North-Eastern Italy: Protocol, Techniques and Results

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Simple Summary

Anoplophora glabripennis (Coleoptera: Cerambycidae) is an extremely polyphagus Asian wood-boring beetle accidentally introduced into North America and Europe. In 2009, an infestation was found in the municipality of Cornuda (Veneto Region, Italy). In order to eradicate the pest, several actions were immediately undertaken in the delimitated infested and buffer zones: tree visual inspections twice a year, felling and chipping of infested and suspected trees, trapping protocols, mitigation plans based on substitution of felled trees with new plants, and citizen alerts. The program lasted 11 years, after which the species was declared eradicated from the region. During the eradication program more than 36,000 trees were surveyed and more than 2000 trees were felled. Trees most affected by the pest were birches, elms, maples, and willows. This paper describes all the actions undertaken during the eradication program, providing a protocol that can also be used for future eradications of the species.

The Asian Longhorn Beetle (ALB), Anoplophora glabripennis (Coleoptera: Cerambycidae), is an important and extremely polyphagous wood-boring beetle native to Asia. In the 1990s, ALB was accidentally introduced into North America and Europe. In 2009, a large ALB infestation was found in the Veneto Region (north-eastern Italy), in the municipality of Cornuda (Treviso province). Eradication actions were immediately undertaken, based on delimitation of infested and buffer zones, tree visual inspections, felling and chipping of infested trees, trapping protocols, and citizen alerts. A total of 36,361 trees, belonging to 16 genera, were surveyed twice a year over an area of 7594 hectares. In 2020, after 11 years of eradication measures, the ALB population of Cornuda was declared eradicated. Overall, 2361 trees belonging to 8 genera were felled and destroyed, of which 1157 were found to be infested by ALB. This paper describes all the actions carried out and the procedures applied in order to eradicate ALB from north-eastern Italy, providing a useful example for current and future ALB eradication programs.

1. Introduction

The Asian Longhorn Beetle (ALB), Anoplophora glabripennis (Motschulsky) (Coleoptera Cerambycidae), is a wood-boring beetle native to China and the Korean Peninsula [ 1 ]. Although in its native range A. glabripennis mainly infests Populus spp., Salix spp., Ulmus spp., and Acer spp. [ 2 ], A. glabripennis is an extremely polyphagous pest able to develop on woody broadleaves of 34 tree species belonging to 14 genera in 10 families [ 1 , 3 ]. A. glabripennis attacks both young and mature trees growing in urban and peri-urban areas [ 3 , 4 , 5 ]. Unlike most longhorn beetles, A. glabripennis develops in apparently healthy plants, although it can also infest stressed trees and fresh logs [ 6 , 7 ]. For these reasons, the introduction of A. glabripennis in new areas represents an enormous threat to urban parks and rural forests [ 8 , 9 ].

Accidentally introduced with infested wood packaging material associated with international trade [ 3 ], A. glabripennis was first found outside its native range in New York City (NY, USA) in 1996 [ 10 , 11 ]. Following this, the pest was found also in other states of the USA (Illinois, New Jersey, Massachusetts, Ohio, and South Carolina) [ 4 , 12 , 13 , 14 , 15 ], in Canada (Ontario) [ 16 ], and in Europe, where the first presence of A. glabripennis was recorded in Austria (2001) [ 17 ], followed by France (continental) (2003) [ 18 ], Germany (2004) [ 19 ], Italy (2007) [ 20 ], the Netherlands (2010) [ 21 ], Switzerland (2011) [ 22 ], the UK (2012) [ 23 ], Corse (2013) [ 24 ], Finland (2015) [ 25 ], and Montenegro (2015) [ 26 ]. After eradication measures undertaken by different states, to date, infestations of the pest still occur in France (both continental and Corse), Germany, Italy, and the USA [ 27 ].

In Italy A. glabripennis was found in the northern regions of Lombardy (2007) [ 20 ], Veneto (2009) [ 28 ], Piedmont (2018) [ 29 ], and in the central region of Marche (2013) [ 30 ]. At these latitudes the whole A. glabripennis life cycle lasts 12–18 months (Faccoli, pers. observ.). Adults emerge in summer, mainly in late June–early July [ 31 ]. After maturation feedings and mating, the female lays eggs in oviposition pits chewed out under the bark of the host tree [ 31 ]. Young larvae initially feed on phloem and, starting from the third instar, they complete their development in the wood [ 32 ]. Pupation lasts a couple of weeks, in a pupal chamber created by mature larvae in the sapwood, and the new adult emerges through a circular hole (about 10 mm diameter, smaller in males) [ 32 ]. In Europe, complete development of A. glabripennis has been recorded on many genera of woody broadleaves.

The A. glabripennis infestation occurring in north-eastern Italy (Veneto Region) was discovered in June 2009 in the municipality of Cornuda [ 28 , 33 ]. A maple was found to be infested in a private garden of the city center following a report of the garden owner to the local phytosanitary office of the Regional Plant Protection Organization (RPPO). The infested tree showed all the typical symptoms of A. glabripennis infestation, with large and circular emergence holes, sectorial dieback of the canopy with dead branches, and oviposition pits along the stem. Immediately after the discovery of the pest, a large-scale intensive monitoring and eradication program started. The eradication program was based on the establishment of buffer zones, visual inspections, felling and destruction of infested trees, trapping protocols, and citizen alerts, following the official guidelines issued by EPPO [ 34 ]. The A. glabripennis population of Cornuda was declared eradicated in 2020 [ 35 ], after 11 years of application of specific control measures. The aim of this paper is to present and describe all the actions and procedures successfully carried out in order to eradicate A. glabripennis from north-eastern Italy.

2. Materials and Methods

2.1. site description.

The municipality of Cornuda (45°830′ N, 12°010′ E; Italy), is located along the southern border of the Italian Alps, approximately 160 m a.s.l., in a transition from continental to Mediterranean climatic conditions, with temperate summers and winters. Annual precipitation ranges from 1100 to 1200 mm, concentrated in the spring and autumn. Cornuda is a small village of about 6200 inhabitants located in a suburban area fallowing within a hilly landscape with patches of agricultural land. The village structure consists mainly of small, isolated houses surrounded by small private gardens often containing ornamental trees represented by woody native or exotic broadleaves. Public tree-lined parks and rows of trees along the streets are also common.

The village is closely surrounded by mixed deciduous forests and riparian habitats which develop along the Piave river. Natural forests are primarily composed of Acer pseudoplatanus L., Carpinus betulus L., Fagus sylvatica L., Fraxinus excelsior L. and Quercus robur L. on shady damp slopes, as well as Betula pendula Roth., Fraxinus ornus L., Ostrya carpinifolia Scopoli and Quercus pubescens Willd. on sunny dry slopes.

2.2. Tree Database

Since the first record of the A. glabripennis in June 2009, an extensive monitoring program started, visually checking one-by-one all of the possible host trees occurring in Cornuda and in the surrounding municipalities (see next paragraphs), with search directions radially oriented from the point of first discovery of the pest. All host trees were also geo-referenced, creating a specific database reporting information about the plant (genera, age, general physiological conditions), geographic position (coordinates), and ownership data (owner name, address and phone number). Some import–export companies work in in the Cornuda territory, with frequent commercial trade with China. As the infestation may have originated from these companies, particular attention was paid to surveying their surrounding areas.

2.3. Visual Tree Survey

After the first survey carried out in summer 2009, with the creation of the tree database, a systemic and continuative survey of the host-plants started, choosing the most susceptible tree genera according to the literature: Acer , Aesculus , Alnus , Betula , Carpinus , Cercidiphyllum , Corylus , Fagus , Fraxinus , Ostrya , Platanus , Populus , Prunus , Salix , Tilia and Ulmus [ 3 , 4 ]. The aim of the survey was to discover the presence of A. glabripennis -infested plants and update the infested and buffer areas accordingly. The survey was conducted by 3 teams of operators each composed by 2 trained technicians of the Regional Plant Protection Organization that checked all the host-trees one-by-one from the ground, with the use of binoculars looking for A. glabripennis infestation symptoms. All host-trees were geo-referenced and checked twice a year, in summer and late-autumn, looking for adult exit holes, larval frass emission, oviposition scars, maturation feedings on twigs, tree decline with branch dieback, and presence of adults on branches and canopies ( Figure 1 ). On one hand, some symptoms are recognizable more easily in summer, such as the presence of recently emerged adults, fresh oviposition pits on the bark, maturation feedings performed by immature adults on twigs, and occurrence of dying branches. On the other hand, the presence of emerging holes is more visible in late autumn after the plants have lost leaves and the upper branches and canopy are easily checkable from the ground. In case of large trees or plants having trunk and branches covered with ivy, difficult to check from ground with accuracy, the inspectors of the Phytosanitary Service were supported both in summer and winter by a team of 4 tree-climbers from the Treviso Forest Service, who also checked for the potential presence of symptoms in the upper part of the canopy.

Main symptoms of A. glabripennis infestation: presence of adults ( a ), adult exit holes ( b ), oviposition scars ( c ), maturation feedings on twigs ( d ).

Forested areas neighboring the village of Cornuda and falling within the infested or buffer areas were surveyed once per year, along the main forest edges (i.e., external edges, forest tracks, and clear-cut edges). A strip at least 30 m deep inside the forest was monitored. Studies conducted in native regions of South Korea suggest that A. glabripennis is not a true forest species but is adapted to riparian habitats characterized by long edges [ 36 ]. Similarly, in countries where they are introduced, A. glabripennis infestations are usually limited to urban trees that are isolated, growing in small groups or rows, in small rural stands or along forest edges [ 3 , 4 ].

2.4. Zone Delimitation

After the checking of all host plants growing in the area, the infested and buffer zones were established ( Figure 2 ). The “infested zone” consisted of a polygon including all the infested plants, where the vertices of the polygon were the more external infected plants. The infested zone was included in the territory of six municipalities (Cornuda, Pederobba, Crocetta del Montello, Maser, Montebelluna and Caerano di San Marco). Then, a “buffer zone” was established with a radius of 2 km around the infested zone, i.e., around the most external infested trees, according to European regulations [ 34 ]. Year-by-year, when new infested trees were found, the delimited zones (infested and buffer zones) were officially extended (or restricted) several times, coming to also include the bordering municipalities where satellite infestations have been discovered since 2010.

Map of the A. glabripennis infestation, with position of infested trees, involved municipalities border and first and maximum buffer zones extension.

2.5. Pheromone Traps

During the eradication program, trapping protocols were carried out in the buffer zone in order to verify the presence of A. glabripennis in the territory and to test the effectiveness of different trapping tools (i.e., trap models and lures). The sites for trap-setting were chosen in relation to the concentration of susceptible hosts and in suitable areas near the edge of the delimited zone.

In 2011, 14 traps of different type, size, and color were placed for one month (August) and checked weekly ( Table 1 ). Traps were baited with a blend produced by ChemTica International (Heredia, Costa Rica). In 2012, 27 black cross-vane traps were set up and baited with either the ChemTica blend used in 2011, or a Russian experimental blend provided in six different formulations by Dr. Oleg Kulinich of the Department of Forest Quarantine, of the All-Russian Center of Plant Quarantine of Moscow. Traps were checked three times during July, the month with the highest emergence of A. glabripennis adults [ 31 ]. In 2013, 24 traps were placed: 6 control unbaited traps, and 6 others for each of three different blends, equally divided (ChemTica blend and two new formulations of the Russian blend). Lastly, in 2019, ten black cross-vane traps were set in the delimited zone and checked every 2 weeks from middle June to middle September, in order to support the action of visual inspections and verify if A. glabripennis was successfully eradicated. Traps were baited with the ChemTica blend tested in the previous year. The used trap types and lure formulations are summarized in Table 1 .

Information about detection tools used during the eradication program. For each year information was reported about the trap model (type and color), the trap number, and the type of lure.

2.6. Sanitation Felling and Tree Destruction

All trees detected during the visual survey as showing A. glabripennis infestation symptoms were cut and destroyed (details below). Trees reporting unclear symptoms possibly confused with those potentially caused by others urban pests infesting the same host-trees (e.g., Cossus cossus (Lepidoptera: Cossidae), Zeuzera pyrina (Lepidoptera: Cossidae), Saperda carcharias (Coleoptera: Cerambycidae)) were cut and destroyed as well.

A. glabripennis adults leave the tree where they emerged only rarely—when they are strongly disturbed, for example, by tree felling. Moreover, adults were found to be active from the end of May to November. Therefore, infested trees found during the spring–summer survey were marked but not cut immediately, to avoid adult dispersal during tree felling and movement of infested wood through the village. Tree felling, carried out by workers of the Regional Forest Service, was hence postponed every year to winter, between December and April, during insect hibernation as mature larvae [ 31 ]. Infested trees, from both public and private properties, were felled at ground level, leaving only stumps, and moved to a fenced and paved storage area falling within the infestation zone, but away from host plants. In winter, cut trees were then chipped in 2-cm long chips, and chips sold to a biomass power station outside the infestation area. Wood chips have been submitted to entomological analysis by the University of Padua in order to exclude the presence of live larvae. However, chips were moved to the biomass power station and burned by the end of winter to reduce the risk of dispersal outside the infestation area of woody material potentially infested with active A. glabripennis stages.

The first 4 years of eradication (2009–2012), tree cutting, and chipping only concerned infested plants and a few infested trees growing close to the attacked ones. Then, in order to make the eradication protocol more effective, the Regional Decree no. 33 of 10 September 2012 of the Veneto Region introduced the “Clear-cut” measure, which involved identification and cutting of all susceptible plants, even if apparently uninfested, occurring in the area within a 50 m radius from each infested plant. Since 2015, considering the imminent approval of the Decision UE/2015/893 of the European Commission, the clear-cut radius has been increased to 100 m.

2.7. Mitigation Plan

In accordance with the owner’s will, plants felled in private gardens were replaced for free with new trees belonging to non-host species. Young trees (3–4 years old) were chosen by the citizens among the species available for reforestation and urban design programs at the forest nursery of the Regional Forest Service, including Cercis siliquastrum , Clerodendrom trychotomum , Ginko biloba , Liquidambar styraciflua , Quercus robur , and Quercus pubescens .

2.8. Communication

Municipalities and other territorial authorities, such as schools, park administrators and citizens’ associations, were immediately involved. Public meetings were organized to inform citizens, with the distribution of informative brochures and poster hanging, and releases were sent to local newspapers. Moreover, specific technical meetings were organized to inform and train local nurserymen, pruners, gardeners, and other professional stakeholders. Lastly, brief lessons were organized for students of primary and secondary schools of the territory, with projection of slides and photos concerning pest biology and symptoms. In this way, citizens were informed about the threat and how to recognize and report signs of the pest presence. A toll-free number was also activated to allow citizens to report suspect symptoms or A. glabripennis sightings. Finally, an internet site was activated providing information for citizens and a platform to upload reports.

The eradication plan started in the summer of 2009 and ended in 2020 when, according to Commission regulation EU/2015/893, the species was declared eradicated from Cornuda and its surrounding municipalities [ 35 ], following four years of no new records of infested plants. Indeed, neither insects nor plants showing infestation symptoms (oviposition pits, emerging holes, or maturation feedings) have been found since 2016.

3.1. Visual Tree Survey

Since the beginning of monitoring in 2009, more than 36,000 trees were surveyed singly over twelve years ( Table 2 ). Among the 16 surveyed genera, the most abundant were Acer (10,277 trees, 28%), Ulmus (6227 trees, 17%), Salix (5271 trees, 15%), Carpinus (4837 trees, 13%), and Betula (2067 trees, 6%); other genera occurred in percentages lower than 5% ( Table 2 ).

Number of monitored and infested trees, with relative percentages of infestation, divided by genera.

During the survey carried out from 2009 to 2020, 1157 trees belonging to 8 genera were found to be infested (3% of the total amount of surveyed trees). The most at-tacked genera were Acer (431 trees infested), Ulmus (337 trees infested), Betula (210 trees infested), and Salix (149 trees infested)—even if, looking at the ratio between in-fested and monitored trees, the most susceptible genera were Cercidiphyllum (18.2%), Aesculus (11.6%) and Betula (10.2%), followed by Ulmus (5.4%), Acer (4.2%), and Salix (2.8%) ( Table 2 ). Monitored but never found infested genera were Alnus, Carpinus, Cor-ylus, Fagus, Fraxinus, Ostrya, Platanus and Tilia ( Table 2 ).

3.2. Zone Delimitation

In 2009, the year of the infestation discovery, the delimited zone had an area of 4105 ha which grew in the subsequent years, reaching the maximum size of 7594 ha in 2013 ( Figure 2 ). In 2016, as a result of the application of the eradication protocol, both infestation and buffer zone began to decline, reaching the minimum value of 1843 ha in 2018 ( Table 3 ).

Extension of buffer zones, number of monitored and felled trees (divided in “infested” and susceptible trees), and clear-cut radius adopted for each year.

* Healthy trees were monitored every year, so the total is not the sum of monitored trees for each year, but the total amount of monitored trees during the eradication period. Susceptible trees: healthy host-trees growing close to an infested tree. During the first three years, no clear-cuts were applied systematically; however, in the case of polychromic trees, or trees very close to infested ones, or trees damaged by felling-infested trees, felling was also carried out on plants that were not directly infested.

3.3. Pheromone Traps

During the three year monitoring with traps (2011–2013), only two A. glabripennis females were caught in 2013. The two individuals were caught by one multi-funnel and one cross-vane trap, both baited with the ChemTica blend. In 2019, after three years without finding any attacked plants, traps were placed to provide further confirmation that eradication had taken place, and no individuals were caught.

3.4. Sanitation Felling, Tree Destruction, and Mitigation Plan

Beside the 1157 infested trees, another 1204 trees were felled because they were inside the “clear-cut area” ( Table 3 ). Overall, a total of 2361 trees were cut in 12 years during the eradication plan applied in Cornuda, of which only 220 plants were from private gardens. The highest peak of felling occurred during the first year (2009), with 630 cut trees. Subsequently, the number was steadily decreasing with time, until no new plants were felled since 2016.

Two samples of wood chips, taken in 2010 and in 2011, have been submitted to an entomological analysis by the University of Padua. The analysis showed that the size of the wood pieces from the chipping operations is incompatible with the development and survival of A. glabripennis larvae in the wood; almost all of the material analysed was less than 2 cm in length, compared to 4–5 cm in length for mature larvae. In fact, several remains of crushed larvae were found during the analysis. Moreover, the few larger elements are subject to rapid deterioration due to tissue dehydration or fermentation, depending on the humidity conditions. In conclusion, the product tested was found to be biologically safe and free from risk of spreading A. glabripennis infestation.

Main costs incurred in carrying out eradication program were:

• €380,000 used by the Regional Plant Protection Organization for the annual tree survey (twice per year).
• €520,000 used by Regional Forest Services for felling and chipping trees.
• €20,000 used by the University of Padua for scientific support and research activities.

The program was initially financed by funds from the Veneto Region, which then accessed European funds for the management and eradication of invasive species.

As compensation for the felling of infested trees occurring in private properties, owners could choose a new tree to plant as a replacement. A total of 217 new trees (over 220 cut) were planted, including Cercis siliquastrum (65), Liquidambar styraciflua (41), Ginko biloba (35), Clerodendrom trychotomum (29), Quercus robur (27), and Quercus pubescens (20). There was no financial compensation.

4. Discussion

Eradication is the numerical reduction of a population in a specific geographic area to prevent its reproduction and, therefore, bring it to local extinction [ 37 , 38 ]. Conditions that support a higher probability of successful eradication include early detection of the pest (i.e., limited spatial distribution), ability to detect and identify the invader or the infested plants, availability of effective tools for pest monitoring and control, and public support [ 39 , 40 ]. Moreover, the target species should have all or most of the following characteristics: low rate of reproduction and dispersal, ease of detection at low population density, and limited host range [ 39 ].

Early detection of the pest plays a key role in a successful eradication program. The earlier the parasite is discovered from the time of actual arrival, the higher the chances of success, as the parasite will have less time to reproduce and spread. In fact, the probability of successful eradication declines with increases in the infested area [ 41 , 42 ]. In particular, Rejmánek and Pitcairn [ 42 ], analyzing data from 53 infestations of 18 pest species, showed that eradication success probability is about 50% between 0.1 and 1.0 hectares and about 25% between 100 and 1000 hectares of infested area. Moreover, as the area of eradication increases, the required efforts (i.e., costs) also increase and the operation may no longer be economically viable [ 40 ]. The Cornuda infestation initially measured about 4000 ha (infested and buffer zones) and expanded to a maximum extension of about 7600 ha. Although the infestation was discovered in 2009, it was verified, by dating the exit holes from the host trees, that the infestation started at least five years previously, in 2005 [ 31 ]. This delay in starting the eradication program caused an effort of eight years of active eradication (2009–2016) and another four years (2017–2020) of surveying in order to eradicate A. glabripennis from the territory. The monitoring protocol involved more than 36,000 trees checked one-by-one twice a year for 11 years; 2361 of these trees were felled because they were found to be infested or simply because they were within the clear-cut radius. Another example of successful A. glabripennis eradication is at Paddock Wood (Kent, UK), although in this case the infestation was much smaller (with an infested zone of only 11.4 ha). After just one year (and another seven years of surveying) the pest was eradicated and about 2200 trees were felled, of which 66 were infested [ 43 , 44 ]. In contrast, a large infestation was detected in Worcester (MA, USA) in 2008 [ 5 ], and is still active [ 45 ]. Until 2015, the extension of the infestation was larger than 20,000 ha with more than 5 million monitored trees, of which approximately 34,000 were removed (both infested, and those deemed to be high-risk) [ 46 ]. Such a wide spread makes a successful eradication challenging [ 5 ].

Besides early detection, a successful eradication is based on the possibility of easily identifying the pest or its infestation symptoms and the disposal of effective tools for its detection. Visual inspections have proved to be effective against A. glabripennis , but they lose effectiveness for recently infested trees [ 43 ], or in the case of large trees or trunks covered by ivy (Faccoli, pers. observ.). Pheromone traps are often used alongside the work of phytosanitary inspectors, both to find pests and for their eradication by mass-trapping and lure-and-kill techniques [ 47 , 48 , 49 , 50 ]. Unfortunately, no long-range pheromone has been reported for A. glabripennis , although both male-produced short-range and female-produced contact recognition pheromones have been identified [ 3 , 4 , 51 , 52 ]. Several studies have tested the effectiveness of these pheromones combined with some host-volatiles (e.g., Z-3-hexen-1-ol and Linalool) in trapping protocols, showing some positive outcomes, but with few catches despite the dozens of traps used [ 53 , 54 , 55 ]. During the eradication program carried out in Cornuda, only two A. glabripennis females were caught by traps in 2013 and no A. glabripennis individuals were caught by traps used at Paddock Wood [ 43 ]. Despite the use of A. glabripennis pheromones remaining indicated for pest interception in areas where it is not yet been detected [ 56 ], our results corroborate the hypothesis that the attraction of pheromones is not strong enough to be used for active eradication actions by mass-trapping, and probably not even for reliable monitoring. The low effectiveness of pheromone-based trapping techniques is probably due to the fact that they mainly attract virgin females and, at close range, females also used other visual and chemical stimuli which require further study [ 57 ].

Despite A. glabripennis having many of ecological and biological characteristics indicated by Brockerhoff et al. [ 39 ] as facilitating their eradication, an effective A. glabripennis eradication is never easy because of its extreme polyphagy and the generic symptoms it causes to host plants. First of all, A. glabripennis has a low fecundity rate [ 3 ]. In China, under natural conditions, 25–40 viable eggs were estimated per female [ 4 ], whereas in the USA that fecundity was estimated to vary between 30–178 eggs per female [ 58 , 59 ]. Additionally, their limited active dispersal capacity is an important factor. The potential dispersal of A. glabripennis adults was estimated at about 2000 m, with a realistic annual spread of about 300 m from the closest infested tree [ 56 , 60 , 61 ]. Moreover, the tendency to reinfest the same tree for several years was usually observed [ 3 , 4 ]. Lastly, according to climatic conditions A. glabripennis takes 1–3 years or even more to fully complete its life cycle [ 3 , 4 ]. All these characteristics (i.e., fecundity, active fly, and life cycle duration) strictly depend on temperature [ 62 ]. In Cornuda, annual temperatures range between −2 and 29 °C, with a mean temperature of 23 °C during warmer months [ 63 ]. In Paddock Wood annual temperatures vary between 2 and 23 °C, with a mean temperature of 17 °C during warmer months [ 64 ]. The first effects of the different climatic conditions at the two sites can be observed on the life cycle. In northern Italy A. glabripennis was considered univoltine [ 61 ], whereas for the UK a 2–3 year life cycle was estimated [ 64 ]. Moreover, research carried out on the effects of temperature on A. glabripennis fecundity estimated that the optimum temperature for maximum fecundity is about 25 °C [ 59 ]. Another study showed that the adult’s flight capacity increases with temperatures from 15 to 30 °C, and that no flight occurs under 15 °C [ 65 ]. In conclusion, lower temperatures of Paddock Wood caused a lower A. glabripennis adult fecundity, lower flight propensity (i.e., lower dispersion of the infestation), and lengthening of development time—doubling or even tripling it compared to the Cornuda infestation. All these factors probably contributed to keeping the pest infestation low in the UK, despite the eradication beginning about ten years after the estimated arrival of the pest [ 66 ], whereas in Italy only five years had elapsed.

One of the major problems in dealing with A. glabripennis eradication concerns its extreme polyphagy on woody broadleaves. An extensive investigation conducted in the Yinchuan region (China) found damage on trees belonging to 14 genera of broadleaves, although complete development has not been confirmed on all species listed as hosts [ 4 ]. However, host suitability differs in different continents: Populus and Salix are more suitable than Ulmus in China [ 4 ]; Acer and Ulmus are generally more suitable than Fraxinus in the USA [ 15 , 67 ]; in Europe the most suitable genera are, in decreasing order, Acer , Betula , Salix , Aesculus and Populus [ 18 ]. In the Cornuda infestation the most infested genera (by percentage) were Betula , Ulmus , Acer , and Salix (excluding Cercidiphyllum and Aesculus because of the small number of trees present)—similar to infestations in other Italian regions (Lombardy, Marche, and Piedmont) [ 35 ] and to the infestation at Paddock Wood, where the most-attacked genera were Acer , Salix , and Betula [ 43 , 44 ]. Interestingly, in both the Cornuda and Paddock Wood infestations, the number of infested poplars was very low; this is particularly evident in Italy, were only 2 out of 1709 poplars (0.1%) were found to be infested. In contrast, poplars are among the most suitable hosts for A. glabripennis in China, even if not all Populus species are equally susceptible to A. glabripennis [ 68 ]. Acer , instead, is confirmed as one of the main hosts for A. glabripennis . Such a large polyphagy has important consequences in the management of A. glabripennis infestations. A higher number of potential hosts means, on one hand, higher chances for the pest to reproduce and proliferate and, on the other hand, a higher number of trees to be surveyed and, if infested, to be felled and replaced. Moreover, A. glabripennis infests healthy and vigorous trees [ 3 , 4 ], which makes prevention more difficult, as keeping plants healthy and in good physiological condition does not prevent infestations.

The eradication of A. glabripennis from the municipality of Cornuda shows the importance of taking prompt, coordinated, and effective actions to contain the spread of the pest and to proceed with its systematic elimination from the infested area. Despite the considerably large size of the Italian infestation, the benefits obtained from the eradication of the pest have far exceeded the costs incurred for its implementation [ 69 ]. Of course, different scenarios may occur. For example, the A. glabripennis infestation occurring in Worcester (Massachusetts) seems to be too widespread now and the eradication, although it remains the goal, is of uncertain outcome and is taking enormous effort [ 46 ].

Of all the actions undertaken, those that proved the most effective were visual survey of susceptible trees (in order to find signs of infestation) and the felling and destruction of all the infested trees and nearby ones (in order to prevent the diffusion of the pest). Additionally, the destruction of felled trees by chipping is a very useful practice in order to kill all the larvae present inside the wood, as demonstrated by the wood chip analyses conducted by the University of Padua and already reported in other works [ 57 , 70 ]. On the other hand, the use of pheromone traps proved useless, as was also the case during the eradication in Paddock Wood [ 43 ]. Finally, the involvement and active participation of citizens and stakeholders is also of paramount importance [ 39 , 71 ]. If not properly motivated, ‘unpopular’ actions such as felling private trees and killing insects may lead to protests and non-cooperation by the population, jeopardizing the outcome of the whole operation. In this respect, the first report of the presence of A. glabripennis in Cornuda was made by a private citizen, demonstrating the importance of citizens’ cooperation in the interception of alien species.

Acknowledgments

We thank the phytosanitary office of the Regional Plant Protection Organization (RPPO) for data provided and their help in fieldwork; the staff of U.O. Forestale of Treviso for data provided and their help in fieldwork; and Oleg Kulinich (Department of Forest Quarantine, of the All-Russian Center of Plant Quarantine of Moscow) for providing Russian lures.

Author Contributions

Conceptualization, M.M. and M.F.; methodology, M.M. and M.F.; formal analysis, M.M.; writing—original draft preparation, M.M.; writing—review and editing, M.F.; funding acquisition, M.F. All authors have read and agreed to the published version of the manuscript.

This research was partially financed by the Regional Plant Protection Organization of the Veneto Region and by the DOR projects of the University of Padua.

Institutional Review Board Statement

Data availability statement, conflicts of interest.

The authors declare no conflict of interest. The funders had no role in the design of the study; in the collection, analyses, or interpretation of data; in the writing of the manuscript; or in the decision to publish the results.

Publisher’s Note: MDPI stays neutral with regard to jurisdictional claims in published maps and institutional affiliations.

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Description of life stages, sampling or scouting procedures, management options, acknowledgments, references cited.

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Asian Longhorned Beetle (Coleoptera: Cerambycidae), an Introduced Pest of Maple and Other Hardwood Trees in North America and Europe

The use of trade, firm, or corporation names in this publication is for the information and convenience of the reader. Such use does not constitute an official endorsement or approval by the U.S. Department of Agriculture or the Forest Service of any product or service to the exclusion of others that may be suitable.

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P. S. Meng, K. Hoover, M. A. Keena, Asian Longhorned Beetle (Coleoptera: Cerambycidae), an Introduced Pest of Maple and Other Hardwood Trees in North America and Europe, Journal of Integrated Pest Management , Volume 6, Issue 1, March 2015, 4, https://doi.org/10.1093/jipm/pmv003

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The Asian longhorned beetle, Anoplophora glabripennis (Motschulsky), threatens urban and forest hardwood trees both where introduced and in parts of its native range. Native to Asia, this beetle has hitchhiked several times in infested wood packaging used in international trade, and has established breeding populations in five U.S. states, Canada, and at least 11 countries in Europe. It has a broad host range for a cerambycid that attacks living trees, but in the introduced ranges it prefers maples. Identification, classification, and life history of this insect are reviewed here. Eradication is the goal where it has been introduced, which requires detection of infested trees using several approaches, including ground and tree-climbing surveys. Several agencies and researchers in the United States and Europe are evaluating the use of pheromone- and kairomone-baited traps. Control options beyond cutting down infested trees are limited. To date, the parasitoids and predators of this beetle have broad host ranges and are unlikely to be approved in the United States or Europe. An effective delivery system under development for entomopathogenic fungi appears promising. Systemic insecticides have been widely used in the United States, but the ability of these chemicals to reach lethal doses in the crown of large trees is disputed by some scientists, and the potential nontarget effects, especially on pollinators, raise concerns. The most practical approach for eradicating Asian longhorned beetle is to optimize trapping methods using semiochemicals for early detection to eliminate the insect before it spreads over large areas.

The Asian longhorned beetle, Anoplophora glabripennis (Motschulsky) (Coleoptera: Cerambycidae) ( Fig. 1 ), is an invasive, polyphagous wood-boring insect that is capable of destroying 30.3% of the urban trees in the United States at an economic loss of US$669 billion ( Nowak et al. 2001 ). The beetle’s ability to attack multiple genera of apparently healthy hardwood trees could dramatically alter urban and forest ecosystems. Early, accurate detection of this nonnative pest is crucial for discovering infestations before they become unmanageable.

Asian longhorned beetle female on Acer twigs, showing the typical black and white body coloration and faint blue iridescence on the legs.

A. glabripennis is a member of the family Cerambycidae, which are commonly known as the longhorned beetles ( Lingafelter and Hoebeke 2002 ). These beetles are characterized by having long antennae, which are approximately 1.5–2 times the length of the insect’s body. The subfamily, Lamiinae, is commonly referred to as flat-faced longhorns, which are exclusively xylophagous as larvae and phytophagous as adults and play an important role in nutrient cycling ( Özdikmen and Hasbenli 2004 ). The genus Anoplophora contains 36 species native to Asia ( Lingafelter and Hoebeke 2002 ).

Common names for Asian longhorned beetle in Asia include the starry sky beetle, basicosta white-spotted longicorn beetle, and smooth shoulder-star longicorn ( Commonwealth Agricultural Bureaux International [CABI] 2014 ). In France and Germany, the beetles are known as longicorne asiatique and Asiatischer Laubholzkäfer, respectively. In the United States, the common name was derived from one of its most notable features, the uniquely long antennae, and its geographic origin. Even though this fairly generic common name is not the most desirable, it became widely used in North America and so has been maintained to prevent any public confusion that would arise by changing it now.

Asian longhorned beetle adults are large, 17–39 mm in length, and although females tend to be larger than males, there is considerable variation in size both within and between the sexes ( Figs. 1 and 2 A and B, Lingafelter and Hoebeke 2002 ). Both sexes have 11-segmented antennae with an alternating white and black banding pattern. The tarsi may have a faint iridescent blue color. The pronotum has two large spines, while the elytra are shiny black and bear white or yellowish tan spots in variable patterns. Older females may have fewer or faded spots following prolonged mating and mate guarding, as the males will chew off the white hairs that create the spots. The decorative setae that create these spots closely resemble setae found on the antennae and other parts of its body.

Adult Asian longhorned beetles are large, have two sharp tubercles (spines) on the pronotum, and a distinct black-white banding pattern on the antennae. The legs and feet are pubescent with a white or blue iridescence. Spots on the elytra are highly variable in color, shape, and size. Males have longer antennae relative to their body and are generally smaller than females. The anterior elytra are smooth. (A) Asian longhorned beetle adult females. (B) Asian longhorned beetle adult males. Note the variation in sizes of males and females.

The sexes can be distinguished as follows: females are generally larger and have shorter antennae than the males, and there are some abdominal tip differences as well. When the antennae are pulled back over the body, five antennomeres (segments) of the male and only one to two antennomeres of the female extend past the apex of the elytra ( Lingafelter and Hoebeke 2002 ). From one-third to one-half of the upper side of each antennomere in both male and female antennae is covered with white hairs, but because male antennae are longer, the black sections appear longer in the male than the female, and this difference is even more pronounced on the underside where the white sections on the males may be almost absent ( Wang et al. 2004 ). The terminal ventral segment on the female has a strong notch in the middle and is densely fringed with hairs, while the terminal ventral segment on the male has a flatter margin, with almost no notch and short hairs ( Lingafelter and Hoebeke 2002 ).

Oviposition often begins on branches or the main trunk of the lower crown of a tree. Females chew an oviposition pit through the bark and lay a single egg just under the bark in the cambium ( Haack et al. 2006 , Fig. 3 ). Under the bark, the egg ultimately sits in the middle of a localized necrotic tissue zone, either caused by a compound the female deposits or by the process of raising the bark during oviposition ( Fig. 3 A). In most cases, this prevents the egg from being crushed by the formation of callus tissue as the tree attempts to heal the wound. Eggs resemble a large grain of rice ∼5–7 mm in length ( Haack et al. 2010 , Fig. 3 A). Often sap will ooze from oviposition pits and can stain the bark. Fresh oviposition pits are initially red to light brown ( Fig. 3 B) but darken as they weather ( Fig. 3 C). The shape of oviposition pits varies from a narrow slit (>1 mm in height) to an irregular oval pit up to 15 mm in diameter, depending on the bark thickness ( Smith et al. 2002 , Ric et al. 2007 ). Eggs require less time to hatch at higher temperatures ( Keena 2006 ). See Table 1 for minimum temperature development thresholds and degree-days required for each life stage.

Asian longhorned beetle oviposition sites and eggs. (A) A. platanoides bolt with bark removed showing eggs. Eggs are inserted beneath the bark. (B) An oviposition pit on A. saccharum. These pits can be found on branches and sometimes the main bole of infested trees. Fresh oviposition pits are red or light brown and may ooze sap. (C) Older oviposition pits on Acer rubrum . After a few months, oviposition pits darken.

Lower temperature development thresholds and degree-day requirements for various Asian longhorned beetle instars and life stages

T L , lower temperature development threshold measured in °C; DD 50 , degree days needed for 50% of insects to complete each life stage. Excerpt from Keena and Moore (2010) and Keena (2006) .

The basic morphology of the Asian longhorned beetle is similar to that of other cerambycids in the Lamiinae, with a cylindrical body and no legs ( Fig. 4 A). Larvae are segmented, elongate, and light yellow or white. The head has a large pair of heavily sclerotized, black mandibles. The pronotum contains a heavily sclerotized, raised plate with shallow pits on the dorsal side ( Cavey et al. 1998 , Fig. 4 A). Asian longhorned beetle larvae have a reported maximum body length and head capsule width of 50 and 5 mm, respectively ( Cavey et al. 1998 ), but head capsule widths up to 5.38 mm have been seen in the laboratory ( Keena and Moore 2010 ). Young larvae create galleries just under the bark, whereas larger larvae tunnel deep into the heartwood of a tree ( Fig. 4 B). Active larvae excrete a sawdust-like frass and wood shavings that are extruded from larval galleries in infested trees.

Larval stages of Asian longhorned beetle. (A) An older larva. Notice the shape of the plate on the pronotum. (B) Galleries created by larvae in Acer rubrum . Older instars are able to tunnel into the heartwood.

Asian longhorned beetle pupae can be found in pupal chambers in the sapwood about 1 cm below the bark in the spring (M.K. unpublished data). Larvae pack wood shavings and frass behind themselves to block the larval tunnel and the partially excavated exit hole in front of them ( Fig. 5 A). Pupae are the same color as larvae but have some traits that resemble those of the adults ( Figs. 2 , 4 A, and 5 B). Sclerotized adult structures begin to darken as pupae approach eclosion ( Fig. 5 B). During metamorphosis, the mandibles and eyes darken first, followed by the legs and antennae ( Fig. 5 B).

Asian longhorned beetle pupae. (A) Pupa in pupal chamber. Frass has been packed into the entrance tunnel with a partially prechewed exit tunnel from the chamber. (B) Various ages of pupae. Older pupae on the left have more sclerotized body parts compared with younger pupae on the right.

Distinguishing Asian Longhorned Beetles From Other Insects

Anoplophora chinensis (Föster), the citrus longhorned beetle, is a closely related beetle that has also gained notoriety as an invasive species ( Haack et al. 2010 , Fig. 6 A). Asian longhorned beetles can be differentiated from A. chinensis by the black scutellum and smooth elytra of the former ( Fig. 7 A vs. B). In the United States, pine sawyers ( Monochamus spp.) can be easily mistaken for the Asian longhorned beetle. Monochamus spp. bear no or only faint markings on their elytra, tend to be smaller, and usually emerge before the Asian longhorned beetle ( Fig. 6 B and C). Another cerambycid native to the United States, the cottonwood borer ( Plectrodera scalator F.), can also be mistaken for the Asian longhorned beetle, but can be distinguished by its black spots against a yellowish-white body ( Fig. 6 D).

Examples of Asian longhorned beetle look-alikes. (A) The invasive citrus longhorned beetle, Anoplophora chinensis . This beetle looks nearly identical to Asian longhorned beetle in shape and size but usually has rough, bumpy anterior elytra. Photo by Holly Raguza, Pennsylvania Department of Agriculture. (B) The white spotted sawyer, Monochamus scutellatus. This beetle is considerably smaller than Asian longhorned beetle and has a distinct, white scutellum. Asian longhorned beetle does not have a white scutellum. (C) The northern pine sawyer, Monochamus notatus , a native U.S. species, can be easily mistaken as Asian longhorned beetle. These beetles are generally smaller than Asian longhorned beetle and have longer antennae. The body of the northern pine sawyer is usually grayish-brown. Faint spots may be present on the elytra. (D) The cottonwood borer, Plectrodera scalator , can also be mistaken for Asian longhorned beetle. This beetle is native to the United States and found east of the Rocky Mountains. P. scalator has black spots against a yellow body.

Comparison of anterior elytra of Asian longhorned beetle and A. chinensis. (A) The anterior elytra of Asian longhorned beetle are smooth. The scutellum is covered with dark hairs. (B) The anterior elytra of A. chinensis . Note the bumps present in this area. The scutellum is also covered with white hairs. Photo by Holly Raguza, Pennsylvania Department of Agriculture.

Sequence-characterized amplified genes from random amplified polymorphic DNA (RAPD) fragments collected from a wing, leg, antennae, or frass can be used to differentiate the Asian longhorned beetle from other native and exotic cerambycid species, including A. chinensis , Monochamus scutellatus Say (white-spotted sawyer), P. scalator , Saperda tridentata Oliver (elm borer), and Graphisurus fasciatus (Degeer) ( Kethidi et al. 2003 ). Additional RAPD markers have been found to be useful in intraspecific differentiation of Asian longhorned beetle populations ( Gao et al. 2007 ). Seven mitochondrial DNA regions of the Asian longhorned beetle and three congenerics found in China have been evaluated and appear to be useful for identifying this species ( An et al. 2004a , b ).

Asian longhorned beetle larvae are not easily distinguished from other species of cerambycid larvae. Their throat ridge on the underside of the head near the mouthparts does not have distinct lateral margins ( Lingafelter and Hoebeke 2002 ). The dorsal abdominal ambulatory ampullae (raised bumps) form two distinct rings around a central lobe that has no bumps, while larvae of Monochamus spp. and P. scalator have indistinct rings. A key “with good color figures” to distinguish Asian longhorned beetle larvae from closely related beetle species found in Europe is available ( Pennacchio et al. 2012 ). Accuracy of larval stage identification can be greatly enhanced with the aforementioned genetic tools.

Mitochondrial DNA sequences have been used to evaluate the genetic diversity of Asian longhorned beetle introductions in the United States to determine if introductions were independent or expansions of previous infestations. Compared with Asian longhorned beetle populations in China, beetles introduced to the United States have limited genetic diversity as a result of bottlenecking, few separate introductions, or both ( Carter et al. 2009a ). This limited genetic diversity may also result from prior bottlenecking events in China where the Asian longhorned beetle has moved into man-made treed landscapes. Genetic analysis of Asian longhorned beetles from Ontario, Canada, found mitochondrial DNA haplotypes not present in U.S. populations, indicating that both of these introductions occurred independently ( Carter et al. 2009b ). Hardy–Weinberg equilibrium was not satisfied in the Canadian beetle populations, indicating nonrandom mating. Using RAPD analysis, Asian longhorned beetle introductions in Illinois and New York have also been determined to be independent ( An et al. 2004a , b ). Further work to refine the ability to determine the origin of each introduction is needed.

Distribution

The Asian longhorned beetle was first discovered in the United States in Brooklyn, New York, during August 1996 ( Haack et al. 1997 , Cavey et al. 1998 ). Following this initial discovery, it was also found in the greater New York City area, New Jersey, Massachusetts, Illinois, and Ohio ( Poland et al. 1998 , Haack 2006 , Haack et al. 2006 , Dodds and Orwig 2011 , U.S. Department of Agriculture, Animal and Plant Health Inspection Service [USDA-APHIS] 2013a ), and most recently in Babylon Township, New York ( USDA-APHIS 2013b ). The Asian longhorned beetle has been declared eradicated from Islip, Manhattan, and Staten Island, New York; Jersey City, New Jersey; Chicago, Illinois; Boston, Massachusetts; and Toronto, Ontario, although the Asian longhorned beetle was recently discovered in Toronto in late 2013 ( Canadian Food Inspection Agency [CFIA] 2013 ). Infestations have been found in other countries where eradication has also been the goal, including Austria, Belgium, England, France, Germany, Italy, Switzerland, and the United Kingdom ( European and Mediterranean Plant Protection Organization [EPPO] 2013 ).

The Asian longhorned beetle has a broad distribution throughout China and the Koreas ( Lingafelter and Hoebeke 2002 , Williams et al. 2004a ). It is most damaging in the Chinese provinces of Liaoning, Jiangsu, Shanxi, Henan, and Hubei ( Yan 1985 ) and can be found in a much larger area spanning from 21°–43°N and 100°–127°E. Global warming has allowed the Asian longhorned beetle to spread northwards in China ( Wang et al. 2011 ). Niche modeling indicates it is most likely to establish in central and eastern China, the Koreas, and Japan ( Peterson et al. 2004 ), with a slight chance of establishment in southeast China and east India.

Niche modeling has been used to predict the susceptibility of U.S. forests to Asian longhorned beetle establishment. Most of the eastern United States is at risk ( Peterson and Vieglais 2001 , Peterson et al. 2004 ). Epidemic simulation models based on habitat suitability and known entry points (ports and warehouses) indicate that the Great Lakes region is most likely to be an initial invasion point ( Peterson et al. 2004 ). Proximity to transportation corridors, such as roads, and a high probability of preferred host presence are significant predictors of an Asian longhorned beetle infestation ( Shatz et al. 2013 ).

Given the lack of suitable hosts outside of urban areas, the western United States is less susceptible to establishment despite the greater volume of cargo arriving from Asia compared with other parts of the United States ( Peterson et al. 2004 ). This species is not predicted to establish in Mexican or northern Canadian forests due to the general lack of the most preferred North American host trees. Cold temperatures are unlikely to limit the range of the Asian longhorned beetle . The beetle has a supercooling point of −25.8°C and is freeze tolerant, with at least 92% of larvae being able to survive temperatures of −25°C or lower for 24 h in laboratory experiments ( Roden et al. 2009 ).

Life History

After a female chews an oviposition pit in the bark during the summer or early fall, she lays an egg beneath the bark into the phloem. Mortality is highest in the egg and first instar, as these life stages occur close to the plant surface, leaving Asian longhorned beetles vulnerable to extreme temperatures, host responses, and natural enemies ( Tang et al. 1996 ). Asian longhorned beetle eggs take 54.4 ± 0.7 to 13.3 ± 0.7 d to hatch at temperatures ranging from 15 to 30°C with less time needed at higher temperatures ( Keena 2006 ). Percent egg hatch is the highest at 25°C (63.6 ± 6.8%) and rapidly decreases at temperatures above or below the optimum. Asian longhorned beetle eggs do not hatch if held at 5, 10 or 35°C and require 239 degree-days for 50% of the eggs to hatch ( Keena 2006 , Table 1 ). Eggs that are laid in late summer or early fall do not have time to hatch; instead these eggs overwinter and hatch when it warms up again the next year. Females usually lay eggs only when the air temperature is between 15 and 30°C.

Larvae require 1–2 yrs to develop before reaching a critical weight and pupating. In colder, northern regions of China, the Asian longhorned beetle is more likely to take 2 yrs to develop ( Hua et al. 1992 ). There are five or more instars, which varies depending on host species, host condition, and temperature. In the laboratory, larval molts have numbered as high as 20 ( Keena and Moore 2010 ). As with eggs, larval development time depends on host species ( Smith et al. 2002 ) and temperature, with time in each instar shortening as temperatures increase up to 30°C and with no development occurring at ≤10 or ≥40°C ( Keena and Moore 2010 ).

Pupal eclosion is temperature-dependent and can take 12–50 d at temperatures ranging from 15–30°C ( Keena and Moore 2010 ). Prepupae kept at low temperatures (10°C) may fail to pupate until exposed to higher temperatures. The lower temperature development threshold for pupae is estimated to be 10°C ( Table 1 ). About 256 degree-days (lower threshold 10°C) are needed for 50% of pupae to develop into adults ( Keena and Moore 2010 ).

Following adult eclosion, beetles spend 4–7 d undergoing sclerotization before initiating chewing out of trees, and another 4–5 d to complete emergence ( Sánchez and Keena 2013 ). In Ningxia, China, peak emergence occurs from late June to early July, with a smaller peak occurring in mid-August ( Zhang and Xu 1991 ). In the New York City infestation the first recorded beetles were found between June 26 and July 29; peak emergence generally also occurred during this time but some occurred even in September ( Auclair et al. 2005 ).

Temperature and host species have significant effects on Asian longhorned beetle longevity and fecundity. In the laboratory, females and males can survive up to 158 and 202 d, respectively, when reared at 20 °C on Acer saccharum Marsh ( Keena 2006 ). Optimal longevity occurs when beetles are kept at 18°C ( Keena 2006 ). Maximum fecundity occurs when beetles are kept between 23–24°C, depending on their geographical origin, and more eggs hatch when held at 25°C than at other temperatures. Using infested bolts cut from quarantine areas in New York and Chicago, fecundity of emerged females was evaluated under laboratory conditions. Lifetime production of viable eggs per female was 45–62 eggs on average ( Keena 2002 ). In another laboratory-based study, the number of viable eggs laid increased with increasing female body size, but decreased as a function of beetle age, the diameter and area of the oviposition bolt, and the thickness of the bark ( Smith et al. 2002 ). Females fed on twigs of Acer platanoides L. or Acer rubrum L. had longer survival and greater fecundity than females fed on Salix nigra Marshall ( Smith et al. 2002 ).

Adults remain inactive and perch in tree crowns during the early morning and late afternoon, and are inactive during inclement weather and high temperatures ( Zhou et al. 1984 ). Under greenhouse conditions, adults were least active in the morning when it was cool, rested in the shade during the heat of the day, and were most active later in the day from 8 p.m. to midnight ( Morewood et al. 2004 ). Both sexes spent most of their time “in decreasing frequency” resting, walking, or feeding. Males mate multiple times with multiple females and guard inseminated females for several hours. Adults feed on leaf petioles and debark small branches, also feeding on the cambium. Adults tend to re-infest their natal host as long as it is alive, rarely fly, and instead walk as their primary means of locomotion ( Zhou et al. 1984 ). Possible reasons that beetles fly are in response to declining host quality (M.K., unpublished data), in response to landing on nonpreferred hosts, after experiencing an aggressive encounter with a conspecific (M.K., unpublished data), to locate a mate, when disturbed by a potential predator, or to find suitable environmental conditions.

Larvae live in and consume the sapwood and heartwood of susceptible host species. A single Asian longhorned beetle larva can consume 1,000 cubic cm of wood ( Yan and Qin 1992 ). When 3-yr-old Populus euramericana Guinier trees were infested with Asian longhorned beetles for 3 yrs, they had a 22–49% and 5–25% decrease in the diameter of the truck at breast height (DBH) and vertical height, respectively ( Gao et al. 1993 ). Mounting evidence suggests that the beetle is able to thrive in the sapwood and heartwood of healthy trees by harboring a diverse gut microbial community that facilitates lignocellulose degradation and nutrient acquisition ( Geib et al. 2008 , 2009 ; Scully et al. 2013 ; Ayayee et al. 2014 ).

The Asian longhorned beetle typically begins attacking the crown of a host tree along main branches ( Haack et al. 2006 ), making it difficult to detect in the first year or two of the infestation. Eggs are laid in an aggregated pattern along main branches and the lower crown ( Li et al. 2012 , Ma et al. 2012 ). In heavily infested trees or those with continuous branches beginning at the base, beetles oviposit on the lower trunk. A female-produced trail pheromone is hypothesized to act as a spacing pheromone that may deter repeated oviposition in the same location ( Hoover et al. 2014 ). Asian longhorned beetles also begin attacking the trunk as the crown dies from a heavy infestation ( Lingafelter and Hoebeke 2002 ).

Adult Dispersal

Spread of the Asian longhorned beetle within a given location has been modeled; in a 30-d mark, release, recapture study in China, the average dispersal of Asian longhorned beetles from the center of a field was 106.3 m ( Junbao et al. 1988 ). Weekly sampling of forest trees in Liu Hua, China, for 3.5 mo revealed that male and female beetles can disperse up to 1,029 and 1,442 m, respectively ( Smith et al. 2001 ). Ninety-eight percent of the recovered beetles were captured within 560 m of the release site. In a follow-up study performed at the same location with more distant recapture points, dispersal potential for males and females was calculated to be 2,394 and 2,644 m, respectively, but 98% of beetles were recaptured within 920 m from the release point ( Smith et al. 2004 ). A study utilizing harmonic radar showed that beetles moved an average of 14 m in a 9–14-d interval with a bimodal distribution of 0 and 90+ meters; males moved six times the distance compared with females ( Williams et al. 2004b ). Recaptured beetles tend to move upwind ( Li et al. 2010 ), perhaps in response to plant or conspecific odors. USDA-APHIS requires quarantine boundaries to extend 2,400 m from the nearest infested tree, which encompasses the dispersal potential of nearly 100% of Asian longhorned beetles ( [USDA-APHIS-PPQ] 2014 ).

In poplar forests, the Asian longhorned beetle is more likely to be found along forest edges than in the forest interior ( Liu et al. 2012 ). High canopy density and high vegetation coverage reduce Asian longhorned beetle densities. In Ningxia, China, populations were lowest 12 m inside a Populus alba L. stand and peaked 2 m from the forest edge ( Wei et al. 1997 ). Trees on the eastern side of the stand were the least damaged by the Asian longhorned beetle. In mixed stands, the distribution of Asian longhorned beetles can vary according to tree species composition ( Wang et al. 2006 ). For example, in a stand containing P. alba var. pyramidalis and Acer negundo L., damage by Asian longhorned beetles followed an aggregated distribution. The oldest oviposition pits were found on the outside of the stand, while the active frass holes were in the center of the stand, indicating the infestation started on the margins and moved toward the center of the stand. In a pure stand of Populus simonii x Populus nigra var. italica , damage caused by the Asian longhorned beetle was randomly distributed. In closed canopy mixed hardwood stands in the Worcester, Massachusetts, regulated area, Asian longhorned beetle-infested Acer trees were found throughout the stand, indicating that although they may have started along the edges, they can quickly move into a stand ( Dodds et al. 2014 ). In addition, in these stands the codominant and dominant size classes of Acer rubrum L. were more frequently attacked compared with other Acer species and size classes that were present.

The Asian longhorned beetle is known to attack >100 different tree species in the wild with preference for those in the genera Acer , Populus , Salix , and Ulmus ( MacLeod et al. 2002 , Haack et al. 2006 , Hu et al. 2009 ). See Table 2 for a list of genera known to serve as Asian longhorned beetle larval hosts (one of which has a single documented infestation on one species) and Supp Supplementary Data (online only) for an extended list of tree species on which Asian longhorned beetle feeds, oviposits, or completes development under field conditions . ( Wu and Zhang 1966 ; Gao et al. 1993 ; Lou et al. 1993 ; Zhao et al. 1993 ; Wang and Zhou 1994 ; Tang et al. 1996 ; Gao et al. 1997 ; Bai and Zhang 1999 ; Wang et al. 2000b , 2009 ; Maspero et al. 2007 ; Qiao et al. 2007 ; Tomicezk and Hoyer-Tomiczek 2007 ; Guo 2008 ; Yan et al. 2008 ; Hu et al. 2009 ; Jin and Sun 2009 ; Tian et al. 2009 ; Zhao et al. 2011 ; Bund Schweizer Baumpflege [BSB] 2012 ; Jiang et al. 2012 ; Yan and Liu 2012 ; U. S. Department of Agriculture, Forest Service [USDA-FS] 2012 ; CFIA 2013 ; EPPO 2013 ; Dodds et al. 2014 ; CABI 2014 ; USDA-APHIS-PPQ-CPHST 2015 ). In the United States, the Asian longhorned beetle has also been documented to infest tree species in the genera Aesculus , Hibiscus , and Betula ( Haack et al. 1996 ), but strongly prefers maples, which are ubiquitous across the American landscape, including Acer platanoides L. (Norway maple), Acer saccharum Marsh (sugar maple), Acer saccharinum L. (silver maple), Acer rubrum (red maple), and Acer negundo (boxelder) ( Haack et al. 2006 ). The most recent new host record was recorded in Worcester, Massachusetts, the katsura tree in the genus Cercidiphyllum (U.S. Department of Agriculture, Animal and Plant Health Inspection Service, Plant Protection and Quarantine [USDA-APHIS-PPQ] 2010 ). In China, the beetle is a major pest in monoculture Populus plantations and has been referred to as the “forest fire without smoke” ( Li and Wu 1993 ), but susceptibility among poplar species and hybrids varies considerably ( Hu et al. 2009 ).

Tree genera (families) that serve as larval hosts for Asian longhorned beetle

a Only one report of a single species from this genus reported as infested.

b Susceptibility to Asian longhorned beetle varies greatly in Populus spp.

Signs of Asian Longhorned Beetle Infestation

Signs of Asian longhorned beetle infestation include oviposition pits ( Figs. 3 B and C and 8 A), frass ( Fig. 8 B), exit holes ( Fig. 8 C), oozing sap, twigs with stripped bark ( Fig. 8 D), and galleries in the sapwood and heartwood ( Haack et al. 2010 , Fig. 4 B). Thus, severe infestations can compromise the structural integrity of a tree. Since older (and bigger) larvae spend most of their time in the heartwood, trees can sustain infestations without visible signs of attack. Asian longhorned beetle infestations are often first detected and brought to the attention of government officials by members of the public.

Signs of Asian longhorned beetle infestation in Betula platyphylla Suk in Harbin, China. (A) Oviposition pits concentrated along a main branch and the trunk in the lower crown. In Betula spp., oviposition pits become black by the following season. Recently chewed pits are brown colored. (B) Frass being extruded from oviposition pits. Frass will also accumulate on the ground beneath a tree. (C) Perfectly round exit hole on a small branch. This is uncommon. Exit holes are typically found on larger branches and the main trunk. (D) Adult feeding damage on A. rubrum twigs. Beetles debark twigs to gain access to the cambium.

Detection and Eradication Methods

Several methods have been developed to detect and delimit Asian longhorned beetle populations. The most commonly used approach utilizes specialized ground surveyors equipped with binoculars to locate signs of Asian longhorned beetle damage on a tree. During these surveys all trees with a stem diameter of >2.5 cm are inspected ( Ric et al. 2007 ), as this beetle has been found in trees as small as 8 cm in diameter in the United States ( Haack et al. 2006 ) and as small as 3.3 cm in diameter in Canada ( Kimoto and Duthie-Holt 2004 ). According to USDA environmental assessment reports, accuracy of detection by ground surveyors is about 30% as determined by quality assurance checks ( USDA-APHIS 2013a ). Effectiveness of ground surveyors in detecting trees with signs was found to improve when sign density increased, signs were below 2.5 m in height on the tree, and when oviposition pits were located on boles and exit holes on branches ( Turgeon et al. 2010 ). Tree climbers can detect signs of Asian longhorned beetle infestation with higher rates of accuracy, ranging from 60 to 75% ( USDA-APHIS 2013a ), but this method is more costly and slower than ground surveys ( Hu et al. 2009 ). Hydraulic lifts are also used to survey for evidence of Asian longhorned beetle infestation and are more effective than ground surveys alone.

Lure-baited traps can be used to detect Asian longhorned beetles in the field. A two-component volatile male-produced pheromone consisting of a 1:1 ratio of 4-(n-heptyloxy)butanal and 4-(n-heptyloxy)butan-1-ol was identified ( Zhang et al. 2002 ) and has been under evaluation for several years. The pheromone alone is not significantly attractive ( Nehme et al. 2009 ), but addition of a plant kairomone mixture containing ( - )-linalool, ( Z )-3-hexen-1-ol, and trans -caryophyllene with or without addition of trans -pinocarveol or linalool oxide significantly increased trap catches of female Asian longhorned beetle in field studies ( Nehme et al. 2010 , 2014 ). This finding is consistent with several cerambycid trapping studies that demonstrated a synergistic effect when pheromone and host plant kairomones were released together, resulting in a larger number of beetles being captured compared with traps baited with pheromone or plant kairomones alone ( Pajares et al. 2004 , 2010 ; Reddy et al. 2005 ).

In a recent field study in China, increasing the release rate of plant volatiles in combination with the male-produced pheromone increased the number of female beetles trapped compared with lower release rates ( Meng et al. 2014 ). Other semiochemicals produced by Asian longhorned beetle include a female-produced, sex-specific trail pheromone ( Hoover et al. 2014 ), a female-produced contact pheromone ( Zhang et al. 2003 ), and a punitive third component to the male-produced pheromone ( Crook et al. 2014 ), but whether these semiochemicals would have any use in management has not been determined.

Acoustic detection of the Asian longhorned beetle has been investigated by two different groups. When Asian longhorned beetle larvae feed inside of trees they produce distinctive sounds that can be picked up by sensors attached to the infested tree and distinguished by use of software calibrated for the particular frequency and pattern of sounds ( Haack et al. 2001 , Mankin et al. 2008 ).

Sniffer dogs have been trained to detect the Asian longhorned beetle and A. chinensis in the United States and Europe ( Errico 2012 , Hoyer-Tomiczek and Sauseng 2013 ). Dogs can detect frass odors on tree trunks and exposed roots. A team of four dogs can inspect 15,000 Acer seedlings in a nursery in 3 d ( Hoyer-Tomiczek and Sauseng 2013 ), and a team of three dogs can search a 5-acre neighborhood in 20 min ( Errico 2012 ). In controlled double-blind laboratory trials, trained dogs were able to detect Asian longhorned beetle frass 80–90% of the time ( Errico 2012 ).

The only effective method to eradicate Asian longhorned beetle is to completely remove and chip or burn infested trees and grind stumps ( Wang et al. 2000a , b ). From 1998 to 2006, the costs to detect, remove, and prophylactically treat host trees with imidacloprid was US$249 million dollars in the United States alone ( United States Government Accountability Office [GAO] 2006 ). Full host removals of infested and adjacent healthy trees may prevent the spread of the Asian longhorned beetle but are controversial with the public. Early detection of invasive pests both increases the likelihood of successful eradication as well as decreases control costs ( Mehta et al. 2007 ). The Asian longhorned beetle infestation in Worcester, Massachusetts, was not detected until 8–10 yrs after the beetle’s likely introduction ( USDA-APHIS 2010 ). As of 2013, the quarantined area now spans 110 square miles in six jurisdictions. Thus, there is a strong need for developing sensitive Asian longhorned beetle detection methods that can delimit the extent of the beetle infestation before it becomes unmanageable.

A review of the Asian longhorned beetle eradication programs suggests that emphasis on minimizing potential pathways for new introductions, maintaining public awareness, and continuing to develop methods for early detection and rapid response are key to reducing the threat posed by this insect ( Smith et al. 2009 ). Public outreach is critical ( GAO 2006 , Ciampitti and Cavagna 2014 ). Members of the public have been responsible for locating and bringing to authorities’ attention the beetle or its signs in each North American location that a breeding population has been found. In addition, cooperation from the public and local governments is critical to eradication efforts.

Cultural or Mechanical Control

Cultural control methods to manage the Asian longhorned beetle have been developed in China. Elaeagnus angustifolia L. and Acer mono Maximowicz are planted in Populus plantations as trap trees and then cut down and burned prior to adult emergence ( Feng et al. 1999 ). Acer negundo can be used to protect Salix plantations from the Asian longhorned beetle and A. chinensis by luring the beetles to a centralized location and then manually removing trees ( Xu and Wu 2012 ). Trap trees not removed from plantations may cause a temporary reduction in Asian longhorned beetle populations for 1 yr and then lead to a sharp increase in beetle populations the following year ( Xu and Wu 2012 ). Tilia mongolica Maximowicz is another trap tree that can be used to help control the Asian longhorned beetle because it draws adults from up to 10 m away and then inhibits egg and larval development ( Tian et al. 2009 ).

Host Plant Resistance

Considerable efforts have been devoted to breeding poplar cultivars resistant to borers. Populus deltoides ‘Lux (I-69/55)’ has a high water content in the sapwood that confers resistance to Asian longhorned beetles ( Qin et al. 1996 ). Egg chambers fill with sap, which inhibits insect development or may kill the eggs. On other resistant poplars, callus tissue overgrows the oviposition wounds quickly and can prolong egg hatch or kill eggs ( Wang and Zhou 1994 ). Planting mixed stands of susceptible and resistant poplar cultivars can dramatically reduce the proportion of infested trees ( Jia and Li 2008 ). In the United States, a list of trees to use for replanting in Asian longhorned beetle-infested areas and a list of nonhosts have been compiled ( USDA FS 2014 and Supp Supplementary Data [online only] ( Wang et al. 2000b , 2009 ; Lazarus 2003 ; Morewood et al. 2004 ; USDA-FS 2014 , respectively).

Biological Control

Entomopathogens and other biological control agents have been evaluated for their efficacy against the Asian longhorned beetle and may be environmentally safer than conventional control methods utilizing pesticides. However, in regulated areas in North America and Europe where eradication is the goal, classical biological control is not the first option. In its native range in China, biological control has been incorporated into an integrated pest management approach. Unfortunately, several of the biological control agents have a broad host range and would be capable of attacking native woodborers if introduced to North America or Europe.

Several entomopathogens have been evaluated for their efficacy against the Asian longhorned beetle and other invasive species ( Hajek and Tobin 2010 ). Beauveria brongniartii (Saccardo) Petch is a fungus native to North America that has been shown to increase mortality and decrease fecundity in the Asian longhorned beetle ( Dubois et al. 2004a , b ). However, the commercial strain of B. brongniartii from Japan was found to belong to the taxon Beauveria asiatica (Saccardo) Petch ( Goble et al. 2014 ). Given the reclassification of B. asiatica and lack of natural North American isolates, introduction of this strain as a biological control in the United States is unlikely. The pathogenicity of Metarhizium anisopliae was also tested against the Asian longhorned beetle but was less promising than the two aforementioned Beauveria spp. ( Hajek et al. 2006 ).

Goble et al. (2014) found that exposure to Metarhizium brunneum and B. asiatica decreased median adult beetle survival time to 7.5–9.5 d compared with 24–31 d for B. brongniartii . Given the high virulence of M. brunneum against Asian longhorned beetle adults and its current registration in the United States, research has focused on the formulation and delivery of this fungus using oil, agar, and different fungal band textures ( Ugine et al. 2013a , b ). Finding an entomopathogenic fungal biological control agent for this beetle that can be used in the United States is still a possibility, but as with all fungi, efficacy when exposed to high temperature and low humidity environments may be diminished.

Bacillus thuringiensis Berliner toxins have been tested in a laboratory setting and were found ineffective against the Asian longhorned beetle when incorporated into artificial diet or delivered directly to beetles with a micropipette ( D’Amico et al. 2004 ). The entomopathogenic nematodes Steinernema feltiae (Filipjev) Wouts, Mracek, Gerdin & Bedding; S. glaseri Ssteiner; S. riobrave Cabanillas, Poinar & Raulston; S. carpocapsae (Weiser) Wouts, Mracek, Gerdin & Bedding; and Heterorhabditis marelata Liu and Berry were tested against early instar Asian longhorned beetles ( Xixiang et al. 1988 , Fallon et al. 2004 ). Only S. feltiae and S. carpocapsae were effective, and only against mid- to late-instars. Methods to apply the nematodes to infested trees so that they can survive and find the beetle larvae successfully are still needed to make this a viable option.

Predators and parasitoids in the native range of the Asian longhorned beetle have also been evaluated for control potential. Dastarcus helophoroides (Fairmaire) is a parasitoid of cerambycids in China with a broad host range and has been used to control the Asian longhorned beetle in its native range ( Li et al. 2007 , Wei et al. 2009 ). Populations of this parasitoid from different regions of China are differentially attracted to Asian longhorned beetle frass ( Wei and Jiang 2011 ). D. helophoroides also parasitizes Monochamus alternatus Hope, a vector of the pinewood nematode ( Bursaphelenchus xylophilus (Steiner & Buhrer) Nickle, Miura et al. 2003 ). Because D. helophoroides has a wide host range, it would be an unlikely candidate for classical biological control. Sclerodermus guani Xiao and Wu, a hymenopteran parasitoid, is highly effective against Cerambycidae in China and can reduce Asian longhorned beetle populations up to 45% in Populus plantations ( Fu et al. 2010 ). However, this parasitoid is known to attack and develop in honey bees so it would not be considered for use in the United States ( Yao and Yang 2008 ).

Some native European hymenopteran parasitoids that immobilize and feed on the outside of Asian longhorned beetle larvae (idiobiont ectoparasitoids) have been found to host-shift to Asian longhorned beetles and could be candidates for biological control. The most common parasitoids of Asian longhorned beetle larvae in Italy were Spathius erythrocephalus Wesmael and Trigonoderus princeps (Westwood) ( Hérard et al. 2013 ). No Asian longhorned beetle egg parasitoids were found in Italy. Similar studies are ongoing in the United States and three natural enemy species can complete some development in Asian longhorned beetle ( Smith et al. 2007 ).

Woodpeckers are well known to prey on Asian longhorned beetle in China ( Gao et al. 1994 ); artificial cloth bird nests can be used to attract the woodpecker Dendrocopos major L. to susceptible trees ( Cheng et al. 2010 ), but woodpeckers alone cannot eliminate the Asian longhorned beetle. Woodpecker activity has been observed in the United States on trees infested with Asian longhorned beetle and can indicate potential infestations .

Chemical Control

Conventional methods using insecticides to control the Asian longhorned beetle have been used in both the native and introduced ranges. In China, cypermethrin is widely used to kill adult Asian longhorned beetles in host tree canopies ( Liu et al. 1999 ). Clothianidin, dinotefuran, and thiamethoxam can also be used to control Asian longhorned beetles via ingestion, contact, and antifeedant effects ( Wang et al. 2005 ).

Neonicotinoids for control of the Asian longhorned beetle have been studied in the laboratory and field for several years. In laboratory studies, 60% of larvae that were fed artificial diet treated with 50 ppm azadirachtin or 1.6 ppm of imidacloprid showed decreased longevity ( Poland et al. 2006a ). One hundred percent of Asian longhorned beetle adults died in 13 and 20 d while feeding on maple twigs dipped in 150 and 15 ppm of imidacloprid, respectively. This contrasts with a report by Ugine et al. (2011) who reported that adult female Asian longhorned beetles fed 1 μl of 10 or 30 ppm of imidacloprid per day failed to die faster than control beetles, but laid 23–38% fewer viable eggs because they spent time recovering from intoxication. However, females administered 1 μl of 40 or 50 ppm of imidacloprid per day died faster than control beetles and laid significantly fewer eggs.

In field studies performed in 2000 and 2001 in China, tree injection with imidacloprid and thiacloprid reduced populations of Asian longhorned beetles in several host species ( Salix spp., Ulmus spp., and Populus spp.), but the efficacy declined over time following treatment ( Poland et al. 2006b ). In a separate study in the United States, beetles were fed twigs from A. platanoides trees that were trunk-injected with imidacloprid in the Worcester, Massachusetts, infestation. Adult beetles fed twigs from treated trees had significantly lower fecundity and survival compared with beetles fed untreated twigs, with an LC 50 over 21 d of 1.3 ppm; yet, during the course of the study, only 35% of all beetles died ( Ugine et al. 2012 ). Imidacloprid concentrations in twig samples varied widely, both seasonally and between trees ( Ugine et al. 2012 ). During the course of the study, over half of twig samples from injected trees contained <1 ppm of imidacloprid and 37% contained no detectable levels. Twigs from trees injected in the spring also showed a decline in insecticide concentration as the season progressed into the fall. Leaves were only sampled in the fall and had a higher concentration of imidacloprid than twigs; adult Asian longhorned beetles feed on both twigs and leaves.

Imidacloprid seemed to produce an antifeedant response in Asian longhorned beetle larvae in the study by Poland et al. (2006a) . However, in a choice test from a different study, adults did not have a stronger preference for control twigs compared with those obtained from imidacloprid-injected trees ( Ugine et al. 2012 ). The discrepancies between these two studies may reflect differences in either methods or host tree species characteristics (e.g., size, species, soil conditions).

Although imidacloprid and other neonicotinoids may be an option to prophylactically treat and protect susceptible trees, these pesticides do not translocate evenly or quickly within a tree and have sublethal effects on nontarget organisms ( Tattar et al. 1998 , Kreutzweiser et al. 2008 ), with particular concern for pollinators ( Biddinger et al. 2013 ), although many Asian longhorned beetle hosts are wind pollinated. In addition, a study of the nearly 250,000 at-risk trees treated in the New York, Illinois, and New Jersey regulated areas found that 11 trees had strong evidence that some Asian longhorned beetles may have escaped the effects of imidacloprid treatments resulting in nine adults emerging (Sawyer 2006). Given the high cost of treating large numbers of trees with systemic insecticides and the potential for nontarget effects, additional controlled studies on a broader scale in the United States are needed to maximize the likelihood of successful control in the context of complex environmental and biological factors that can impact this approach.

Summary of Management Options

Successful eradication efforts for the Asian longhorned beetle hinge on accurate and timely detection of infested trees, methods to kill the beetle in infested trees, and ways of preventing it from spreading. Currently, detection of infested trees is accomplished through ground and tree-climbing surveys, but over the past decade a lure and trap combination has been developed that can aid pest management practitioners and eradication programs in locating infestations and allocating resources to halt the beetle’s spread. Control options beyond cutting down infested trees are limited. To date, the parasitoids and predators that have been found attacking this beetle have broad host ranges and are unlikely to be approved for release in the United States or Europe. An effective delivery system under development for entomopathogenic fungi seems promising. Systemic insecticides have been widely used in the United States, primarily to protect trees from becoming infested, but the ability of these chemicals to reach lethal doses in the crown of large trees is disputed by some scientists, and the potential nontarget effects, especially on pollinators, raise concerns. Managing Asian longhorned beetle via other chemical methods holds some promise, but widespread chemical treatments would be cost prohibitive and could cause unpredictable nontarget effects.

Concluding Remarks

Failure to eradicate Asian longhorned beetle where it has been introduced or to control it in altered habitats in the native range could be devastating to both forest and urban trees. In its native China, Asian longhorned beetle causes nearly 12% of the total losses attributable to forest pests, costing an estimated US$1.5 billion annually ( Hu et al. 2009 ). In the United States, it is estimated that 12–61% of all urban trees are at risk, with total estimated value of US$669 billion dollars ( Nowak et al. 2001 ). In 2002, two of the maple hosts favored by Asian longhorned beetle were among the top 10 most common forest trees in U.S. forests: Acer rubrum (7.6% of all trees) is the most common, and Acer saccharum (3.1% of all the trees) the sixth most common ( USDA-FS 2002 ). Thus, the value of the trees at risk in the United States alone if the Asian longhorned beetle were to freely infest all susceptible hosts is monumental, serving as an important factor in the decision to eradicate this insect where it has been introduced both in North America and Europe.

We thank R. Haack for reviewing the manuscript and providing very useful suggestions. We thank Alphawood Foundation for funding to K.H. for Asian longhorned beetle research at Pennsylvania State University. The Frost Entomological Museum and Pennsylvania Department of Agriculture provided specimens for imaging.

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Invasive Species Centre

Asian Longhorned Beetle

Asian longhorned beetle (ALHB) has been eradicated from the cities of Mississauga and Toronto in the province of Ontario. After five years of surveys with no detection of this plant pest, the  Asian Long-horned Beetle Infested Place Order  has been repealed effective June 9, 2020.

Asian Longhorned Beetle (Anoplophora glabripennis) French common name: Longicorne asiatique

Asian longhorned beetle has irregular white spots along its body.

ALB attack maple and other hardwood trees.

ALB are shiny and black with prominent, irregular white spots, 2-4 cm in length.

Order:  Coleoptera

Asian longhorned beetle (ALB) was first discovered in North America at several ports in the early 1990s. ALB was detected in the Toronto, Ontario area in 2003 and quick action lead to eradication of the pest over the following years. Unfortunately, a new infestation was discovered in Mississauga, Ontario in 2013. After five years of surveys with no detection of this plant pest, the  Asian Long-horned Beetle Infested Place Order has been repealed effective June 9, 2020 .

The destructive nature and wide variety of hardwood tree hosts of the ALB make it especially concerning for Canada. The ALB readily attacks and kills maple trees, which are widespread across natural and urban areas in Canada. Maple trees are culturally and economically important to Canadians, and the loss of this species could have devastating impacts.

It is important to remain vigilant as individual detections (not established populations) do occur.

Physical Description

Adult : Shiny black with prominent, irregular white spots, 2-4 cm in length. It has distinct bluish-white legs and long black and white segmented antennae. In Canada, the adult ALB can easily be mistaken for several native beetles that look similar, such as the white-spotted sawyer ( Monochamus scutellatus).  ( OFAH/OMNR Invading Species Awareness Program, 2012 )

Larvae :  Round, grub-like, and creamy white-coloured.

Photo: Taylor Scarr, OMNRF

Female adult beetles chew a shallow, oval-shaped pit into the bark or branches of a host tree to deposit an egg. Single eggs will be laid at multiple sites, with the possibility of one female laying up to 100 eggs. Eggs resemble grains of rice, and could take a week to several months to develop into larvae. The developed larvae initially feed in the cambium region under the bark, and eventually tunnel deeper into the tree. The tunnels created by the larvae hinder the tree’s ability to transport water and nutrients and eventually kills it. The larvae overwinter in the tree, until pupation occurs in April or May. Adult beetles emerge from late May to July, through large (1.5-2 cm diameter) exit holes in the bark ( City of Toronto, 2015 ). The rate of development depends on temperatures during the growing season; therefore, a portion of the population can take two years, rather than one, to complete its life cycle. ( OMRNF, 2014 )

“Photos: (1)Melody Keena, USDA Forest Service, Bugwood.org, (2) Kenneth R. Law, USDA APHIS PPQ, Bugwood.org (3) Pennsylvania Department of Conservation and Natural Resources – Forestry , Bugwood.org and (4) Donald Duerr, USDA Forest Service, Bugwood.org

Signs of infestation by ALB include ( City of Toronto, 2015 ): 

• Oval-shaped egg pits dug on trunk, branches, or exposed roots
• Leaking sap
• Large round exit holes (1.5-2 cm diameter)
• Yellowing leaves
• Premature leaf drop
• Branch dieback

Asian longhorned beetles create large (1.5 – 2 cm diameter) exit holes on the trunk and branches of host trees.

Asian longhorned beetle (ALB) is an insect native to several Asian countries, including China and Korea. The beetle was introduced into Canada when infested wood from plantations was used as packaging material for cargo being shipped to North America. ALB was first detected in the US and Canada in the early 1990s at several ports. To date, 15 states, British Columbia, and Ontario have reported ALB interceptions ( City of Toronto, 2015 ). Currently there are ALB infestations in only three states (Ohio, New York, and Massachusetts) and in Ontario (see map, below). All ALB infestations are under active eradication programs.

Locations of Asian longhorned beetle found since 1996.  Note:  The “active infestation” in the Greater Toronto Area is under an active eradication program with no recent detections.

The map below is the EDDMapS (Early Detection & Distribution Mapping System) Ontario distribution map for the Asian longhorned beetle as of May 2017.  To see the current EDDMapS distribution map, click on the map below.

There are restrictions on the movement of nursery stock, trees, logs, lumber, wood, wood chips and bark chips from trees identified as potential hosts of the ALB and firewood of all species in the regulated area (see map below).  It is also recommended that potential host trees (maple, birch, poplar, willow, elm, horse chestnut, sycamore, katsura, mountain ash, silk tree, hackberry, and goldenrain tree) should not  be planted within the regulated area until the ALB has been declared eradicated.

The Asian longhorned beetle regulated area

Map:   CFIA (2022) Click on the map to see the current regulated areas.

The following Canadian Food Inspection Agency (CFIA) plant protection policies relate to the ALB:

• D-11-05 – Phytosanitary Requirements for Non-Manufactured and Non-Propagative Wood Products to Prevent the Introduction from the Continental United States and Spread Within Canada of the Asian Long-horned Beetle, Anoplophora glabripennis (Motschulsky)
• D -11-01 – Phytosanitary Requirements for Plants for Planting and Fresh Branches to Prevent the Entry and Spread of  Anoplophora   spp.
• D -08-04 – Plant Protection Import Requirements for Plants and Plant Parts for Planting: Preventing the Entry and Spread of Regulated Plant Pests Associated with the Plants for Planting Pathway
• D-02-12 – Import requirements of non-manufactured wood and other non-propagative wood products, except solid wood packaging material, from all areas other than the continental United States
• D-01-12 – Phytosanitary Requirements for the Importation and Domestic Movement of Firewood
• D-98-08 – Entry Requirements for Wood Packaging Materials Produced in All Areas Other Than the Continental United States

In 2003, adult ALB were discovered in areas of Toronto and Vaughan.  The Canadian Food Inspection Agency (CFIA) acted quickly to address the infestation. Many susceptible trees around the infested areas were removed in hopes of stopping the spread of the beetle. As a result of early detection and quick response, the spread of ALB was prevented and no beetles were detected within Ontario between 2007 and 2013 ( OFAH/OMNR Invading Species Awareness Program, 2012 ). Unfortunately, a new infestation was detected in Mississauga in 2013.

Economic impacts

If the beetle spreads within Ontario and Canada, it can cause large reductions in wood supply and reduce the availability and quality of hardwood species to the forest industry. This would mean large revenue losses, as the hardwood forest industry is worth billions of dollars in wood products ( NRCan, 2014 ). For instance, Canada’s commercial hardwood forests produce $11 billion in wood products annually ( City of Toronto, 2015 ).

Also, as the beetle favours maple species, the damage and death it causes to these trees could greatly hurt the maple syrup industry, which is worth about $100 million each year in Canada ( NRCan, 2014 ;  City of Toronto, 2015 ).

Further, healthy forests support tourism and recreation, so damage to the natural environment could negatively impact these industries as well.

Ecological impacts

Since ALB has no effective predators in North America, it could cause great damage to native ecosystems if it is not contained. The hardwood species that the beetle attacks make up a large proportion of both natural and urban forest canopies across Canada. For example, in the City of Toronto, 42% of the street trees are preferred host species for the beetle and thus susceptible to its attack ( City of Toronto, 2015 ).

In natural deciduous forests, hardwood species such as maple serve as a foundation species in their environment. These trees hold a disproportionate control over the structure and function of the ecosystem, and their loss can cause widespread ecological damage. According to Ellison et al. (2005), the loss of a foundation species from a forest can result in a disruption of fundamental ecosystem processes, such as decomposition rates, nutrient cycling, carbon sequestration, energy flow, and impacts adjacent aquatic ecosystems in riparian areas. If ALB is not contained, it has the potential to devastate Canadian forests, especially when combined with the damage caused by emerald ash borer and other invasive forest pests. Many species at risk are found within Ontario’s hardwood forests, so their survival is also at risk with this potential decline of forest health ( City of Toronto, 2015 ).

Social impacts

Any widespread tree loss due to ALB has the potential to impact tourism and recreation values, due to the loss of aesthetic values that trees and forests possess.

After its first detection in 2003, the CFIA implemented an eradication program in 2004 to quickly respond to the threat of ALB. First, a quarantine zone of about 150km 2  was established around the infested trees where the beetle was discovered. This restricted the movement of wood and wood products out of this zone ( NRCan, 2014 ).

If you have found a suspect Asian longhorned beetle and/or its habitat, download and complete the following sampling protocols:

• Survey Guidelines for Asian Longhorned Beetle ( Anoplophora glabripennis ) – ENGLISH
• Lignes directrices pour les enquêtes sur le longicorne asiatique ( Anoplophora glabripennis ) – FRANÇAIS

When conducting surveys in accordance with this protocol, please contact the CFIA so all partner activities can be tracked as part of the national surveillance efforts.  Suspect finds should also be reported to the CFIA  [email protected].

Respond and Control

All infested trees found between 2003 and 2004 were removed and the wood was chipped. In addition, another 12,500 nearby trees that were considered susceptible to infestation were also cut down and chipped  ( NRCan, 2014 ) . This was done to prevent the further spread of the beetle. As trees were removed, full documentation of tree species, tree size, and signs and location of beetle attacks on each tree was carried out. This was done to develop a comprehensive understanding of the beetle’s behaviour in the Canadian context, to be used for future detection, training, and management  ( NRCan, 2014 ) .

An infestation of ALB was detected in Mississauga, Ontario in 2013. Using knowledge gathered in the first eradication of the insect, an eradication plan was quickly put in place. Monitoring efforts concluded in 2020 when the Canadian Food Inspection Agency declared ALB had been eradicated from Mississauga and Toronto.

Towards successful eradication of the Asian longhorned beetle: Early detection & rapid response 

When prevention is not possible, ‘early detection and rapid response’ is advocated as the best strategy to combat an introduced invasive species. As the name suggests, the success of this strategy depends on how early the species is detected, and how rapid and effective the response is. In some cases, when an invasive species is first detected in a forest or urban area, it has already spread much further than expected, and it may not be possible to eradicate. For example, when the emerald ash borer (EAB) was first detected in Windsor, Ontario in 2002, a quarantine barrier was established. The EAB attacks and kills >99% of ash trees, so an ‘ash-free zone’ was established where all potential host trees were removed in an attempt to stop EAB spread. Unfortunately, the EAB had already spread beyond the geographical range of this barrier long before the quarantine was established. Scientists now believe that EAB could have been present in Ontario up to 10 years prior to its initial detection. Even though the response was rapid, it was not effective at stopping the spread of the insect due to late detection.

In some cases, however, early detection and rapid response can be very effective and successful. This is best exemplified in the case of the Asian longhorned beetle (ALB). ALB was first detected in an industrial area north of Toronto in September 2003. ALB attacks and kills a wide range of native trees, including the symbolic maple tree. This one invasive species had the potential to impact tourism and social values, create losses in the billion dollar hardwood and maple syrup industries, and cause widespread and immeasurable impacts on native forests. Officials were already aware of the potential impacts of ALB on many of Canada’s native hardwood tree species, so were quick to make the decision to eradicate. Immediately, a 150 km 2 quarantine zone was established around the initial detection site to restrict the movement of wood materials. By the following spring, 531 infested or likely infested trees were removed and destroyed, in addition to 12,500 high-risk trees in the regulated area.

After tree removal, the Canadian Forest Service and multiple survey crews continued to monitor trees within the regulated area looking for any signs of ALB. From 2004-2007, an additional 40 infested trees were detected and removed each year, less than 15 infested trees were found in 2007, and no infested trees were found for the remainder of the surveys (NRCAN, 2011). Over five years passed with no detectable signs of ALB in the area, and it was officially declared to be eradicated in April 2013 by the CFIA – a great success!

Unfortunately, the celebrations were short-lived. In December 2013, an adult beetle was collected and positively identified as ALB in an industrial area of Mississauga, outside of the original regulated area. Again, early detection of the new infestation allowed for rapid response by dedicated researchers and technicians. The area surrounding the detection was extensively surveyed, and technicians were able to locate two Norway maples and three Manitoba maples that showed signs of infestation. One of the Manitoba maples was heavily infested with over 900 exit holes!

Photo: Pennsylvania Department of Conservation and Natural Resources bugwood.org. A tree showing damage from ALB 

Over the remainder of the winter, extensive tree removal took place in the new infested area. Tree removal efforts focused on four preferred tree hosts for ALB: maple, poplar, willow, and birch. Roughly 7,500 trees were removed in an 800 m buffer surrounding known infested sites. Each tree that was removed was carefully inspected by researchers for signs of ALB damage and then destroyed. Throughout this process, 25 more infested trees and 17 suspect trees were discovered. A new 46 km 2  regulated area was established surrounding the infestation (below), and again, it is prohibited to move any tree materials out of the area.

Photos: (1) Thomas B. Denholm, New Jersey Department of Agriculture, Bugwood.org  and (2) USDA Forest Service – Forest Operations Research , USDA Forest Service, Bugwood.org.

Following positive detection of Asian longhorned beetle, trees identified to be infested or high-risk are removed and destroyed by chipping.

This area continues to be very closely monitored for any new signs of ALB, and if discovered, any suspect trees will be promptly removed and destroyed. This is the quick action that is required to control an invasive pest outbreak, and prevent the species from becoming established on the Canadian landscape. The intentional removal of host trees from these regulated areas is a labour-intensive and expensive operation, however these impacts are only a small fraction of the potential economic, ecological, and social impacts created if the ALB were to permanently establish in Canada. With this continued effort, ALB can be successfully and permanently eradicated from Ontario.

You can help stop an Asian longhorned beetle invasion by:

• Never moving firewood, nursery stock, or other tree materials outside of regulated areas.
• Learn to identify the Asian longhorned beetle.
• Report any suspect ALB signs or symptoms – photos help! For more information on reporting an invasive species, click here!
• Spread the word! Tell your friends, family, and neighbours about the threat from ALB.
• Replant removed trees with a native non-host species. Contact your local forestry professionals for information on what to plant.

For more tips,  check out the Forest Invasives Canada Quick Tips Page !

Fact Sheets

Asian Longhorned Beetle Fact Sheet

Longicorne Asiatique

CFIA Fact Sheet

USGA Fact Sheet

New York State Fact Sheet

ALB Survey Protocol ENG

ALB Survey Protocol FR

Managing invasive populations of  asian longhorned beetle  and citrus  longhorned beetle : a worldwide perspective, asian longhorned beetle  (coleoptera: cerambycidae), an introduced pest of maple and other hardwood trees in north america and europe, asian longhorned beetle  anoplophora glabripennis (motschulsky): lessons learned and opportunities to improve the process of eradication and management, current research and knowledge gaps.

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Further Reading

The Invasive Species Centre aims to connect stakeholders. The following information below link to resources that have been created by external organizations.

Invading Species – Asian Long-horned Beetle Profile

Asian Long-horned Beetle: An unwanted invasive species

Natural Resources Canada – Asian Long-horned Beetle Profile 

Ontario Government – Asian Long-horned Beetle Profile 

Tree Canada – Asian Long-horned Beetle Profile

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High trunk truncation as a potential sustainable management option for asian longhorned beetle on salix babylonica.

Simple Summary

1. introduction, 2. materials and methods, 2.1. survey sites, 2.2. insects, 2.3. biological characteristics of alb, 2.3.1. spatial distributions of frass and emergence holes, 2.3.2. bark consumption by alb adults on branches with different diameters, 2.3.3. oviposition selection on branches with different diameters, 2.4. control effectiveness of high trunk truncation for alb in a controlled field experiment, 2.5. control effectiveness of high trunk truncation for alb in different areas, 2.6. statistical analysis, 3.1. biological characteristics of alb, 3.1.1. spatial distributions of frass and emergence holes, 3.1.2. bark consumption by alb adults on branches with different diameters, 3.1.3. oviposition selection on branches with different diameters, 3.2. control effectiveness of high trunk truncation for alb in a controlled field experiment, 3.3. control effectiveness of high trunk truncation for alb in different areas, 4. discussion, supplementary materials, author contributions, data availability statement, acknowledgments, conflicts of interest.

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Huang, C.; Wang, H.; Hai, X.; Wang, Z.; Lyu, F. High Trunk Truncation as a Potential Sustainable Management Option for Asian Longhorned Beetle on Salix babylonica . Insects 2024 , 15 , 278. https://doi.org/10.3390/insects15040278

Huang C, Wang H, Hai X, Wang Z, Lyu F. High Trunk Truncation as a Potential Sustainable Management Option for Asian Longhorned Beetle on Salix babylonica . Insects . 2024; 15(4):278. https://doi.org/10.3390/insects15040278

Huang, Chen, Hualing Wang, Xiaoxia Hai, Zhigang Wang, and Fei Lyu. 2024. "High Trunk Truncation as a Potential Sustainable Management Option for Asian Longhorned Beetle on Salix babylonica " Insects 15, no. 4: 278. https://doi.org/10.3390/insects15040278

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Asian Longhorned Beetles: What to Know

The Asian longhorned beetle is a type of longhorn beetle that poses a serious problem to hardwood forests. Although native to Asia, this invasive species has spread throughout parts of North America and Europe. To prevent further spread, many countries have established eradication programs to assess infected trees and destroy infestations of this insect. 

What Is the Asian Longhorned Beetle? 

Asian longhorned beetle facts: 

• Asian longhorned beetles likely came to North America and Europe through infested wood that was used to build shipping crates.
• Adult female beetles eat small grooves out of trees so that they can lay their eggs. Once eggs hatch, the larvae live in and eat the tree for up to two years. 
• It’s estimated that rampant infestation of the Asian longhorned beetle could kill over 30% of urban trees in the United States. 
• Trees and forests infested with Asian longhorned beetles have to be quarantined and systematically eradicated to get rid of the insects.

Asian longhorned beetle appearance. Asian longhorned beetles are named for their long antennae, which are usually about 1.3 to 1.5 times as long as their bodies. The antennae are black with white stripes. These beetles are fairly large and can measure up to 1.5 inches long. 

Adult Asian longhorned beetles are all black with white dots on their wings, earning them the nickname “starry sky beetle”. Adult beetles have six legs and can fly but typically only do so for short distances. 

Asian longhorned beetle life cycle. Adult female beetles lay eggs in tree bark. The eggs hatch after about 11 days. After hatching, larvae eat their way into the tree. Larvae can stay in this stage for one to two years. 

Once larvae are ready to mature, they create a hollow chamber in the tree where they can pupate. The pupa develops for two to three weeks before an adult beetle emerges and tunnels itself out of the tree. 

What does the Asian longhorned beetle eat? Asian longhorned beetles in the larval stage burrow through trees, eating the wood as they go. Adult beetles are herbivores and tend to eat twigs and leaves.

Types of Asian Long-Horned Beetles

Several species may be mistaken for Asian longhorned beetles. Citrus longhorned beetles, another invasive species, look the same as Asian longhorned beetles from a distance. If you look very closely, citrus longhorned beetles have bumps on their back that Asian longhorned beetles don’t have. 

Other species that look somewhat similar include:

• White-spotted sawyers, which are much smaller than Asian longhorned beetles but colored similarly
• Northern pine sawyers, which have grayish bodies rather than black
• Cottonwood borers, which have a light body with black spots (the opposite of the black body with light spots seen on Asian longhorned beetles)

Where Do Asian Long-Horned Beetles Live? 

Asian longhorned beetles native range. Asian longhorned beetles are natively found in East Asia, most commonly throughout China, Japan, and Korea.

Asian longhorned beetles invasive range. Asian longhorned beetles have been present in North America since at least 1996, when they were first spotted in New York and Chicago. It’s believed that the beetles came from Asia through untreated wooden shipping crates.

These beetles spread easily, and are now found in most of the northeastern United States and in parts of eastern Canada. They have also been found in almost a dozen countries in Europe, including Austria, Italy, France, and the United Kingdom.

Asian longhorned beetles habitat. These beetles spend most of their life as larvae eating their way into trees. As adults, Asian longhorned beetles typically stay around hardwood forests and other areas with large amounts of hardwood, such as warehouses that store wooden crates.

Signs You Have Asian Longhorned Beetles

Adult Asian longhorned beetles are hard to miss. Look for large, black beetles with white spots, particularly around any trees in the yard. 

Asian longhorned beetle damage. Adult Asian longhorned beetles are typically most active in the summer and fall. If you aren’t seeing any beetles but are worried about an infestation, you can look for certain signs of damage on trees, including:

• Round exit holes in the trunk that adult beetles have emerged from
• Frass , a byproduct of burrowing that looks like sawdust scattered around the branches and base of the tree trunk
• Dark, circular stains on the trunk where eggs have been laid
• Branch dieback , dead branches at the top of an otherwise healthy-looking tree, since infested trees tend to die from the top down

Why Do You Get Asian Longhorned Beetles? 

Unfortunately, there is not much you can do to prevent an infestation of Asian longhorned beetles. It can be difficult to tell if you have an infestation, as an infested tree may not die for 10 to 15 years, and larvae can infest a tree for a year or two before emerging as adult beetles. 

Health Risks of Asian Longhorned Beetles

Asian longhorned beetles are not directly dangerous to humans. These beetles will not bite, poke, or sting you. However, Asian longhorned beetles can have significant indirect health impacts. 

The wood-boring nature of these beetles kills trees from the inside. Dead trees pose a significant hazard if left standing and are likely to drop limbs that can injure pedestrians and damage vehicles or even topple over entirely during storms.

How to Get Rid of the Asian Longhorned Beetle

Asian longhorned beetles are an invasive species that are usually dealt with by governmental agencies like the United States Department of Agriculture. These agencies typically have eradication programs that will quarantine affected areas and take down infected trees at no cost.

Asian longhorned beetle treatment. If you’re worried about a beetle infestation:

• Contact your local eradication program before removing yard waste.
• Do not sell or move any wood, including firewood, from potentially infested trees.
• Use a compliant tree removal or landscaping company to dispose of any woody material.
• Allow eradication program officials to inspect and remove infested trees.

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