ECOLOGY AND MANAGEMENT OF FOREST INSECTS IN MICHIGAN

• Sponsoring Institution: National Institute of Food and Agriculture
• Project Status: COMPLETE
• Funding Source: MCINTIRE-STENNIS
• Reporting Frequency: Annual
• Accession No.: 0160464
• Project Start Date: Jul 1, 2008
• Project End Date: Jun 30, 2013
• Program Code: [(N/A)]- (N/A)

Recipient Organization
MICHIGAN STATE UNIV
(N/A)
EAST LANSING,MI 48824

Performing Department
Entomology

Non Technical Summary
While native insects play important roles in maintaining healthy forest ecosystems over the long-term, exotic forest insects are a different story. In the past 15 years, Michigan forests have been impacted by several alien forest insect pests. Beech scale and beech bark disease were identified in 2000 and in 2002, the emerald ash borer was discovered. Exotic forest insects have not evolved with their North American hosts, which are often much less resistant than congeners in the insects? native range. Exotic insects also typically arrive without the complement of natural enemies that helped to control populations in their native habitat. Information on ecology and impacts of these new pests, the factors that contribute to forest susceptibility or vulnerability, and interactions between these pests and native insects is needed to develop appropriate management strategies to reduce damage. More generally, we need to learn more about exotic forest insect pest invasions and to develop mitigation strategies to reduce the arrival and establishment of potentially damaging nonindigenous forest insects.

Animal Health Component

(N/A)

Research Effort Categories

Basic

(N/A)

Applied

(N/A)

Developmental

(N/A)

Classification

Knowledge Area (KA)	Subject of Investigation (SOI)	Field of Science (FOS)	Percent
211	0610	1130	100%

Knowledge Area
211 - Insects, Mites, and Other Arthropods Affecting Plants;

Subject Of Investigation
0610 - Conifer forests of the North;

Field Of Science
1130 - Entomology and acarology;

Keywords

invasive species

invasive forest pests

emerald ash borer

beech scale

beech bark disease

invasion ecology

forest insect management

Goals / Objectives
Project Objective 1: Increase our understanding of the ecology of native and nonindigenous forest insects including their interactions with host trees, natural enemies and potential competitors, the ecological roles that native insects play in forested ecosystems, and the effects of disturbance and management activities on forest insects. Project Objective 2: Acquire information on the impacts of native and nonindigenous forest insects, the factors that contribute to insect damage, and apply this knowledge towards the development of appropriate mitigation and management strategies. SPECIFC OBJECTIVES: Over the next five years, I anticipate that major areas of focus will include (1) ecology, impact and management of nonindigenous forest insects; (2) biology, ecology and management of emerald ash borer and (3) biology and spread of beech scale in Michigan. These studies were described in detail in Form A of the Research Project Outline. OUTPUTS: We expect to publish results in peer-reviewed scientific journals and to disseminate information via trade and extension publications, web sites and workshops. Results from this work will increase our understanding of forest insect invasion ecology, economic and ecological impacts of invasive forest insects and management options for newly discovered invasive forest pests. New information on emerald ash borer will be provided to regulatory officials, forest managers, arborists, Native American tribes, environmental groups and private landowners to assist in detection, regulatory activities and management of EAB. Results from studies on beech scale and beech bark disease will be provided to forest managers and landowners to assist in long-term planning efforts.

Project Methods
I will collaborate with other scientists to address ecology, impacts and management of nonindigenous forest insects. A list of nonindigenous forest insects in the US will be developed and used to assess patterns in invasion or establishment rates. Current and potential distribution of nonindigenous species will be mapped by combining information on the host range of the pests and tree distribution. Estimates of costs associated with exotic forest pests will be generated. A second group will assess the feasibility of eradication programs for nonindigenous forest pests. We will continue our efforts to reconstruct the temporal and spatial dynamics of ash mortality caused by emerald ash borer (EAB) across southeastern MI. This work relies on dendrochronological analyses of increment cores and cross-sections from ash trees growing across six counties. Data were imported into a GIS to spatially represent areas of EAB-induced tree mortality. Results will provide a unique description of the spatial and temporal characteristics of the EAB invasion. Results from this and other field studies will be used to model EAB population dynamics and ash mortality. An accurate model can be used to evaluate alternative management scenarios for any outlier site. We will continue to evaluate persistence and efficacy of insecticides used to protect landscape ash trees from EAB. We anticipate additional EAB projects will include evaluating suitability of western ash species for EAB adult feeding and larval development, continuing our work on EAB detection and survey methods and determining the biology, distribution and impact of the native Atanycolus sp. parasitoid on EAB populations. We also have a major study underway in central Michigan to assess how EAB density and distribution varies over time in a forested setting. We plan to continue our efforts to monitor and delineate the advancing and killing fronts of beech bark disease (BBD) in lower and upper Michigan. STo track spread of beech scale, we will return to pre-established sites near the advancing fronts in upper and lower Michigan. Beech trees in sites where beech scale was absent in the previous year will be examined to assess progress of the advancing front. Within each site, we will quantify overstory conditions (species composition, DBH, crown condition, canopy transparency) using fixed radius plots. Results will be imported into a GIS and maps indicating locations of the advancing front (stands with beech scale) will be developed and provided to state and national forest managers. We will also use these data to validate and refine the model we developed in 2007 to predict rate of spread of the advancing fronts in upper and lower Michigan.

Progress 07/01/08 to 06/30/13

Outputs
Target Audience: Forest and land managers, municipal foresters and arborists, personnel from regulatory agencies, Christmas tree growers, private property owners Changes/Problems: Nothing Reported What opportunities for training and professional development has the project provided? Results from our projects and new information on exotic forest pests of concern have been presented to foresters, arborists, regulatory officials and other target audiences at a workshops, conferences and meetings. How have the results been disseminated to communities of interest? Oral presentations, articles in magazines, websites, journal articles What do you plan to do during the next reporting period to accomplish the goals? Our work on the ecology and management of native and exotic forest insects will continue. Several projects related to EAB are ongoing. Results from our beech bark disease work will be submitted for publication. Additional funding will be sought to continue our studies.

Impacts
What was accomplished under these goals? Our projects addressed the ecology & management of exotic as well as native forest insect pests. Much of our work in this reporting period focused on emerald ash borer biology, ecology and management in forest and urban settings. We assessed EAB spread, determined the origin of the EAB infestation in North America and evaluated interactions between this insect and native ash host trees. We monitored parasitism of EAB by native and introduced species of natural enemies, and quantified woodpecker predation on EAB populations. We began a statewide survey with baited traps designed to identify native and non-native woodboring beetle species in hardwood & conifer forests, plant nurseries and sawmills. We completed a survey & have mapped the 2013 distribution of beech scale and beech bark disease across the state, quantified impacts on tree mortality and assessed changes in species composition of regeneration and down woody debris in affected stands. We published our results in scientific journals, websites and trade magazines and gave presentations to a wide range of forestry, arboriculture and regulatory personnel at local, state and national meetings

Publications

• Type: Other Status: Published Year Published: 2013 Citation: Wieferich, D., D. Hayes, D.G. McCullough. 2013. Evaluation of digital photography for quantifying beech scale (Cryptococcus fagisuga Lind.) (Hemiptera: Eriococcidae) density on American beech trees. Journal of Economic Entomology 106: 1324-1330. Tobin, P.C., J.M. Kean, D.M. Suckling, D.G. McCullough, D.A. Herms, L.D. Stringer. 2013. Determinants of successful eradication. Biological Invasions. In press. Mercader, R.J., D.G. McCullough and J.M. Bedford. 2013. A comparison of girdled ash detection trees and baited artificial traps for emerald ash borer (Agrilus planipennis Fairmaire) detection. Environmental Entomology 42: 1027-1039. Liebhold, A.M, D.G. McCullough, L.M. Blackburn, SJ. Frankel, B.Von Holle and J.E.Aukema. 2013. A highly aggregated geographical distribution of forest pest invasions in the USA. Diversity and Distributions 19: 1208-1216. Kovacs, K.F., R.G. Haight, R.J. Mercader, and D.G. McCullough. 2013. A bioeconomic analysis of an emerald ash borer invasion of an urban forest with multiple jurisdictions. Resource and Energy Economics. In press. Klooster, W.S., D.A. Herms,K.S. Knight, C.P. Herms, D.G. McCullough, A. Smith, K.J.K. Gandhi, and J. Cardina. 2013. Ash (Fraxinus spp.) mortality, regeneration, and seed bank dynamics in mixed hardwood forests following invasion by emerald ash borer (Agrilus planipennis). Biological Invasions. In press. Nagle, A.M., R. Usborne, A. Stone, D. McCullough and C. S. Sadof. 2013. Power hours: invasive species communication through collaborative webinars. Journal of Extension. In press. (Note: JoE is peer-reviewed). McCullough, D.G. and R. Usborne. 2003-Current. National emerald ash borer website; includes photos, information related to EAB and ash tree identification, research updates, control options for homeowners and landscapers, regulations, bulletins and many other pages. Available at www.emeraldashborer.info. McCullough, D.G., R. Usborne and A. Ziegler. 2010-Current. Developed, populated and maintain multi-agency SLAMEAB.info website, which provides information, interactive map viewers, trap and detection tree data, outreach materials and other features related to the SLAM (SL.ow A.sh M.ortality) Pilot Project in the Upper Peninsula. www.slameab.info. McCullough, D.G. 2013. Will we kiss our ash goodbye? Featured article; American Forests 118: 16-23. Wieferich, J.B. and D.G. McCullough. 2013. Impacts and spread of beech bark disease: another invasive pest clobbers Michigan forests. Newsletter of the Michigan Entomological Society.

Progress 01/01/12 to 12/31/12

Outputs
OUTPUTS: We continue to study emerald ash borer (EAB) biology, detection and management options. In a recent collaboration, we showed EAB, now present in 18 states and two provinces, is the most destructive and costly forest insect to invade North America. We used our simulation models to project EAB population growth, spread and ash mortality over time to evaluate the large scale SLAM (SLowing Ash Mortality) pilot project in Upper MI. On a local scale, we simulated effects of treating 10 to 50% of ash trees in a neighborhood with a highly effective insecticide. Treating 20% of trees protected 90% of the ash over 10 years at the lowest cumulative cost. We evaluated three trap designs and lures comprised of ash volatiles or an EAB contact pheromone for EAB detection in green ash forests. We continued to assess effects of systemic insecticide products on adult and larval EAB, and the ability of trees to recover from varying levels of EAB-caused damage when treated with emamectin benzoate. We are screening EAB resistance of ash species from several US regions, Europe and Asia. We determined native blue ash is as resistant to EAB as Asian ash species. Sara Tanis, a PhD student, assessed effects of plant growth regulators and fertilization on EAB resistance of multiple ash species. Stephen Burr completed his MS project to quantify EAB density, overstory composition and regeneration in green ash forests at three stages of EAB invasion. James Wieferich, an M.S. student, is quantifying impacts of beech bark disease and documening the expansion of the advancing front of BBD in lower and Upper MI. I collaborated with an international group of scientists to populate a global database of arthropod eradication programs and to evaluate factors that contribute to successful eradication. We continued to host and maintain the national EAB website. Results from our projects are distributed through this and other websites, presentations and posters at state, regional and national meetings, publications in scientific journals, conference proceedings, and other formats. PARTICIPANTS: Collaborators include Therese Poland, Andrew Liebhold, Patrick Tobin, Steven Katovich, Robert Haight (USDA Forest Service), Phil Lewis (USDA APHIS), John Kean, Maxwell Suckling (New Zealand), Robert Heyd, Roger Mech and Ron Murray (MI DNR), John Bedford, Michael Philip, Robin Rosenbaum (MDARD), Bert Cregg (MSU), Dan Herms (Ohio State Univ); Visiting scientis: Rodrigo Mecader (Washburn University); Post-doctoral RA: Elizabeth Graham (MSU); Graduate students: Sara Tanis, Stephen Burr, James Wieferich; Technicians: Andrea Anulewicz, Jacob Bournay, Andrew Tluczek; Undergraduates: Kyle Redilla, Molly Robinett, Russell Kibalt, EAB Outreach: Robin Usborne, Amy Kennedy. TARGET AUDIENCES: Audiences include forest managers, forest health specialists, consulting foresters, arborists, municipal and urban foresters, regulatory personnel from state, provincial or federal agencies, conservation organizations, extension educators, Christmas tree growers, undergraduate students PROJECT MODIFICATIONS: Nothing significant to report during this reporting period.

Impacts
In cooperation with international collaborators, we identified combinations of control tactics and the influence of Allee effects on potential eradication of non-native insects. Results from our research have been incorporated into the ongoing SLAM (SLow Ash Mortality)pilot project in the Upper Peninsula and similar projects are underway in IN, PA and NY. Results from our insecticide projects continue to be provided to private landowners, forest managers, municipal arborists, city foresters and regulatory officials in many states as well as Ontario and Quebec. Our studies showed EAB populations remain in southeast MI, the original core of the EAB infestation, despite nearly complete ash mortality. Blue ash, a native species, survived the EAB invasion in southeast MI and appears to be at least as resistant to EAB as Asian ash species. We continue to document the spread of the advancing front of beech bark disease. Updated maps were distributed to state forest health professionals in November and are currently used to establish firewood and harvest regulations for state forest land. We determined the cerambycid (longhorned beetle) communities vary considerably between ground and canopy levels in forests. We showed the influence of spring weather on the gypsy moth fungal pathogen Entomophaga maimaiga in a large scale field study.

Publications

• Graham, E.E., T.M. Poland, D.G. McCullough and J. G. Millar. 2012. A comparison of trap type and height for capturing cerambycid beetles (Coleoptera). Journal of Economic Entomology 105(3): 837-846.
• McCullough, D.G. and R.J. Mercader. 2012. SLAM in an urban forest: evaluation of potential strategies to SLow Ash Mortality caused by emerald ash borer (Agrilus planipennis). International Journal of Pest Management 58: 9-23.
• Mercader, R.J., N.W. Siegert and D.G. McCullough. 2012. Estimating the influence of population density and dispersal behavior on the ability to detect and monitor Agrilus planipennis (Coleoptera: Buprestidae) populations. Journal of Economic Entomology 105: 272-281.
• Siegert, N.W., D.G. McCullough, M.M. Wheeler and A.E. Hajek. 2012. Evaluation of potential versus realized primary infection of gypsy moth (Lepidoptera: Lymantriidae) by Entomophaga maimaiga (Zygomycetes: Entomophthorales). Environmental Entomology 41(5):1115-1124.
• Suckling, D.M., D.G. McCullough, D.A. Herms and P.C. Tobin. 2012. Combining tactics to exploit Allee effects for eradication of alien insect populations. Forum article; Environmental Entomology 105: 1-13.
• Tanis, S.R. and D.G. McCullough. 2012. Differential persistence of blue ash and white ash following emerald ash borer invasion. Canadian Journal of Forest Research 42: 1542-1550.
• Tanis, S. R., B. M. Cregg, D. Mota-Sanchez, D. G. McCullough, and T. M. Poland. 2012. Spatial and temporal distribution of trunk-injected 14C-imidacloprid in Fraxinus trees. Pest Management Science 68: 529-536.
• Bournay J., D.G. McCullough, N.J. Gooch, A. Anulewicz and P. Lewis. 2012. Lethal trap trees: a potential tool for managing emerald ash borer. In: Proceedings of the Emerald Ash Borer Research and Technology Development Meeting. October 14-16, 2011. Wooster, Ohio. USDA Forest Service.
• Burr, S.J. and D.G. McCullough. 2012. The Future of green ash behind, within, and ahead of the advancing front of emerald ash borer. In: K. McManus and K.W. Gottschalk, eds. Proceedings, USDA Interagency Research Forum on Invasive Species 2012. January, Annapolis. USDA Forest Service, Northern Research Station.
• Burr, S.J. and D.G. McCullough. 2012. The rise and fall of emerald ash borer populations and the ash overstory at three stages of the invasion wave. In: Proceedings of the Emerald Ash Borer Research and Technology Development Meeting. October 14-16, 2011. Wooster, Ohio. USDA Forest Service.
• Herms, D.A., J. Cardina, K.S. Knight, K.J.K. Gandhi, W. Klooster, K. Rice, C.P. Herms, D. Hartzler, A. Smith, and D.G. McCullough. 2012. Direct and indirect ecological impacts of emerald ash borer and ash mortality. In: Proceedings of the Emerald Ash Borer Research and Technology Development Meeting. October 14-16, 2011. Wooster, Ohio. USDA Forest Service.
• Mercader, R.J. and D.G. McCullough. 2012. SLAM in an urban forest: can we optimize strategies for protecting landscape ash trees from emerald ash borer Proceedings of the Emerald Ash Borer Research and Technology Development Meeting. October 14-16, 2011. Wooster, Ohio. USDA Forest Service.
• Poland T.M. and D.G. McCullough. 2012. Emerald ash borer attraction to artificial traps: influence of trap design and color at different population densities. In: Proceedings of the Emerald Ash Borer Research and Technology Development Meeting. October 14-16, 2011. Wooster, Ohio. USDA Forest Service.
• Tanis, S.R. and D.G. McCullough. 2012. Five Fraxinus species and one Agrilus beetle: adult emerald ash borer survival, host preference and larval density. In: K. McManus and K.W. Gottschalk, eds. Proceedings, USDA Interagency Research Forum on Invasive Species 2012. January, Annapolis. USDA Forest Service, Northern Research Station. Tanis, S.R. and D.G. McCullough. 2012. Good to be blue: evidence that blue ash may persist in North American forests. Proceedings of the Emerald Ash Borer Research and Technology Development Meeting. October 14-16, 2011. Wooster, Ohio. USDA Forest Service.
• Tluczek, A.R. and D.G. McCullough. 2012. Are impacts of native enemies on emerald ash borer larvae increasing Proceedings of the Emerald Ash Borer Research and Technology Development Meeting. October 14-16, 2011. Wooster, Ohio. USDA Forest Service.
• Wieferich, J.B. and D.G. McCullough. 2012. Comparison of beech bark disease distribution and impacts in Michigan from 2002 to 2011. In: K. McManus and K.W. Gottschalk, eds. Proceedings, USDA Interagency Research Forum on Invasive Species 2012. January, Annapolis. USDA Forest Service, Northern Research Station.
• Anulewicz, A.C. and D.G. McCullough. 2012. Development of emerald ash borer (Agrilus planipennis) in novel ash (Fraxinus spp.) hosts. In: Proceedings of the Emerald Ash Borer Research and Technology Development Meeting. October 14-16, 2011. Wooster, Ohio. USDA Forest Service.

Progress 01/01/11 to 12/31/11

Outputs
OUTPUTS: We continue to study emerald ash borer (EAB) biology, detection and management options. This beetle has killed tens of millions of trees in 15 states and two provinces. It is the most destructive and costly forest insect to ever invade North America. Along with my post-doc, Rodrigo Mercader, we developed, refined and used simulation models to project EAB population growth, spread and ash mortality over time. These models were applied to evaluate the SLAM (SLowing Ash Mortality)approach in a large scale pilot project in the Upper Peninsula. We also used our models to evaluate how EAB populations are likely to build and spread across the US, the economic costs and the influence of satellite populations on spread and costs over the next ten years. On a local scale, we simulated effects of using a highly effective insecticide in a residential neighborhood. Optimal treatment regimes and cumulative costs of different treatment protocols were identified. We continued to evaluate trap designs and lures for EAB detection in field sites with varying EAB densities. We continue to evaluate one, two and three-year control of trunk-injected systemic insecticides. We are also studying effects of insecticide products on adult and larval EAB, and the ability of trees to recover from varying levels of EAB-caused damage when treated with emamectin benzoate. We continued to record presence and parasitism rates of a native Atanycolus sp. wasp that has begun to attack EAB larvae. We are studying EAB host preference and resistance of ash species from several US regions, Europe and Asia. We recently determined that native blue ash is as resistant to EAB as Asian ash species. Sara Tanis, a PhD student, is continuing to assess effects of plant growth regulators and other treatments on EAB resistance of multiple ash species. Stephen Burr is continuing his MS project to quantify EAB density, overstory composition and regeneration in green ash forests at three stages of the EAB invasion wave (core, crest, cusp). He completed his fieldwork sthis summer. James Wieferich is working on his MS research to assess impacts of beech bark disease and to document the expansion of the advancing front of BBD in lower and Upper MI. I continued to collaborate with a multi-disciplinary group of scientists to identify and quantify the economic impacts of non-native and invasive forest insects. I am also working with another group to evaluate factors that contribute to successful eradication efforts for non-native insects around the world. We continued to develop and maintain the national EAB website, which is hosted at MSU. Results from our projects are distributed through the EAB website, presentations and posters at state, regional and national meetings, publications in scientific journals, conference proceedings, and other formats. I also provided information to a variety of media ranging from Time Magazine to local news and radio programs. PARTICIPANTS: Collaborators include Therese Poland, Andrew Liebhold, Steven Katovich, Robert Haight and Leah Bauer (USDA Forest Service), Kent Kovacs (University of Minnesota), Phil Lewis (USDA APHIS), Juliann Aukema (The Nature Conservancy), Betsy Von Holle (Univ of Central FL), Robert Heyd, Roger Mech and Ron Murray (MI DNR), John Bedford and Robin Rosenbaum (MI Dept of Agriculture), Andrew Storer (MI Tech Univ), Bert Cregg, Dan Hayes and Amos Ziegler (MSU), Dan Herms (Ohio State Univ). Post-doctoral research associates: Rodrigo Mecader; Graduate students: Sara Tanis, Stephen Burr, James Wieferich; Technicians: Andrea Anulewicz, jacob Bournay, Andrew Tluczek; Undergraduate student workers: Elliott Berlin, Kyle Redilla, Molly Robinett, Russell Kibalt, Heather Surface; EAB Outreach and Communications: Robin Usborne, Amy Kennedy, Roger Lintemuth. Opportunities for professional development and training were provided to undergraduate and graduate students, technicians and post-docs associated with my project. We also provided hands on training for survey crews working on the SLAM project and for regulatory and natural resource professionals from other states. TARGET AUDIENCES: Forest health specialists, forest managers and natural resource managers in the Lake States, upper Midwest and northeastern US; Regulatory personnel from state and federal agencies; Conservation organizations; Arborists, municipal foresters, landscapers and property owners; Extension educators; Christmas tree growers PROJECT MODIFICATIONS: Nothing significant to report during this reporting period.

Impacts
Results from our research, including modeling work, have been incorporated into the ongoing SLAM (SLow Ash Mortality)pilot project in the Upper Peninsula, which brought $2.2 million in ARRA funding and 36 jobs to MI for 2010-2011. Similar projects are now underway in IN and PA. Results from our insecticide projects continue to be provided to private landowners, forest managers, municipal arborists, city foresters and regulatory officials in many states where EAB is established. Our studies also showed EAB populations remain in southeast MI despite nearly complete ash mortality. The double-decker EAB trap, which we designed, was used operationally in at least 3 states in 2011 and led to new county records in PA. We identified a native ash species that appears to be at least as resistant to EAB as Asian ash species. We continue to document the spread of the advancing front of beech bark disease. Updated maps were distributed to state forest health professionals in November and are used to establish firewood and harvest regulations for state forest land. An notable paper quantifying the economic costs of non-native forest insects, currently in press, showed that EAB is the most costly invasive forest pest. We assessed five economic sectors and found that homeowners and municipalities bear most of these costs, while federal expenditures and lost timber account for a relatively small proportion of costs.

Publications

• Poland, T.M., D.G. McCullough and A.C. Anulewicz. 2011. Evaluation of an artificial trap for Agrilus planipennis (Coleoptera: Buprestidae) incorporating olfactory and visual cues. Journal of Economic Entomology 104:517-531.
• Tluczek, A.R., D.G., McCullough and T.M. Poland. 2011. Influence of host stress on emerald ash borer (Agrilus planipennis Fairmaire) (Coleoptera: Buprestidae) adult density, development, and distribution in Fraxinus pennsylvanica trees. Environmental Entomology 40:357-366.
• Wieferich, D., D. Hayes, D.G. McCullough and N. Schwalm. 2011. Distribution of beech scale (Cryptococcus fagisuga Lind) in Michigan from 2004 to 2009. Northern Journal of Applied Forestry. In press.
• McCullough, D.G. and R. Usborne. 2011. International multi-agency emerald ash borer website. www.emeraldashborer.info.
• McCullough, D.G. and A. Ziegler. 2011. SLowing Ash Mortality (SLAM) webiste. SLAMEAB.info.
• McCullough, D.G., N.W. Siegert, T.M. Poland, S.J. Pierce and S.Z. Ahn. 2011. Effects of trap type, placement and ash distribution on emerald ash borer captures in a low density site. Environmental Entomology 40:1239-1252.
• McCullough, D.G., T.M. Poland, A.C. Anulewicz, P. Lewis and D. Cappaert. 2011. Evaluation of Agrilus planipennis control provided by emamectin benzoate and two neonicotinoid insecticides, one & two seasons after treatment. Journal of Economic Entomology 104(5): 1599-1612.
• Mercader, R.J., N.W. Siegert, A.M. Liebhold and D.G. McCullough. 2011. Estimating the effectiveness of three potential management options to slow the spread of emerald ash borer populations in localized outlier sites. Canadian Journal of Forest Research 41:254-264.
• Mercader, R.J., N.W. Siegert, A.M. Liebhold and D.G. McCullough. 2011. Simulating the influence of the spatial distribution of host trees on the spread of the emerald ash borer, Agrilus planipennis, in recently colonized sites. Population Biology 53:271-285.
• Hahn, J., D.A. Herms and D.G. McCullough. 2011. Frequently asked questions regarding potential side effects of systemic insecticides used to control emerald ash borer. Multi-state extension bulletin. 4 pages. Available at www.emeraldashborer.info.
• Aukema, J.E., Leung B, Kovacs K., Chivers, C., Britton K.O., Englin J., Frankel S.J., Haight R.G., Holmes T.P., A. Liebhold, McCullough, D.G. and Von Holle, B. 2011. Economic impacts of non-native forest insects in the continental United States. PLoSONE 6(9). In press.
• Kovacs, K., R.J. Mercader, R. Haight, N. Siegert; D.G. McCullough, A. Liebhold. 2011. The influence of satellite populations of emerald ash borer on projected economic damage in U.S. communities, 2010-2020. Environmental Management. 92:2170-2181.

Progress 01/01/10 to 12/31/10

Outputs
OUTPUTS: We continue to study emerald ash borer (EAB) biology, detection and management options. This beetle has killed tens of millions of trees in at least 15 states and is the most destructive forest insect to ever invade North America. We evaluated effectiveness of four trap designs for EAB detection in six field sites with varying EAB densities. We continued to evaluate trunk-injected systemic insecticides to assess multi-year control, effects of products on adult and larval EAB, and the ability of trees to recover from varying levels of EAB-caused damage when treated with emamectin benzoate. We determined that trees injected with emamectin benzoate then girdled 3 weeks later serve as lethal trap trees; egg-laying females are attracted but larvae do not survive. We continued to record presence and parasitism rates of a native Atanycolus wasp that has begun to attack EAB larvae. Chenin Limback completed her MS and determined that while EAB is more attracted to green ash than white ash, girdled white ash are as attractive as green ash. Enhancing vigor through fertilization did not increase ash tree resistance. We continued to study EAB host preference and resistance of ash species from several US regions, Europe and Asia. Sara Tanis, a PhD student, is continuing to assess effects of plant growth regulators and other treatments on host resistance to EAB. Stephen Burr is continuing his study to quantify EAB density, overstory composition and regeneration in green ash forests at three stages of the EAB invasion wave (core, crest, cusp). A post-doc, Rodrigo Mercader, is continuing to develop simulation models to project EAB population growth, spread and ash mortality over time. We also continue to study the progression and impact of beech bark disease (BBD) in Michigan. James Wieferich began his MS research to assess impacts of beech bark disease and to document the expansion of the advancing front of BBD in lower and Upper MI. I continued to collaborate with a multi-disciplinary group of scientists to identify the historical accumulation of non-native forest pests, the proportion that cause damage and the economic impacts of invasive forest insects. I am also working with another group to evaluate ecological and economic factors that contribute to successful eradication efforts for non-native forest insects. We continued to develop and maintain the national EAB web site, which is hosted at MSU. PARTICIPANTS: Collaborators include Therese Poland, Andrew Liebhold, Steven Katovich, Robert Haight, Leah Bauer (USDA Forest Service), Phil Lewis and John Molongoski (USDA APHIS), Kent Kovacs (Univ of MN), Juliann Aukema and Frank Lowenstein (The Nature Conservancy), Betsy Von Holle (Univ of Central FL), Robert Heyd and Roger Mech (MI DNR), John Bedford and Ken Rauscher (MI Dept of Agriculture), Andrew Storer (MI Tech Univ), Bert Cregg (MSU), Dan Herms (Ohio State Univ), Dan Hayes (MSU). Post-doctoral research associates: Rodrigo Mecader and Nathan Siegert; Graduate students: Sara Tanis, Stephen Burr, James Wieferich, Chenin Limback; Technicians: Andrea Anulewicz, jacob Bournay, Nicholas Gooch, Emily Pastula, Andrew Tluczek; Undergraduate student workers: Jacob Bournay, James Wieferich, Paul Nelson, Rachel Posavetz; EAB Outreach and Communications: Robin Usborne, Amy Kennedy, Roger Lintemuth. TARGET AUDIENCES: Forest health specialists, forest managers and natural resource managers in the Lake States, upper Midwest and northeastern US; regulatory personnel from state and federal agencies; conservation organizations; arborists, landscapers, property owners, extension educators, Christmas tree growers PROJECT MODIFICATIONS: Nothing significant to report during this reporting period.

Impacts
Data from several studies have been incorporated into the ongoing SLAM (SLow Ash Mortality)pilot project in the Upper Peninsula, which brought $2.2 million in ARRA funding and 36 jobs to MI for 2010-2011. Similar projects are now underway in IN, MO and potentially in NY. Our simulation models, currently in development, will be the primary means to evaluate the effectiveness of SLAM strategies. Results from our insecticide projects continue to be provided to private landowners, municipal arborists, foresters and regulatory officials in many states where EAB is established. The double-decker EAB trap, which we designed, was used operationally in at least 3 states in 2010. We identified a native ash species that appears to be at least as resistant to EAB as Asian ash species. We continue to document the spread of the advancing front of beech bark disease. Maps were distributed to state forest health professionals and used to establish firewood and harvest regulations for state forest land. An article describing the numbers, accumulation rates and proportion of damaging forest insects by feeding guild was featured in Bioscience. In a related study, our group quantified the economic costs of non-native forest insects on five economic sectors in the US.

Publications

• Mercader, R.J., N.W. Siegert, A.M. Liebhold and D.G. McCullough. 2010. Simulating the influence of the spatial distribution of host trees on the spread of the emerald ash borer, Agrilus planipennis, in recently colonized sites. Population Biology. In press.
• Cappaert, D. and D.G. McCullough. 2010 (2009). Phenology of Atanycolus cappaerti (Hymenoptera: Braconidae), a native parasitoid of emerald ash borer, Agrilus planipennis (Coleoptera: Buprestidae). Great Lakes Entomologist 41:141-154.
• Kovacs, K.F., R.G. Haight, D.G. McCullough, R.J. Mercader and N.W. Siegert. 2010. Cost of potential emerald ash borer damage in U.S. communities, 2009-2019. Ecological Economics 69:569-578.
• Liebhold, A.M., T.T. Work, D.G. McCullough and J.F. Cavey. 2010. Response to Carey (2010). American Entomologist 56:164-165.
• Siegert, N.W., D.G. McCullough, D.W. Williams, I. Fraser and T.M. Poland. 2010. Dispersal of Agrilus planipennis (Coleoptera: Buprestidae) from discrete epicenters in two outlier sites. Environmental Entomology 39(2): 253-265.
• Herms, D.A. and D.G. McCullough. 2010. Pesticides and Insect Eradication. In: The Encyclopedia of Invasive Introduced Species. D. Simberloff and M. Rejmanek, eds. University of California Press. Berkeley, CA.
• Liebhold, A.M. and D.G. McCullough. 2010. Forest Insects. In: The Encyclopedia of Invasive Introduced Species. D. Simberloff and M. Rejmanek, eds. University of California Press. Berkeley, CA.
• Anulewicz, A.C., D.G. McCullough, T.M. Poland, P. Lewis and J. Molongoski. 2010. Evaluation of multi-year application of neonicotinoid insecticides for EAB control. p. 66-67, In: Proceedings of the Emerald Ash Borer Research and Technology Development Meeting. October 20-21, 2009. Pittsburg, PA. USDA Forest Service, Forest Health Technology Enterprise Team, FHTET-2010-01. 136 p.
• Anulewicz, A.C., D.G. McCullough, S. Tanis, C. Limback, R. Hofstetter, A. Mayfield and S. Munson. 2010. Coast to coast ash mortality Potential susceptibility of selected western and southern ash species. p. 49-50, In: Proceedings of the Emerald Ash Borer Research and Technology Development Meeting. October 20-21, 2009. Pittsburg, PA. USDA Forest Service, Forest Health Technology Enterprise Team, FHTET-2010-01. 136 p.
• McCullough, D.G., A.C. Anulewicz, T.M. Poland, P. Lewis and J. Molongoski. 2010. Evaluation of emamectin benzoate and neonicotinoid insecticides: two year control of EAB P. 68-70, In: Proceedings of the Emerald Ash Borer Research and Technology Development Meeting. October 20-21, 2009. Pittsburg, PA. USDA Forest Service, Forest Health Technology Enterprise Team, FHTET-2010-01. 136 p. .
• Mercader, R.J., N.W. Siegert, A.M. Liebhold and D.G. McCullough. 2010. Comparing potential management options to slow the spread of EAB in localized sites. p. 39-40, In: Proceedings of the Emerald Ash Borer Research and Technology Development Meeting. October 20-21, 2009. Pittsburg, PA. USDA Forest Service, Forest Health Technology Enterprise Team, FHTET-2010-01. 136 p.
• Poland, T.M., D.G. McCullough and G. Grant. 2010. Evaluation of traps and attractants for EAB: effects of trap color, placement, host volatiles, release rate and release devices on male and female responses. p. 119-122, In: Proceedings of the Emerald Ash Borer Research and Technology Development Meeting. October 20-21, 2009. Pittsburg, PA. USDA Forest Service, Forest Health Technology Enterprise Team, FHTET-2010-01. 136 p.
• Siegert, N.W., D.G. McCullough, R.J. Mercader and S.J. Pierce. 2010. Dispersal of emerald ash borer at newly colonized sites. p. 22, In: Proceedings of the Emerald Ash Borer Research & Technology Meeting. October 20-21, 2009. Pittsburgh, PA. USDA Forest Service, Forest Health Technology Enterprise Team, In: Proceedings of the Emerald Ash Borer Research and Technology Development Meeting. October 20-21, 2009. Pittsburg, PA. USDA Forest Service, Forest Health Technology Enterprise Team, FHTET-2010-01. 136 p.
• Siegert, N.W., D.G. McCullough, T. M. Poland, R.L. Heyd, S.J. Pierce and Y. Zue. 2010. Effect of clusters of girdled trees on the spread of emerald ash borer in low density infestations. p. 23, In: Proceedings of the Emerald Ash Borer Research & Technology Meeting. October 20-21, 2009. Pittsburgh, PA. USDA Forest Service, Forest Health Technology Enterprise Team, In: Proceedings of the Emerald Ash Borer Research and Technology Development Meeting. October 20-21, 2009. Pittsburg, PA. USDA Forest Service, Forest Health Technology Enterprise Team, FHTET-2010-01. 136 p.
• Tanis, S.R., B.M. Cregg, D. Mota-Sanchez, D.G. McCullough and T.M. Poland. 2010. Sectored flow and reservoirs: a synthesis of 14C-imidacloprid trunk-injection research. p. 78-79, In: Proceedings of the Emerald Ash Borer Research & Technology Meeting. October 20-21, 2009. Pittsburgh, PA. USDA Forest Service, Forest Health Technology Enterprise Team, In: Proceedings of the Emerald Ash Borer Research and Technology Development Meeting. October 20-21, 2009. Pittsburg, PA. USDA Forest Service, Forest Health Technology Enterprise Team, FHTET-2010-01. 136 p.
• Tluczek, A.R., D. Cappaert and D.G. McCullough. 2010. Life cycle of Atanycolus sp. nr. hicoriae, a newly described native parasitoid of emerald ash borer. p. 91-92, In: Proceedings of the Emerald Ash Borer Research and Technology Development Meeting. October 20-21, 2009. Pittsburg, PA. USDA Forest Service, Forest Health Technology Enterprise Team, FHTET-2010-01. 136 p.
• Tluczek, A.R., D.G. McCullough, T.M. Poland and A.C. Anulewicz. 2010. The fast and the furious: influence of host vigor on development and mortality of emerald ash borer larvae. P. 49-50, In: Proceedings of the Emerald Ash Borer Research and Technology Development Meeting. October 20-21, 2009. Pittsburg, PA. USDA Forest Service, Forest Health Technology Enterprise Team, FHTET-2010-01. 136 p
• McCullough, D.G. 2010. Ask the expert: balsam twig aphid control. Great Lakes Christmas Tree Journal 5(3):4-5.
• Poland, T. and D.G. McCullough. 2010. SLAM: A Multi-Agency pilot project to SL.ow A.sh M.ortality caused by emerald ash borer in outlier sites. Newsletter of the Michigan Entomological Society 55(1-2):4-8
• Herms, D.A., W. Klooster,, K.S. Knight,K.J.K. Gandhi, C.P. Herms, A. Smith, D.G. McCullough, J. Cardina. 2010. Ash regeneration in the wake of emerald ash borer: will it restore ash or sustain the outbreak P. 17-18, In: In: Proceedings of the Emerald Ash Borer Research and Technology Development Meeting. October 20-21, 2009. Pittsburg, PA. USDA Forest Service, Forest Health Technology Enterprise Team, FHTET-2010-01. 136 p.
• Limbeck, C.K., D.G. McCullough, Y. Chen, T.M. Poland and B. Cregg. 2010. Host vigor and emerald ash borer larvae on white ash and green ash. p. 44-46, In: Proceedings of the Emerald Ash Borer Research and Technology Development Meeting. October 20-21, 2009. Pittsburg, PA. USDA Forest Service, Forest Health Technology Enterprise Team, FHTET-2010-01. 136 p.
• McCullough, D.G. 2010. Invasive forest insects: trends and impacts. Abstract, In: Proceedings of the USDA Interagency Research Forum on Gypsy Moth & Other Invasive Species. January 12-15, 2010. Annapolis, MD. Published by USDA Forest Service, Northern Research Station, Newtown Square, PA. Gen. Tech. Rep. In press.
• McCullough, D.G., N.W. Siegert, T.M. Poland, S.J. Pierce and S.Z. Ahn. 2010. Detection of a low density emerald ash borer infestation: trap design, placement and ash distribution. p. 116-117, In: Proceedings of the Emerald Ash Borer Research and Technology Development Meeting. October 20-21, 2009. Pittsburg, PA. USDA Forest Service, Forest Health Technology Enterprise Team, FHTET-2010-01. 136 p.

Progress 01/01/09 to 12/31/09

Outputs
OUTPUTS: We continued research on emerald ash borer (EAB) biology, detection and management in 2009. This insect has become the most destructive forest insect to ever invade North America and at least 15 native ash species are threatened. Our studies included comparisons of different trap designs and lures for EAB detection. We continued to evaluate systemic insecticides for EAB control. A two-year study comparing neonicotinoid and emamectin benzoate products was completed. Another project addressed efficacy of lethal trap trees that were injected with emamectin benzoate then girdled 3 wks later. Three additional studies to assess long term effectiveness of insecticide products are underway. Funding was acquired for a major project we developed to slow the onset and progression of ash mortality in localized EAB outlier sites in Upper Michigan. This pilot project, referred to as SLAM (SLowing Ash Mortality), now involves multiple state and federal agencies and institutions. We continued studies to determine the biology, life cycle and parasitism rate of Atanycolus cappaerti, a newly described native parasitoid wasp that we discovered attacking EAB larvae. Andrew Tluzek completed his MS project that addressed EAB larval develpment in relation to host plant stress. Chenin Limback continued her study to assess EAB preference and performance on green ash and white ash and how both ash species respond to stress. Sara Tanis began her fieldwork to identify anatomical differences in leaf and woody tissues among four North American and 1 Asian ash species. She will be evaluating EAB adult and larval performance on the five ash species. Nate Siegert, a post-doc, and I continued a large scale project to assess EAB population dynamics in forests and how girdled trees can be used to redirect spread or reduce EAB density. Another post-doc, Rodrigo Mercader, developed models to predict how EAB populations will grow and spread over time and how management options affect spread. Stephen Burr began his MS program to quantify EAB populations and tree regeneration behind and ahead of areas where EAB is at peak densities. Dan Wieferich continued to evaluate the advance of beech scale and beech bark disease across the state and how beech scale populations change over time. Nancy Schwalm completed her MS thesis on the rate of beech scale spread. I collaborated with a multi-disciplinary team to assess the historical accumulation of non-native forest pests and the economic damage associated with EAB and with all high impact invaders. We continued development and maintenance of the national EAB web site, housed at MSU, along with other outreach and extension efforts. Previous studies related to gypsy moth and effects of weather and climate on the entomopathogen Entomophaga maimaiga, an important biological control of gypsy moth, were completed and published. PARTICIPANTS: Collaborators include Therese Poland, Andrew Liebhold, Noel Schneeberger, Robert Haight and Leah Bauer (USDA Forest Service), Phil Lewis and John Molongoski (USDA APHIS), Kent Kovacs (Univ Nevada-Reno), Juliann Aukema (The Nature Conservancy), Betsy VonHolle (Univ of Central FL), Bob Heyd (MI DNR), John Bedford and Ken Rauscher (MI Dept. of Agriculture), Andrew Storer (MI Tech Univ), Bert Cregg (MSU Dept. of HORT), David Mota-Sanchez and Robert Hollingworth (MSU Dept. of ENT), Frank Telewski (MSU Dept. of BOT) and Dan Hayes (MSU Dept. of F&W). Post-doctoral research assistants: Nathan W. Siegert and Rodrigo Mercader; Graduate students: Sara Tanis, Chenin Limbach, Andrew Tluczek, Dan Wieferich and Stephen Burr; Technicians: Nicholas Gooch, Andrea Anulewicz, Emily Pastulae; Undergraduate assistants: Tara Dell, James Wieferich and Jake Bournay; EAB Outreach and Communications: Robin Usborne, Roger Lintemuth, Amy Kennedy TARGET AUDIENCES: Forest health specialists, forest managers and natural resource managers in the Lake States, the upper Midwest, the northeast and the Plains states regions of the US; Regulatory personnel from state and federal agencies; Conservation organizations (e.g. The Nature Conservancy); arborists, landscapers, property owners, extension educators. PROJECT MODIFICATIONS: Nothing significant to report during this reporting period.

Impacts
Data collected from many of our previous studies formed the basis for the EAB SLAM (Slowing Ash Mortality) pilot project, underway in Upper MI. This project brought more than $2.2 million in federal funding (mostly ARRA) into Michigan, primarily for jobs in the Upper Peninsula. State agencies in at least 2 other states have initiated similar projects on a smaller scale. Our research on EAB population dynamics, models of EAB population growth and spread, girdled trap trees and insecticides have all been incorporated into the national SLAM effort. Results from our EAB insecticide studies were presented to EPA and used to support registration requests for two products. A non-invasive trunk spray application with dinotefuran was approved for full registration. Full registration of emamectin benzoate is pending but expected to be approved by spring 2010. Results from studies with emamectin benzoate, which provided 2 years of nearly 100% control, were shared with collaborators in other states to support special registrations for emamectin benzoate in at least 10 states. I co-authored a multi-state bulletin about insecticides for EAB control and secured funding for printing. It has been widely distributed across the Midwest and eastern US. We collaborated with forest economists to project EAB spread and economic costs likely to be sustained by 2020. Municipalities can expect to spend at least $10 billion treating or replacing ash trees on public property and if suburban areas are included, the costs nearly double. Our research on the Atanycolus parasitoid wasp we found attacking EAB led to the identification and description of a new species. The Latin species name cappaerti was derived from David Cappaert, my former technician who initially recognized the importance of this wasp. We mapped the current distribution of the advancing front of beech bark disease and distributed maps to state and federal forest managers. Our maps are used to develop harvest plans and minimize transport of infested logs or firewood during high risk periods.

Publications

• McCullough, D.G., T. M. Poland, D. Cappaert and A.C. Anulewicz. 2009. Emerald ash borer (Agrilus planipennis) attraction to ash trees stressed by girdling, herbicide and wounding. Canadian Journal of Forest Research 39:1331-1345.
• McCullough, D.G., T. M. Poland, A.C. Anulewicz and D. Cappaert. 2009. Emerald ash borer (Agrilus planipennis Fairmaire) (Coleoptera: Buprestidae) attraction to stressed or baited ash (Fraxinus spp.) trees. Environmental Entomology 38:1668-1679.
• Mercader, R., N.W. Siegert, D.G. McCullough and A.M. Liebhold. 2009. Dispersal of the emerald ash borer, Agrilus planipennis, in newly colonized sites. Agricultural and Forest Entomology 11:421-424.
• Mota-Sanchez, D., B. M. Cregg, D. G. McCullough, T. M. Poland, and R. M. Hollingworth. 2009. Distribution of trunk-injected 14C imidacloprid in ash trees and effects on emerald ash borer (Coleoptera: Buprestidae) adults. Crop Protection 28: 655-661.
• Siegert, N.W., D.G. McCullough, A.E. Hajek and J.A. Andresen. 2009. Effect of microclimatic conditions on primary transmission of the gypsy moth fungal pathogen Entomophaga maimaiga (Zygomycetes: Entomophthorales) in Michigan. The Great Lakes Entomologist 41:111-128.
• Siegert, N.W., D.G. McCullough, R.C. Venette, A.E. Hajek and J.A. Andresen. 2009. Assessing the climatic potential for epizootics of the gypsy moth pathogen Entomophaga maimaiga in the North Central United States. Canadian Journal of Forest Research 39:1958-1970.
• Siegert, N.W., D.G. McCullough, J. Bedford, S.A. Katovich and R.J. Mercader. 2009. Can we slow the spread of isolated emerald ash borer populations A brief overview of the SLAM concept. Abstract, In: Proceedings of the North Central Forest Pest Workshop. September 28 - October 1, 2009. Winnipeg, Manitoba, Canada. Siegert, N.W., D.G. McCullough, T.M. Poland, R.L. Heyd, S.J. Pierce and Y. Xue. 2009. Manipulating the spread of emerald ash borer using clusters of girdled trees in forested areas. Abstract, In: Proceedings of the North Central Forest Pest Workshop. September 28 - October 1, 2009. Winnipeg, Manitoba, Canada.
• Siegert, N.W., D.G. McCullough, R.C. Venette, A.E. Hajek and J.A. Andresen. 2009. Assessing the climatic potential for epizootics of the gypsy moth fungal pathogen Entomophaga maimaiga in the North Central United States. Abstract, In: Proceedings of the North Central Forest Pest Workshop. September 28 - October 1, 2009. Winnipeg, Manitoba, Canada.
• Storer, A.J., J.M. Marshall, M. Philip, D.G. McCullough and N.W. Siegert. 2009. The history of emerald ash borer discoveries in the Upper Peninsula of Michigan from 2005-2007. Abstract, In: Proceedings of the USDA Interagency Research Forum on Gypsy Moth & Other Invasive Species. January 9-13, 2008. Annapolis, MD. Published by USDA Forest Service, Northern Research Station (Gen. Tech. Rep. NRS-P-36), Newtown Square, PA.
• McCullough, D.G., N.W. Siegert and J. Bedford. 2009. SL.owing A.sh M.ortality: a potential strategy to SLAM emerald ash borer in outlier sites. Abstract, In: Proceedings of the USDA Interagency Research Forum on Gypsy Moth & Other Invasive Species. January 9-13, 2008. Annapolis, MD. Published by USDA Forest Service, Northern Research Station (Gen. Tech. Rep. NRS-P-36), Newtown Square, PA.
• Herms, D.A., D.G. McCullough, D.R. Smitley, C.S. Sadof, R.C. Williamson and P.L. Nixon 2009. Insecticide options for protecting ash trees from emerald ash borer. National IPM Center, Illinois. 12 p. Available for download on the national EAB.info web site.
• McCullough, D.G. and T.M. Poland. 2009. Using double-decker traps to detect emerald ash borer. Developed and distributed as a pdf on the national EAB.info web site and to forest management personnel in multiple states (MD, MN, PA, WI, WV). 10 p.

Progress 01/01/08 to 12/31/08

Outputs
OUTPUTS: Major research projects in 2008 focused on biology, detection & control of emerald ash borer (EAB), an invasive pest that threatens forest and urban forest ash trees across North America. Andrew Tluczek's M.S. research compared survival, density & development rate of EAB larvae on healthy and girdled ash trees. Results showed that overall EAB density increased by roughly ten-fold in one year, larval density & development rates were greater on girdled trees, & woodpecker predation was the most important mortality factor affecting larvae. Chenin Limback examined adult EAB feeding behavior on ash seedlings treated by girdling, fertilization, reduced nitrogen or left as controls as part of her M.S. research. Results suggested adult beetles compensate for low nutrient levels by increasing consumption. Chenin quantified photosynthetic capacity, foliar nutrient levels & EAB larval density & development on green and white ash trees in a plantation we established in 2007. Trees were treated by girdling, fertilization or left as controls. EAB larvae were more abundant on green than white ash and on girdled trees of both species. Nathan Siegert, a post-doc, continued to apply dendrochronological analysis to reconstruct the progression of ash mortality from 1998-2003 across southeast Michigan. We also established nine areas, each 40 acres in size, in central MI. Within each area, 50 x 50 m grid cells were established & all ash trees in each cell were tallied by diam class. EAB density was sampled across all areas and found to be at barely detectable densities. Studies including a low-density trapping experiment and an effort to re-direct EAB dispersal with girdled sink trees were implemented. Results were applied to develop a strategy for SLAM (Slowing Ash Mortality), a national pilot project initiated in 2008 in the Upper Peninsula in cooperation with several state & federal agencies. Rodrigo Mercader, a post-doc, began using data acquired from many of our previous EAB studies to develop an empirically based, spatially explicit model of EAB population growth and expansion. The model will be applied to evaluate success of the SLAM approach. We continued to evaluate insecticide products in multiple sites. Results showed that emamectin benzoate applied as a trunk injection provided two years of nearly 100% EAB control. We evaluated trap design & lures for EAB detection. Our multi-component, double-decker trap baited with a leaf blend lure was more effective than traps currently used by federal agencies. Biology, host range & phenology of a native parasitoid attacking EAB in east MI were evaluated. Dan Wieferich continued to work on his M.S. project to validate a model of beech scale spread, assess population dynamics of beech scale and quantify impacts of beech bark disease. Sara Tanis started her Ph.D. project, which will address differences among ash species in resistance to EAB. Nancy Schwalm continued to revise her M.S. thesis, while working full-time. Number and volume of jack pine trees and woody debris were re-measured in long-term plots in upper MI to assess impacts of jack pine budworm. PARTICIPANTS: Individual participating in projects: Nathan W. Siegert, Post-doctoral research associate; Rodrigo Mercader, Post-doctoral research associate; Andrew Tluczek, M.S. graduate student; Chenin Limbach, M.S. graduate student; Dan Wieferich, M.S. graduate student; Nancy Schwalm, M.S. graduate student; Sara Tanis, Ph.D. graduate student; Research technicians: Andrea Anulewicz, David Cappaert, Jacob Baker; Nicholas Gooch; Undergraduate students: Tara Dell, Ben Schmidt, James Wieferich, Erin Burkett, Kaeli Chambers, Joanna Bloese, Jacob Bournay. MSU faculty: Dan Hayes (F&W), Bert Cregg (Hort), Frank Telewski (Bot); MI Dept Natural Resources: Bob Heyd; MI Dept. of Agriculture: Field & supervisory personnel; USDA Forest Service: Therese Poland, Leah Bauer, Noel Schneeberger; USDA APHIS: Phil Lewis, John Molongoski TARGET AUDIENCES: TARGET AUDIENCES: forest health specialists, forest managers, natural resource managers in MI, WI, MN (the Lake States region), the Plains States (SD, ND, NE, KS), the northeastern US (ME, VT, CT, MA); regulatory personnel from MI & other states & from federal agencies (USDA APHIS), foresters, natural resource managers, landowners, arborists, landscapers, extension educators. PROJECT MODIFICATIONS: No major changes.

Impacts
Results from our EAB studies continue to be incorporated into operational EAB management programs in MI & other states affected by this tree-killing pest. The SLAM pilot study initiated in Upper MI utilizes results from our studies on EAB population dynamics, girdled trees for detection & management, & insecticide control. Other states have expressed interest in implementing SLAM to manage their EAB outlier sites. The emamectin benzoate insecticide we began working with in 2007 was granted a special registration in 9 states in 2008, largely because of the data we generated. Many, if not most, landscapers & arborists in MI & surrounding states began using this product in 2008. Full EPA registration is expected for the product in 2009. Dendrochronological analyses & population modeling are providing a means to predict how newly established EAB populations will grow & spread over time. We are collaborating with other scientists to predict distribution & economic impacts of EAB over the next ten years. We documented a native parasitoid, Atanycolus hicoriae, attacking EAB larvae in multiple sites in MI & 1 site in OH. The basic life cycle, phenology & host range were evaluated in 2008. Data & maps of beech scale spread were provided to state & federal forest health personnel. Maps are currently being used by state resource managers to develop harvest and mitigation plans.

Publications

• Anulewicz, A.C., D.G. McCullough, T.M. Poland, D. Cappaert. 2008. The 06 trap trees in 07. In: Mastro, V. &R. Reardon, compilers. Proc. of the Emerald Ash Borer Research and Technology Development Meeting. October 2007. Pittsburg, PA. USDA Forest Service, Forest Health Technology Enterprise Team, FHTET-2008-07. p. 71-71.
• Cappaert, D. and D.G. McCullough. 2008. The anticipated host switch: a new braconid parasitoid in Michigan. In: Mastro, V. and R. Reardon, compilers. Proceedings of the Emerald Ash Borer Research and Technology Development Meeting. October 2007. Pittsburg, PA. USDA Forest Service, Forest Health Technology Enterprise Team, FHTET-2008-07. p. 51-55.
• Cappaert, D., D.G. McCullough, T.M. Poland, P. Lewis and J. Molongoski. 2008. Imidacloprid concentration effects on adult EAB: a greenhouse study. In: Mastro, V. and R. Reardon, compilers. Proceedings of the Emerald Ash Borer Research and Technology Development Meeting. October 2007. Pittsburg, PA. USDA Forest Service, Forest Health Technology Enterprise Team, FHTET-2008-07. p. 43-44.
• Herms, D.A., K.J.K. Gandhi, J. Cardina, R.P. Long, K.S. Knight, A. Smith and D.G. McCullough. 2008. Impacts of emerald ash borer-induced gap formation on forest communities. In: Mastro, V. and R. Reardon, compilers. Proceedings of the Emerald Ash Borer Research and Technology Development Meeting. October 2007. Pittsburg, PA. USDA Forest Service, Forest Health Technology Enterprise Team, FHTET-2008-07. p. 10.
• McCullough, D.G. 2008. Putting the pieces together: can we solve the EAB management puzzle In: Proceedings of the 19th USDA Interagency Research Forum on Invasive Species. USDA Forest Service. Distributed as CD.
• McCullough, D.G. and J. ODonnell. 2008. Zimmerman pine moth. Great Lakes Christmas Tree Journal 3(1):17-19.
• McCullough, D.G. and N. Schneeberger. 2008. Pest Alert: Emerald Ash Borer, Revision III. Publication No. NA-PR-02-04. USDA Forest Service, State and Private Forestry, Northeastern Area. 2 pp.
• McCullough, D.G., T. M. Poland, D. Cappaert. 2008. Emerald ash borer (Agrilus planipennis) attraction to ash trees stressed by girdling, herbicide or wounding. Canadian Journal of Forest Research. Submitted.
• McCullough, D.G., D.A. Cappaert, T.M. Poland, A. Anulewicz, P. Lewis and J. Molongoski. 2008. Evaluation of non-invasive trunk sprays and trunk-injected emamectin benzoate. In: Mastro, V. and R. Reardon, compilers. Proceedings of the Emerald Ash Borer Research and Technology Development Meeting. October 2007. Pittsburg, PA. USDA Forest Service, Forest Health Technology Enterprise Team, FHTET-2008-07 p. 40-42.
• McCullough, D.G., T.M. Poland, A. Anulewicz and D.A. Cappaert. 2008. Double-deckers and towers: EAB traps in 2007. In: Mastro, V. and R. Reardon, compilers. Proceedings of the Emerald Ash Borer Research and Technology Development Meeting. October 2007. Pittsburg, PA. USDA Forest Service, Forest Health Technology Enterprise Team, FHTET-2008-07. p. 73-76.
• Poland, T.M., D.G. McCullough and L.S. Bauer. 2008. Developing tools to detect and manage emerald ash borer: an invasive pest of ash in urban and natural forests in North America. Abstract (Keynote presentation) In Proceedings, Entomological Society of Canada national meeting, November 2008, Edmonton, Alberta. In press.
• Siegert, N.W., A. M. Liebhold and D.G. McCullough. 2008. Modeling the spatial and temporal dynamics of isolated emerald ash borer populations. In: Mastro, V. and R. Reardon, compilers. Proceedings of the Emerald Ash Borer Research and Technology Development Meeting. October 2007. Pittsburg, PA. USDA Forest Service, Forest Health Technology Enterprise Team, FHTET-2008-07. p. 6-7
• Siegert, N.W., D.G. McCullough, A.M. Liebhold and F.W. Telewski. 2008. Dendrochronological reconstruction of the establishment and spread of emerald ash borer. In: Mastro, V. and R. Reardon, compilers. Proceedings of the Emerald Ash Borer Research and Technology Development Meeting. October 2007. Pittsburg, PA. USDA Forest Service, Forest Health Technology Enterprise Team, FHTET-2008-07. p. 4-5.
• Tanis, S.R., B.M. Cregg, D. Mota-Sanchez, D.G. McCullough, T.M. Poland, R.M. Hollingworth. 2008. Distribution of trunk-injected 14C imidacloprid in ash trees: variation between spring &fall injections. In: Mastro, V. and R. Reardon, compilers. Proc. of the Emerald Ash Borer Research and Technology Development Meeting. October 2007. Pittsburg, PA. USDA Forest Service, FHTET-2008-07. p.45-47. Tluczek, A.R., D.G. McCullough, T.M. Poland and A.C. Anulewicz. 2008. Effects of host stress on EAB development: what makes a good home In: Mastro, V. and R. Reardon, compilers. Proceedings of the Emerald Ash Borer Research and Technology Development Meeting. October 2007. Pittsburg, PA. USDA Forest Service, Forest Health Technology Enterprise Team, FHTET-2008-07. p. 32-33.

Progress 01/01/07 to 12/31/07

Outputs
OUTPUTS: Major research efforts in 2007 focused on biology, detection & control of emerald ash borer (EAB), an invasive pest that threatens forest and urban forest ash trees across North America. Andrew Tluczek completed the second year of his M.S. research that addresses effects of tree vigor on EAB larval survival & development. Results show that larval density and development rates are much greater on girdled trees than on healthy trees. We planted 450 ash trees (1-3 inches diam) in a block design that incorporates four North American and two Asian ash species. Trees will be used to evaluate effects of ash species and tree condition on EAB host selection, feeding and development. Chenin Limbach, who began her M.S. program in fall, will study EAB performance on green ash and white ash trees treated with fertilizer, girdled or left as controls. Nathan Siegert, a post-doc, used dendrochronological analysis to reconstruct the progression of ash mortality from 1997-2003 across a 5800 square mile area of southeast Michigan. Models to estimate ash phloem area & potential EAB production were developed & published. Results of studies on chipping, grinding & heat treatment for ash wood infested with EAB were completed & published. EAB population and rate of spread models are in development. Insecticide products and trap designs for EAB detection were evaluated in multiple sites. We initiated a large scale project in two northern MI forests to evaluate effectiveness of girdled sinks and other options for slowing population growth and spread of EAB in forested site. Dan Wieferich began his M.S. research in fall to validate a model of beech scale spread and to quantify impacts of beech bark disease. Nancy Schwalm, also studying beech bark disease, is revising her M.S. thesis. Number and volume of jack pine trees and woody debris were re-measured in long-term plots in upper MI to assess impacts of jack pine budworm. PARTICIPANTS: Nathan W. Siegert, Post-doctoral research associate; Andrew Tluczek, M.S. graduate student; Chenin Limbach, M.S. graduate student; Dan Wieferich, M.S. graduate student; Nancy Schwalm, M.S. graduate student; Research technicians: Andrea Anulewicz, David Cappaert, Jacob Baker; Undergraduate students: Tara Dell, Ben Schmidt, James Wieferich, Erin Burkett, Kaeli Chambers; MSU faculty: Dan Hayes (F&W), Bert Cregg (Hort), Frank Telewski (Bot); MI Dept Natural Resources: Bob Heyd; MI Dept. of Agriculture: Field & supervisory personnel; US Forest Service: Therese Poland, Leah Bauer, Noel Schneeberger; USDA APHIS: Phil Lewis, John Molongoski TARGET AUDIENCES: forest health specialists in MI, Lake States region, northeastern US; regulatory personnel from MI, other states, federal agencies (APHIS), foresters, natural resource managers, landowners, arborists, landscapers, extension educators

Impacts
Results from our EAB research continue to be incorporated into operational programs for EAB underway in states affected by this tree-killing pest. Our research on trap trees, trap design & lures continues to serve as the foundation for national EAB detection & monitoring programs. Dendrochronological analysis identified the actual area in SE MI where EAB first became established, the rate & distance at which satellite populations occurred & the rate of EAB spread. Our research resulted in a special registration in MI & OH for an efficient, inexpensive non-invasive application of neo-nicotinoid insecticide products. Efficacy of this application is similar to widely used methods but avoids wounding trees or affecting soil. Results from 2007 studies identified a highly effective insecticide product, emamectin benzoate, that controlled > 99% of EAB. Special registration of this product will be requested by MI & other states in 2008. We discovered a previously undocumented parasitoid of EAB larvae established in at least 2 sites. Species identification by systematic experts is pending. Data, maps & predictions of beech scale spread were provided to state & federal forest health personnel and used to develop harvest and mitigation plans.

Publications

• McCullough, D.G., D. Cappaert, T.M. Poland, P. Lewis and J. Molongoski. 2007. Evaluation of neo-nicotinoid insecticides applied as non-invasive trunk sprays. Pp. 52-54, In: Mastro, V., R. Reardon and G. Parra, compilers. Proceedings of the Emerald Ash Borer Research and Technology Development Meeting. October 31-November 1, 2006. Cincinnati, Ohio. USDA Forest Service, Forest Health Technology Enterprise Team, FHTET publ. 2007-04.
• McCullough, D.G., N.W. Siegert, D. Cappaert, T.M. Poland and R. McDonald. 2007. Sinks, bark and Garlon: applied studies for emerald ash borer management. Pp. 92-95, In: Mastro, V., R. Reardon and G. Parra, compilers. Proceedings of the Emerald Ash Borer Research and Technology Development Meeting. October 31-November 1, 2006. Cincinnati, Ohio. USDA Forest Service, Forest Health Technology Enterprise Team, FHTET publ. 2007-04.
• Anulewicz, A.C., D.G. McCullough, T.M. Poland and D.L. Cappaert. 2007. Attraction of emerald ash borer to trap trees: can methyl jasmonate or Manuka oil compete with girdling? Pp. 83-84, In: Mastro, V., R. Reardon and G. Parra, compilers. Proceedings of the Emerald Ash Borer Research and Technology Development Meeting. October 31-November 1, 2006. Cincinnati, Ohio. USDA Forest Service, Forest Health Technology Enterprise Team, FHTET publ. 2007-04.
• Cappaert, D., D.G. McCullough and T.M. Poland. 2007. Effects of trunk injection on emerald ash borer density and ash survival: a four-year study. Pp. 48-50, In: Mastro, V., R. Reardon and G. Parra, compilers. Proceedings of the Emerald Ash Borer Research and Technology Development Meeting. October 31-November 1, 2006. Cincinnati, Ohio. USDA Forest Service, Forest Health Technology Enterprise Team, FHTET publ. 2007-04.
• Anulewicz, A.C., D.G. McCullough and D.L. Miller. 2008. Host range studies of the emerald ash borer (Agrilus planipennis) (Coleoptera: Buprestidae): no-choice bioassays. Great Lakes Entomologist. In press.
• Anulewicz, A.C., D.G. McCullough, D.L. Cappaert and T.M. Poland. 2008. Host range of the emerald ash borer (Agrilus planipennis Fairmaire) (Coleoptera: Buprestidae) in North America: results of multiple-choice field experiments. Environmental Entomology. In press.
• Lindell, C., D.G. McCullough, D. Cappaert, N.M. Apostolou and M.B. Roth. 2008. Factors influencing woodpecker predation on emerald ash borer. American Midland Naturalist. In press.
• Anulewicz, A.C., D.G. McCullough and D.L. Cappaert. 2007. Emerald ash borer (Agrilus planipennis) density and canopy dieback in three North American ash species. Arboriculture and Urban Forestry 33:338-349.
• McCullough, D.G. and N.W. Siegert. 2007. Estimating potential emerald ash borer (Agrilus planipennis Fairmaire) populations using ash inventory data. Journal of Economic Entomology 100:1577-1586.
• McCullough, D.G., T. M. Poland, D. Cappaert, E.L. Clark, I. Fraser, V. Mastro, S. Smith and C. Pell. 2007. Effects of chipping, grinding and heat on survival of emerald ash borer (Agrilus planipennis Fairmaire) (Coleoptera:Buprestidae) in chips. Journal of Economic Entomology 100(4):1304-1315.
• McCullough, D.G. 2007. Rising from the ashes: what might we see after emerald ash borer? p. 20-23, In: F.P. Hain, R.N Coulson, K.D. Klepzig and J. Rhea, eds., Proceedings of the North American Forest Insect Work Conference. May 22-26, Asheville, North Carolina. North Carolina State University, distributed as CD.
• McCullough, D.G. and N.W. Siegert. 2007. Estimating emerald ash borer density at local, landscape or regional scales. p. 90-92, In: Mastro, V., R. Reardon and G. Parra, compilers. Proceedings of the Emerald Ash Borer Research and Technology Development Meeting. October 31-November 1, 2006. Cincinnati, Ohio. USDA Forest Service, Forest Health Technology Enterprise Team, FHTET publ. 2007-04.
• Poland, T.M. and D.G. McCullough. 2007. Evaluation of a multicomponent trap for emerald ash borer incorporating color, silhouette, height, texture and ash leaf and bark volatiles. pp. 74-76, In: Mastro, V., R. Reardon and G. Parra, compilers. Proceedings of the Emerald Ash Borer Research and Technology Development Meeting. October 31-November 1, 2006. Cincinnati, Ohio. USDA Forest Service, Forest Health Technology Enterprise Team, FHTET publ. 2007-04.
• Siegert, N.W., D.G. McCullough, A.M. Liebhold and F. Telewski. 2007. Resurrected from the ashes: a historical reconstruction of emerald ash borer dynamics through dendrochronological analyses. pp. 18-19, In: Mastro, V., R. Reardon and G. Parra, compilers. Proceedings of the Emerald Ash Borer Research and Technology Development Meeting. October 31-November 1, 2006. Cincinnati, Ohio. USDA Forest Service, Forest Health Technology Enterprise Team, FHTET publ. 2007-04.
• Siegert, N.W., D.G. McCullough, A. Liebhold and A. Tluczek. 2007. Two years under the bark: towards understanding multiple-year development of emerald ash borer larvae. P. 20, In: Mastro, V., R. Reardon and G. Parra, compilers. Proceedings of the Emerald Ash Borer Research and Technology Development Meeting. October 31-November 1, 2006. Cincinnati, Ohio. USDA Forest Service, Forest Health Technology Enterprise Team, FHTET publ. 2007-04.
• Mota-Sanchez, D. B.M. Cregg, , D.G. McCullough, T.M. Poland, Tanis, S.R., R. Isaacs and R. Hollingworth. 2007. Mortality, feeding and behavior of male and female emerald ash borer (Agrilus planipennis) adults in response to ingestion of imidacloprid. Pp. 38-41, In: Mastro, V., R. Reardon and G. Parra, compilers. Proceedings of the Emerald Ash Borer Research and Technology Development Meeting. October 31-November 1, 2006. Cincinnati, Ohio. USDA Forest Service, Forest Health Technology Enterprise Team, FHTET publ.2007-04.
• Tanis, S.R., B.M. Cregg, D. Mota-Sanchez, D.G. McCullough, T.M. Poland and R. Hollingworth. 2007. Distribution of trunk-injected 14C imidacloprid in Fraxinus trees: a test of the sectored-flow hypothesis. Pp. 34-38, In: Mastro, V., R. Reardon and G. Parra, compilers. Proceedings of the Emerald Ash Borer Research and Technology Development Meeting. October 31-November 1, 2006. Cincinnati, Ohio. USDA Forest Service, Forest Health Technology Enterprise Team, FHTET publ. 2007-04.
• McCullough, D.G. 2007. Hug your ash now, while you still can. Forest Landowners magazine. Forest Landowner , July/August volume: 30-32.
• McCullough, D.G., S. Evans, R. Usborne. 2007. Debarking ash trees for emerald ash borer detection. DVD of multi-state workshops held for survey crews from state and federal agencies, tribal government and universities involved with emerald ash borer detection and survey. 1 hour long. 1000 copies distributed to date.

Progress 01/01/06 to 12/31/06

Outputs
Major research efforts in 2006 focused on emerald ash borer (EAB)m an exotic pest that threatens the ash resource across the US. Andrew Tluczek completed his first year of research for his M.S. study of how tree vigor affects larval development. Manuscripts describing research on EAB host range and preferences are in preparation. We published a paper describing potential impacts of EAB and problems associated with invasive forest pests in urban settings. Research continues to refine methods for trap tree surveys, currently the primary EAB detection tool, and to develop attractive lures and traps. Nathan Siegert continues his post-doctoral studies to evaluate EAB population dynamics and rate of spread using dendrochronological data and model development is in progress. We identified the location of the initial EAB outbreak in North America. Nancy Schwalm completed field work as part of her M.S. project to identify the spatial extent and density of beech bark disease and developed a model to estimate rate of spread of the Advancing Front. We published a paper summarizing numbers, origins and commodities associated with exotic plant pest interception data at US ports and border crossings from 1984-2000. Another paper describes exotic plant pests associated with travelers baggage and potential re-introductions of economically important pests. We collected carabid beetles in three age classes of jack pine forests. Species identification is in progress to assess difference in carabid beetle diversity in stands regenerating after fire or harvest.

Impacts
Results from our emerald ash borer (EAB) studies continue to be directly incorporated into the operational EAB programs in all states affected by this tree-killing exotic pest. Our research on trap trees, beetle dispersal and extended larval development have been the basis for nearly all efforts to detect, monitor and contain EAB. More than 60 previously unknown EAB outlier populations were discovered in 2006 using trap tree and survey methods we developed. Results from our dendrochronological work have identified the area where ash trees were first killed by EAB and have defined the time when EAB likely arrived in the US. Results from our 2006 research on non-invasive application of systemic neo-nicotinoid insecticides can provide arborists with a simple, inexpensive method that avoids tree wounds or soil or water contamination. Our studies of the 17-year history of exotic plant pest interception records is being used by national environmental organizations, resource management agencies and regulatory officials to identify high risk pathways and commodities associated with exotic plant pest invasion. Data and maps resulting from our studies on beech beech bark disease were provided to state and federal forest managers in MI to assist them in developing harvest and management plans. Our work on ground beetles in jack pine stands will help resource managers compare the impacts of intensive harvesting and wildfire on biological diversity.

Publications

• Siegert, N.W., McCullough, D.G. and Andresen, J. 2006. Using degree day accumulation maps for improved timing of insect pest management. Published on MSU IPM web site at http://www.ipm.msu.edu/landscapeipm/gddland.htm and http://www.ipm.msu.edu/christmastreeipm/gddtree.htm.
• McCullough, D.G., Poland, T.M. and Cappaert, D.L. 2006. Attraction of emerald ash borer to trap trees: effects of stress agents and trap type. p. 61-62, In: Mastro, V., R. Reardon and G. Parra, compilers. Proceedings of the Emerald Ash Borer Research and Technology Development Meeting. September 26-27, 2005. Pittsburg, Pennsylvania. USDA Forest Service, Forest Health Technology Enterprise Team, FHTET-2005-16.
• McCullough, D.G., Cappaert, D.L., Poland, T.M., Lewis, P. and Molongoski, J. 2006. Long-term (3 year) evaluation of trunk injections for emerald ash borer control in landscape ash trees. p. 31-33, In: Mastro, V., R. Reardon and G. Parra, compilers. Proceedings of the Emerald Ash Borer Research and Technology Development Meeting. September 26-27, 2005. Pittsburg, Pennsylvania. USDA Forest Service, Forest Health Technology Enterprise Team, FHTET-2005-16.
• Anulewicz, A., McCulloug, D.G. and Cappaert, D.L. 2006. Emerald ash borer host range and preference studies. p. 15-16, In: Mastro, V., R. Reardon and G. Parra, compilers. Proceedings of the Emerald Ash Borer Research and Technology Development Meeting. September 26-27, 2005. Pittsburg, Pennsylvania. USDA Forest Service, Forest Health Technology Enterprise Team, FHTET-2005-16.
• Cappaert, D.L., McCullough, D.G., Poland, T.M., Lewis P. and Molongoski, J. 2006. Non-invasive neo-nicotinoids: treatments for ash logs and trees. p. 34-36, In: Mastro, V., R. Reardon and G. Parra, compilers. Proceedings of the Emerald Ash Borer Research and Technology Development Meeting. September 26-27, 2005. Pittsburg, Pennsylvania. USDA Forest Service, Forest Health Technology Enterprise Team, FHTET-2005-16.
• McCullough, D.G., Work, T.T., Cavey, J.F., Liebhold, A.T. and Marshall, D. 2006. Interceptions of nonindigenous plant pests at U.S. ports of entry and border crossings over a 17 year period. Biological Invasions 8:611-630.
• Poland, T.M. and McCullough, D.G. 2006. Emerald ash borer: invasion of the urban forest and the threat to North Americas ash resource. Journal of Forestry 104(3):118-124.
• Liebhold, A.M., Work, T.T., McCullough, D.G. and Cavey, J.F. 2006. Airline baggage as a pathway for alien insect species entering the United States. American Entomologist 52:48-54.
• Siegert, N.W., McCullough, D.G., Liebhold, A.M. and Telewski, F. 2006. Spread and dispersal of emerald ash borer: a coupled map lattice model approach. p. 11. In: Mastro, V., R. Reardon and G. Parra, compilers. Proceedings of the Emerald Ash Borer Research and Technology Development Meeting. September 26-27, 2005. Pittsburg, Pennsylvania. USDA Forest Service, Forest Health Technology Enterprise Team, FHTET-2005-16.
• Cregg, B.M., Mota-Sanchez, D., McCullough, D.G., Poland T. and Hollingworth, R.. 2006. Distribution of 14C-imidacloprid in Fraxinus spp. and effects of imidacloprid on the emerald ash borer. p. 24-25, In: Mastro, V., R. Reardon and G. Parra, compilers. Proceedings of the Emerald Ash Borer Research and Technology Development Meeting. September 26-27, 2005. Pittsburg, Pennsylvania. USDA Forest Service, Forest Health Technology Enterprise Team, FHTET-2005-16.
• Siegert, N.W., McCullough, D.G., Liebhold, A.M. and Telewski, F. 2006. Spread and dispersal of emerald ash borer: a dendrochronological approach. p. 10, In: Mastro, V., R. Reardon and G. Parra, compilers. Proceedings of the Emerald Ash Borer Research and Technology Development Meeting. September 26-27, 2005. Pittsburg, Pennsylvania. USDA Forest Service, Forest Health Technology Enterprise Team, FHTET-2005-16.
• Fraser, I., McCullough,D.G., Poland,T.M., Cappaert, D. and Mastro, V. 2006. Survival of emerald ash borer in chipped and ground ash. p. 24-25, In: Mastro, V., R. Reardon and G. Parra, compilers. Proceedings of the Emerald Ash Borer Research and Technology Development Meeting. September 26-27, 2005. Pittsburg, Pennsylvania. USDA Forest Service, Forest Health Technology Enterprise Team, FHTET-2005-16.
• Poland, T.M., Rodriguez-Sanoa, C., Grant,G., Buchan,L., Miller, J.E. and McCullough, D.G. 2006. Trapping and detection of emerald ash borer: identification of stress-induced volatiles and tests of attraction in the field and lab. p. 64-65, In: Mastro, V., R. Reardon and G. Parra, compilers. Proceedings of the Emerald Ash Borer Research and Technology Development Meeting. September 26-27, 2005. Pittsburg, Pennsylvania. USDA Forest Service, Forest Health Technology Enterprise Team, FHTET-2005-16.

Progress 01/01/05 to 12/31/05

Outputs
Major research efforts in 2005 focused on methods to detect, survey and control emerald ash borer (EAB), a new exotic pest that threatens ash species throughout North America. Studies are underway to address methods for EAB detection and survey, population dynamics and rate of spread, and evaluation of insecticides for adult and larval control. Andrea Anulwicz completed field and lab studies for her M.S. research that addresses EAB host range and preference. Andrew Tluczek began his M.S. program in fall 2005 and started research on the influence of EAB density and tree stress on larval development rate. Nancy Schwalm continued her research on the spread and impacts of beech scale and beech bark disease in northern Michigan. Nathan Siegert continues his post-doctoral research on EAB spatial and temporal dynamic. We collected ground beetles (Family Carabidae) in jack pine sites in northern Michigan that regenerated following fire, light harvest and intensive harvest. Beetle abundance and species diversity will be determined and compared among sites to assess effects of natural and human-caused disturbance. A comprehensive record of EAB discovery, biology, impacts and research efforts was published. A second manuscript that addresses EAB effects on ash in forested settings is in press. A paper that summarizes 17 years of exotic pest interception records at US ports of entry is in press and another manuscript that addresses the rate of exotic insect arrival in the US was published. A paper that reports results of research on pine needle scale including biological control was published.

Impacts
Results from many of our emerald ash borer studies have been directly incorporated into the multi-agency EAB programs in the U.S. More than 20 previously unknown EAB outlier populations were discovered using trap tree and survey methods we developed. Results from our dendrochronological work showed that most EAB outlier populations were established prior to the identification of this exotic pest in 2002. Recommendations based on results from our insecticide studies are widely used by landscapers, arborists and regulatory officials to control or contain EAB in affected areas. Our research on arrival rates of exotic plant-feeding insects and our studies of the 17-year history of exotic plant pest interception records will help regulatory agencies identify high risk pathways and commodities associated with exotic plant pest invasion. Data and maps resulting from our studies on beech scale and beech bark disease will continue to be used to assist public and private land managers develop harvest and management plans. Our work on ground beetles in jack pine stands will help resource managers compare the impacts of intensive harvesting and wildfire on insect biological diversity.

Publications

• Kim, J.W., L.M. Matuana and D.G. McCullough. 2005. Emerald ash borer infested ash trees as raw material for wood-based composites. Forest Products Journal 55(11):89-92.
• Cappaert, D., D.G. McCullough, T.M. Poland and N.W. Siegert. 2005. Emerald ash borer in North America: a research and regulatory challenge. American Entomologist 51(3):152-165.
• McCullough, D.G., D.L. Cappaert and T.M. Poland. 2005. Evaluation of insecticides for control of emerald ash borer: summary of 2004 trials. Published on www.emeraldashborer.info.
• Cook, W.E. and D.G. McCullough. 2005. Emerald ash borer and your woodland. MSU Extension Bulletin E-2943. 4 p.
• Rebek, K.A., E.J. Rebek and D.G. McCullough. 2005. Dont be fooled by look-alikes of the emerald ash borer and Asian longhorned beetle. MSU Extension bulletin E-2944. 2 p.

Progress 01/01/04 to 12/31/04

Outputs
Graduate students = In 2004, Nathan W. Siegert completed his Ph.D. in Entomology, Andrew Klein completed his M.S. in Forestry and Andrea Agius continued work on her M.S. in Forestry. Amy Kearney defended her M.S. in Entomology and is revising her thesis. Major research projects included analysis of a 17-year database on interceptions of nonindigenous insects and other plant pests at US ports of entry and borders. Results include summaries of origins, commodities and arrival points associated with high numbers of plant pest interceptions. We also analyzed 4 years of data from random samples of cargo arriving in four pathways and predicted that at least 42 new species of plant-feeding insects may have become established between 1997-2001. One manuscript is in press and another was accepted, pending minor revision. Major research efforts in 2004 focused on emerald ash borer, a new exotic pest that threatens all ash species in North America. Studies addressed rate of spread and population dynamics of the borer, its host range and host preference, methods for detection, survey and trapping, and evaluation of insecticides for its control. A manuscript that addresses alternatives to conventional insecticides to control pine needle scale in Christmas tree fields is in press. A second manuscript that addresses biological control of balsam twig aphid in Christmas tree plantations was published. Two manuscripts are in preparation that summarize field and lab studies addressing the gypsy moth fungus Entomophaga maimaiga, an important biological control agent. Meteorological modeling using research results indicates that the fungus may not strongly regulate gypsy moth populations in northern portions of the Lake States region. Data collection from a field study of the impacts of beech bark disease was completed and analysis is underway. We are continuing to incorporate results of our recent jack pine research into models that can be used by resource managers for planning and decision support.

Impacts
Results of our emerald ash borer research addressing rate of spread, population dynamics, insecticide evaluations, trap tree utilization and related studies continue to form the basis for the multi-agency EAB eradication projects in the U.S. and Canada. Results from our analysis of exotic pest interception data identified the relative risk associated with four pathways including marine and air cargo and border crossings. Results will help federal and state regulatory agencies identify high risk commodities and revise exotic pest interception procedures. This should reduce the likelihood of new exotic pest introduction and establishment. Models and maps of climatic suitability generated in our research on Entomophaga maimaiga have been requested by forest health managers in other North Central states. This information is being used to asses potential benefits of introducing this attractive biological control agent and to identify areas where it may not be effective. Data collected from our beech bark disease research has been provided to federal, state and private landowners to assist them in developing salvage and management plans. Results from research on the health of jack pine stands and impacts of jack pine budworm defoliation is being integrated with national growth and yield models, which will be used by forest managers to predict budworm impact, fire risks and prioritize stands for harvest, pre-salvage or salvage activities.

Publications

• Fondren, K.M. and D.G. McCullough. 2004. Phenology, natural enemies and horticultural oil for control of pine needle scale (Chionapsis heterophylla) (Fitch) (Homoptera: Diaspidae) on Christmas tree plantations. Journal of Economic Entomology. In press.
• Fondren, K.M., D.G. McCullough and A.J. Sommers. 2004. Insect predators and augmentative biological control of balsam twig aphid (Mindarus abietinus) (Homoptera: Aphididae) on Christmas tree plantations. Environmental Entomology. In press.
• Work, T.T., D.G. McCullough, J.F. Cavey and R. Komsa. 2004. Approach rate of nonindigenous insect species into the United States through cargo pathways. Biological Invasions. In press.
• Hajek, A.E., N.W. Siegert, M.M. Wheeler and D.G. McCullough. 2004. Using bioassays to estimate the abundance of Entomophaga maimaiga resting spores in soil. Journal of Invertebrate Pathology 86:61-64.
• McCullough, D.G., D.R. Smitley, D. Cappaert and T. Poland. 2004. Preliminary results from 2003 research on insecticides for emerald ash borer control. Distributed via the multi-agency EAB web site at www.emeraldashborer.info, and in printed form.
• McCullough, D.G. and S.A. Katovich. 2004. Pest Alert - Revision. Emerald ash borer. Publication No. NA-PR-02-04. USDA Forest Service, State and Private Forestry, Northeastern Area. 2 pp.
• McCullough, D.G. and S.A. Katovich. 2004. Focus on invasive species: how to identify emerald ash borer. Journal of Forestry 102:4-5.

Progress 01/01/03 to 12/31/03

Outputs
Graduate students = Laura Lazarus completed her M.S. in Entomology; 3 other students are working on their M.S.degrees; 1 student is working on his Ph.D. Major research projects currently in progress include studies related to biology, control and management of the emerald ash borer, a newly discovered exotic pest. We monitored the phenological development of emerald ash borer in the field, continued to evaluate insecticide products, timing and application methods, monitored survival of larvae in wood chips and initiated studies to assess host range of this tree-killing pest. We cooperated in studies to evaluate potential detection and survey tools for emerald ash borer including girldling and herbicide treatments, and lure/trap combinations. We analyzed a major dataset consisting of exotic insects detected in random samples of cargo entering U.S. ports of entry from 1997-2001. A manuscript detailing results, including potential numbers of species arriving via four cargo pathways, was accepted for publication, pending revision. Additional analyses of a related exotic pest interception dataset are continuing. We completed field and laboratory work and climate modeling in a study to evaluate factors limiting the success of the gypsy moth biological control agent Entomophaga maimaiga. Results suggest that spring precipitation has strong effects on resting spore germination and climatic variability may limit the effectiveness of this biocontrol in northern regions of Michigan and surrounding states. Baseline data has been collected from 62 northern hardwood stands to document the long-term impacts of beech bark disease on species composition, forest productivity and wildlife habitat in stands with varying levels of beech overstory. Data collected in jack pine stands is currently being analyzed to assess live and dead jack pine volume, effects of jack pine budworm defoliation on tree growth, tree mortality and accumulation of coarse woody debris in seven regions of northern Michigan and central Wisconsin. Research on pine shoot beetle colonization of native and exotic pine species in three regions of Michigan was published. Results from research on phenology, damage and economic injury associated with balsam twig aphid, a major Christmas tree pest, were published.

Impacts
Results of emerald ash borer research related to larval survival in chips, insecticide evaluations, trap tree utilization and related studies have already been incorporated into the development and implementation of the multi-agency EAB eradication projects in the U.S. and Canada. Results from our analysis of exotic pest interception data identified the relative risk associated with four pathways including marine and air cargo and border crossings. Results will help federal and state regulatory agencies revise exotic pest interception procedures. This should reduce the likelihood of new exotic pest introduction and establishment. Models and maps of climatic suitability generated in our research on Entomophaga maimaiga have been requested by forest health managers in other North Central states. This information is being used to asses potential benefits of introducing this attractive biological control agent and to identify areas where it may not be effective. Data collected from our beech bark disease research has been provided to federal, state and private landowners to assist them in developing salvage and management plans. Results from research on the health of jack pine stands and impacts of jack pine budworm defoliation can be used by forest managers to prioritize stands for harvest, pre-salvage or salvage activities.

Publications

• Fondren, K.M. and McCullough, D.G. 2003. Phenology and density of balsam twig aphid (Mindarus abietinus Koch) (Homoptera: Aphididae) in relation to bud break, shoot damage and value of fir Christmas trees. Journal of Economic Entomology. In press.
• Siegert, N.W. and McCullough, D.G. 2003. Colonization of Scotch, red and jack pine logs by Tomicus piniperda (Coleoptera: Scolytidae) in Michigan pine stands. Canadian Journal of Forest Research 33:2238-2244.
• McCullough, D.G., Poland, T. and Cappaert, D. 2003. Dispersal of emerald ash borer: a case study at Tipton, Michigan. Abstract, In: Proceedings of the joint U.S.-Canada emerald ash borer research meeting. October, Port Huron, Michigan.
• McCullough, D.G., Cappaert, D., Poland, T. and Smitley, D. 2003. Control of emerald ash borer adults and larvae with insecticides. Abstract, In: Proceedings of the joint U.S.-Canada emerald ash borer research meeting. October, Port Huron, Michigan.
• McCullough, D.G., Poland, T. and Cappaert, D. 2003. Survival of emerald ash borer in chips. Abstract, In: Proceedings of the joint U.S.-Canada emerald ash borer research meeting. October, Port Huron, Michigan.
• Agius, A., McCullough, D.G., Cappaert, D., Poland, T., Miller, D. and Bauer, L. 2003. Host range and host preference of emerald ash borer. Abstract, In: Proceedings of the joint U.S.-Canada emerald ash borer research meeting. October, Port Huron, Michigan.
• Herms, D., Smitley, D., McCullough, D. and Bonello, P. 2003. Evaluation of resistance of Asian and North American ashes to emerald ash borer. Abstract, In: Proceedings of the joint U.S.-Canada emerald ash borer research meeting. October, Port Huron, Michigan.
• Poland, T.M., De Groot, P., Grant, G., MacDonald L. and McCullough, D.G. 2003. Developing attractants and trapping techniques for the emerald ash borer. Abstract, In: Proceedings of the joint U.S.-Canada emerald ash borer research meeting. October, Port Huron, Michigan.
• Lazarus, L.L. 2003. Monochamous scutellatus host preference and suitability of four conifer species for larval development. M.S. thesis, Dept. of Entomology, Michigan State University. 111 pp.

Progress 01/01/02 to 12/31/02

Outputs
Graduate Students = Kirsten Fondren and Linda Williams completed their MS degree; 3 other students working on MS; 1 student working on PhD. Project objectives are to increase our understanding of forest insect ecology and impacts, identify factors that contribute to insect damage in forests, acquire information on ecological roles of forest insects, and apply this knowledge to develop sound management strategies for healthy, productive forests. A report on predicting invasions of exotic plant pests was completed and published by the National Academy of Science. Research on biology, survey and eradication methods for emerald ash borer, a new exotic forest pest, was initiated this summer. Fieldwork was completed on a study to assess the ability of the fungal pathogen Entomophaga maimaiga to persist, germinate and infect gypsy moth larvae. Modeling to evalulate suitability of climate in Michigan and surrounding states for epizootics of E. maimaiga is in progress. Analysis of exotic plant pest interception records from U.S. ports of entry is in progress to identify biological or economic factors associated with high pest arrival rates. Results of these analyses include prediction of arrival rates of exotic plant-feeding insect species; a manuscript was submitted for publication. Fieldwork is underway to collect baseline data and quantify impacts of beech bark disease on forest productivity, species composition and wildlife habitat in Michigan. Field and lab studies to evaluate host preference and host suitability of North American trees for the exotic Asian longhorned beetle and native pine sawyer longhorned beetles was completed and one manuscript has already been published. Alternative methods to control balsam twig aphid and pine needle scale, the most important insect pests on fir and Scotch pine Christmas trees, respectively, was completed. Results of field studies on preference of the exotic pine shoot beetle for native and exotic pines were submitted for publication. Research demonstrating competition between the exotic pine shoot beetle and native phloem-feeding insects and predation of the exotic pine shoot beetle by native beetles and wasps was published.

Impacts
Our results will improve the ability of regulatory officials and ecologists to develop sound policies to reduce risk and impact of exotic pests. Our data on emerald ash borer will be critical for success of the the eradication program proposed for this major pest. Information we acquired on Entomophaga maimaiga will assist managers in predicting the effectiveness of this pathogen as gypsy moth expands into north central states. Estimates of jack pine budworm impact on volume and coarse woody debris will enable forest managers to prioritize stands for harvest, salvage or conversion and to predict impacts of future budworm outbreaks. Our data on beech bark disease will be used to assess changes in hardwood forests over time and to develop silvicultural guidelines as beech bark disease spreads. Information on longhorned beetle host preference and suitability of hosts for larval development will be used in the Asian longhorned beetle eradication program and expands our understanding of longhorned beetle behavior and ecology. Results of studies on balsam twig aphid and pine needle scale will reduce insecticide use and increase profits in Michigan Christmas tree production. Results of studies on pine shoot beetle show that importation and release of a European bark beetle predator, as proposed by federal agencies, was unnecessary, especially given the risks associated with this generalist, exotic predator. Proposed releases of the exotic predator were rejected.

Publications

• National Research Council, Board of Agriculture and Natural Resources, National Academy of Science. 2002. Predicting invasions of nonindigenous plants and plant pests. National Academy Press, Washington, DC. 198 pp. Kennedy, A.A. and D. G. McCullough. 2002. Phenology of the exotic pine shoot beetle (Tomicus piniperda [L.]) (Coleoptera: Scolytidae) in relation to native bark beetles and natural enemies in red pine stands. Environmental Entomology 31:261-272.
• Work, T.T. and D.G. McCullough. 2002. Lepidopteran community response to gypsy moth outbreaks is robust to guild reclassification: a reply to comments of Summerville and Crist. Environmental Entomology 31:584-587.
• Ludwig, S., L. Lazarus, D.G. McCullough, K. Hoover, S. Montero and J.C. Sellmer. 2002. Methods to examine host tree susceptibility to the Asian longhorned beetle Anoplophora glabripennis. J. Environmental Horticulture 20:175-180.
• Hajek, A.E., N.W. Siegert, D.G. McCullough and L. Butler. 2002. Activity of Entomophaga maimaiga in the field. In: Proceedings, National Exotics Review. January 16-18, 2002, Annapolis, Maryland. USDA Forest Service, Northeastern Res. Sta. Gen. Tech. Report.
• Poland, T.M., D.G. McCullough, T.R. Petrice and N.W. Siegert. 2002. Status of beech bark disease establishment and research in Michigan. Abstract, In: Proceedings, National Exotics Review. January 16-18, 2002, Annapolis, Maryland. USDA Forest Service, Northeastern Res. Sta. Gen. Tech. Report.
• McCullough, D.G. and J. Zablotney. 2002. Directory of exotic forest insect and disease pests. MSU Extension Bulletin E-2811. 42 p.
• McCullough, D.G. and D. Roberts. 2002. Pest Alert - Emerald Ash Borer. Publication No. NA-PR-07-02. USDA Forest Service, State and Private Forestry, Northeastern Area. Newtown Square, PA. 2 pp.
• Fondren,K.M. 2002. Alternative controls for two important pests of Christmas trees. M.S. thesis. Dept. of Entomology, Michigan State University. 163 pp.
• Fondren, K. and D.G. McCullough. 2002. Biology and management of pine needle scale. MSU-Extension Bulletin E-2812. 4 pp.
• Fondren, K. and D.G. McCullough. 2002. Biology and management of balsam twig aphid. MSU-Extension Bulletin E-2813. 4 pp.
• Scarr, T., D.G. McCullough and G. Howse. 2002. New forest pest - emerald ash borer. Canadian Forest Service and Ontario Ministry of Natural Resources. 4 pp
• Williams, L.K. 2002. White pine regeneration in Michigan: evaluation of white pine weevil and white pine blister rust. M.S. thesis. Dept. of Entomology, Michigan State University. 127 pp.

Progress 01/01/01 to 12/31/01

Outputs
Graduate Students = 1 Ph.D. and 4 M.S. students in progress. Objectives are to increase our knowledge of forest insect ecology, develop sound managment strategies for forest insect pests, and acquire information on forest insect communities, ecological roles of insects in forested ecosystems, and effects of disturbance on forest insects. Results of research on the impact and host preference of the exotic bark beetle Tomicus piniperda (pine shoot beetle) were published. A manuscript reporting phenology of the exotic pine shoot beetle and its interactions with native bark beetles and native predators is in press. A paper presenting results of a major field study on effects of winter weather on survival of gypsy moth egg masses is in press. I wrote several sections of a National Academy of Science NRC report on predicting invasiveness of nonindigenous plant-feeding insects, plant pathogens and weeds; the report is in press. Analysis of the USDA Animal and Plant Health Inspection Service (APHIS) Port-Interception-Network database was intiated. This database includes nearly 1 million records of interceptions of exotic plant pests at US borders and ports of entry and ecological patterns in interceptions have not been previously examined. Field and lab work to address effects of weather, stand and site-related variables on the distribution and persistence of the fungus Entomophaga maimaiga and the gypsy moth nucleopolyhedrosis virus, the two most important biological controls of gypsy moth, is continuing. Fieldwork on a project to examine alternative methods of controlling balsam twig aphid and pine needle scale in Christmas tree fields was completed and data analysis is underway. Results to-date indicate that biological controls, host resistance and horticultural oil are effective alternatives to conventional chemical insecticides for these two pests. Host preference of the pine sawyer, a native longhorned wood-boring beetle, was evaluated in field and lab studies. Competition between this species and other phloem-feeding insects is currently being examined. Establishment of a state and regional network of plots to assess impact of jack pine budworm outbreaks was initiated. Tree mortality, topkill, growth and volume loss were related to budworm defoliation and site or stand variables. Studies were begun to evaluate impact and potential rate of spread of beech bark disease in Michigan forests. Effects of beech scale and the associated Nectria fungi on forest composition, productivity and wildlife habitat will be examined.

Impacts
Results of this research will improve our ability to reduce damage from major exotic forest pests such as gypsy moth, pine shoot beetle and beech bark disease, and from native pests such as jack pine budworm and bark beetles. Our studies will lead to improved recommendations for managing forest insect pests effectively using silviculture and biological control in forested ecosystems and in Christmas tree production, reducing the need for microbial or conventional insecticide application. Christmas tree growers in Michigan reduced insectide use for balsam twig aphid control in 2001 because of our research results. Results will help identify temporal and spatial patterns in the population dynamics and distribution of important forest insects. This information will help forest managers to assess risks of pest damage under different harvesting or management scenarios, predict impacts on productivity, and ensure that forests are managed on a sustained yield basis.

Publications

• McCullough, D.G., R.L. Heyd and J.G. O'Brien. 2001. Biology and management of beech bark disease. MSU-Extension Bulletin E-2746. 12 pp.
• Katovich, S.A., A.S. Munson, J. Ball and D. McCullough. 2001. Bronze birch borer. USDA Forest Service, Forest Insect and Disease Leaflet 111. 8 pp. St.Paul, Minnesota.
• Kennedy, A.A. and D. G. McCullough. 2002. Phenology of the exotic pine shoot beetle (Tomicus piniperda [L.]) (Coleoptera: Scolytidae) in relation to native bark beetles and natural enemies in red pine stands. Environmental Entomology. In press.
• Andresen J. A., D.G. McCullough, B.E. Potter, N. Koller, L.S. Bauer, D.P. Lusch and C.W. Ramm. 2002. Effects of winter temperatures on gypsy moth egg masses in the Great Lakes region. Agricultural and Forest Meteorology. In press.
• Siegert, N.W. and D.G. McCullough. 2001. Survey of shoot damage caused by the exotic pine shoot beetle in Michigan pine stands. Northern Journal of Applied Forestry 18:101-109.
• Siegert, N.W. and D.G. McCullough. 2001. Host preference of pine shoot beetle (Tomicus piniperda [L.])(Coloeptera: Scolytidae) parent and progeny adults. Canadian Entomologist 133:343-353.
• Mack, R.N. et al. (Multiple authors). 2001. Scientific basis for predicting invasiveness of plant pests in natural and managed ecosystems. Board of Agriculture and Natural Resources, National Research Council, National Academy of Sciences.Washington, D.C. In press.
• McCullough,D.G., A.E. Hajek, N.W. Siegert, M. Wheeler, R.C. Venette, W.C. Kaufmann. 2001. Factors influencing success of the gypsy moth biological control Entomophaga maimaiga. Abstract, p. 91-92.In: Proceedings, National Exotics Review. January 16-18, 2001, Annapolis, Maryland. USDA Forest Service, Northeastern Res. Sta. Gen. Tech. Rep. NE-285.
• Siegert, N.W., D.G.McCullough, A.E.Hajek and M.M.Wheeler. 2001. An evaluation of the gypsy moth fungus Entomophaga maimaiga (Zygomycetes:Entomophthorales) in Michigan forests. Abstract, p. 207. In: Proc. North American Forest Insect Work Conference. May 14-18. Edmonton, Alberta, Canada.
• McCullough, D.G. 2001. Comparison of eastern tent caterpillar, forest tent caterpillar and gypsy moth. MSU-Extension Bulletin E-2299. 2 pp.

Progress 01/01/00 to 12/31/00

Outputs
Graduate students = 1 completed M.S.; 4 in progress. Objectives are to increase our knowledge of forest insect ecology, develop sound management strategies, and acquire information on forest insect communities, ecological roles of insects in forest ecosystems and disturbance effects on forest insects. Results of research on forest insect communities in two hardwood ecosystems and effects of gypsy moth invasion on those communities were published. Fieldwork addressing effects of weather, parasitism and other factors on gypsy moth egg survival was completed; data analysis is in progress. Distribution of the Entomophaga maimiaga fungus, a gypsy moth biocontrol agent, its interactions with the other major gypsy moth pathogen, and its impact on gypsy moth populations is currently being examined. We also initiated work to model suitability of climate in Michigan and north central states for this recently introduced fungus. We completed two additional white pine plantings to establish long-term stands to evaluate associations between regeneration methods and white pine weevil damage. A project on biocontrol, host resistance and other alternative methods to control pine needle scale and balsam twig aphid on Christmas trees is in progress. Preliminary results were presented in trade publications and presentations to growers. Host preference of Asian longhorned beetle was examined; further studies with similar, native beetles are planned. Research on pine shoot beetle host preference was completed; 2 papers were submitted and one more is in preparation. Funding to initiate a major study of exotic insect, plant pathogen and weed introductions using federal port interception data was obtained. Work with several collaborators will begin in 2001. Jack pine budworm population dynamics, including historical outbreak patterns in the Lake States, were reviewed and a paper was published. Futher jack pine budworm research will occur in 2001. Beech bark scale and beech bark disease were discovered in Michigan for the first time; efforts were made to cooperate in surveys, development of management recommendations and training sessions. More research on dispersal, impact and management of these pests will occur in 2001.

Impacts
Results should improve our ability to use silvicultural methods to effectively reduce damage from major pests including exotic species such as gypsy moth, pine shoot beetle and Asian longhorned beetle, and native insects such as jack pine budworm and white pine weevil. Results will help managers understand temporal and spatial dynamics of gypsy moth, jack pine budworm and pine shoot beetle populations, improving our ability to predict density or impacts of these pests. Results will also enhance efforts to use biological control to effectively manage pests in forest and Christmas tree systems, reducing the need for microbial or conventional insecticide use. Increased understanding of specific exotic pests, and the impacts of disturbance caused by their introduction will be used to develop management options for these and future introductions.

Publications

• McCullough, D.G. 2000. A review of factors affecting the population dynamics of jack pine budworm (Choristoneura pinus pinus Freeman). Population Ecology. In press.
• Work, T.T. and D.G. McCullough. 2000. Effects of gypsy moth invasion on Lepidopteran communities in two forest ecosystems in northern Michigan. Environmental Entomology 29:884-900.
• McCullough, D.G. and L.A. Leefers. 2000. Economic consequences of jack pine budworm outbreaks. Final Report. Status of Michigan Natural Resources Special Report. Michigan Agric. Exp. Stn. Report No. 105.
• McCullough, D.G., J.A. Andresen, N. Koller, L. Bauer, D. Miller and C.W. Ramm. 2000. Factors affecting survival of gypsy moth eggs in Michigan. Abstract. In: Proceedings, National Gypsy Moth Review. November 1-4, 1999, Madison, Wisconsin. National Gypsy Moth Management Board. In press.
• McCullough, D.G., R.L. Heyd, J.G. O'Brien and W. Cook. 2000. Beech bark disease. Michigan Society of American Foresters web site -(hot topic). http://forestry.msu.edu/msaf/main
• Bauer, L.S. and D.G. McCullough. 2000. Bt - one option for gypsy moth management. MSU-Extension Bulletin E-2724. 4 pp.

Progress 01/01/99 to 12/31/99

Outputs
Graduate students = 4 in progress. Objectives are to increase our understanding of forest insect ecology, and impacts and factors that contribute to insect damage in forests; apply this knowledge to develop sound management strategies; and acquire information on forest insect communities, ecological roles of insects in forest ecosystems, and effects of disturbance on forest insects. Research on economic and biological impacts of jack pine budworm defoliation was completed and results were published. A study on accuracy and efficiency of gypsy moth egg mass survey methods was completed and results were published. Research on factors affecting survival of gypsy moth eggs is in progress; early results indicate that intense solar radiation in autumn has more impact on egg survival than winter temperatures. A new project was initiated to evaluate the distribution, abundance and impact of the Entomophaga maimaiga fungus on gypsy moth populations in Michigan, and how this biocontrol agent is affected by weather and site or stand conditions. We continued surveys to quantify the impact of white pine weevil and white pine blister rust in existing white pine stands. Establishment of new white stands to evaluate alternative regeneration methods is in progress. Host preference of the exotic pine shoot beetle and its impact on red, jack and scotch pine forests was examined; data analysis is in progress. We began a new project to examine host preference of the exotic Asian longhorned beetle, using quarantine facilities in Canada. A major manual on the use of biological control in forests, Christmas tree plantations and urban forests was completed and published.

Impacts
Results of research should improve the ability of forest managers to use silviculture methods to effectively reduce damage from jack pine budworm, white pine weevil, pine shoot beetle and gypsy moth. New information will also help gypsy moth program managers and forest planners anticipate population size or impacts of these pests. Results will improve understanding and interest in using biological controls in forest and Christmas tree systems, reducing the use of microbial or conventional insecticides.

Publications

• McCullough, D.G., N.W. Siegert and A.A. Kennedy. 1999. Pine shoot beetle in Michigan forests. Abstract, In: Proceedings of the 10th USDA Interagency Research Forum on Gypsy Moth and Other Invasive Species. January 19-22, Annapolis, Maryland.
• McCullough, D.G., J.A. Andresen, L. Bauer, N. Koller, D. Miller and C.W. Ramm. 1999. Effects of abiotic factors on gypsy moth egg survival in Michigan. Abstract, In: Proceedings of the 10th USDA Interagency Research Forum on Gypsy Moth and Other Invasive Species. January 19-22, Annapolis, Maryland.
• McCullough, D.G., J.A. Andresen, N. Koller, L. Bauer, D. Miller and C.W. Ramm. 1999. Factors affecting survival of gypsy moth eggs in Michigan. Abstract. In: Proceedings, National Gypsy Moth Review. November 1-4, 1999, Madison, Wisconsin. National Gypsy Moth Management Board. (In press).
• McCullough, D.G., S. Katovich, D. Neumann, D. Mahr, C. Sadof and M. Raupp. 1999. Biological control of insect pests in forested ecosystems: a manual for foresters, Christmas tree growers and landscapers. MSU Bulletin E-2679. 123 p.
• Buss, L.J., D.G. McCullough and C.W. Ramm. 1999. Evaluation of three egg mass survey methods in relation to gypsy moth (Lepidoptera: Lymantriidae) defoliation in Michigan. Environmental Entomology 28:485-495.
• Conway, B.E., L.A. Leefers, and D.G. McCullough. 1999. Financial evaluation of a jack pine budworm outbreak and implications for management. Canadian Journal of Forest Research 29:382-392.
• Conway, B.E., D.G. McCullough and L.A. Leefers. 1999. Long-term effects of jack pine budworm outbreaks on the growth of jack pine trees in Michigan. Canadian Journal of Forest Research 29:1510-1517.
• McCullough, D.G. 1999. Diversity of native natural enemies of pine shoot beetle and their potential impact on biological control. Abstract. In: W.D. Hutchison, (ed.). Symp. on exotic pests: new opportunities for classical biological control in the Midwest. Entomological Society of America, North Central Branch meeting. March 28-31, Des Moines, Iowa.

Progress 01/01/98 to 12/31/98

Outputs
Graduate students = 2 (B.E. Conway, MS 98; A.A. Kennedy, MS 1998). Objectives are to increase our understanding of forest insect ecology and impacts andfactors that contribute to insect damage, apply this knowledge to develop sound management strategies, and acquire information on forest insect communities, ecological roles of insects in forested systems, and effects of disturbance on forest insects. Results of research on jack pine budworm impacts were used to develop models and subsequently a decision support system (DSS). The DSS enables forest managers to use stand inventory data to predict volume and economic loss due to tree mortality and reduced growth, and identify stands that will sustain the greatest loss during a budworm outbreak. The DSS was delivered and is currently in use by the Hiawatha National Forest. Effects of parasitism and weather variables on gypsy moth egg mortality were experimentally evaluated in an ongoing study. A study on effects of gypsy moth invasion on native insect communities was completed; one report was published and another manuscript is in review. We began a study to assess methods of white pine regeneration in open plantations and below an existing overstory to reduce white pine weevil damage to young trees. We continued to evaluate the impacts of the exotic pine shoot beetle in Michigan pine forests, its phenology, host preference, and interactions with native bark beetles. We published data used to develop the Pine Shoot Beetle Compliance Program for Christmas Trees, and further refined recommendations for Christmas tree growers affected by the pine shoot beetle quarantine. A major reviw paper addressing interactions between fire and insects in forested systems was published.

Impacts
(N/A)

Publications

• McCullough, D.G. and C.S. Sadof. 1998. Evaluation of an integrated management and compliance program for Tomicus piniperda (Coleoptera: Scolytidae) in pine Christmas tree fields. Forum Article, Journal of Economic Entomology 91:785-795.
• McCullough, D.G., R.A. Haack and W.H. McLane. 1998. Control of Tomicus piniperda (Coleoptera: Scolytidae), the pine shoot beetle, in pine stumps and logs. Journal of Economic Entomology 91:492-499.
• Smitley, D., J. Andresen, R. Priest, R. Mech and D.McCullough. 1998. Winter mortality of gypsy moth (Lepidoptera: Lymantriidae) eggs in Michigan. Environmental Entomology 27:700-708.
• Work, T.T., D.G. McCullough and W.J. Mattson. 1998. Moth and carabid beetle species associated with two ecological landtype phases in northern Michigan. USDA Forest Service, North Central Research Station, General Technical Report NC-201. 25 pp.
• Bauer, L., N. Koller, D. McCullough, J. Andresen and C. Ramm. 1998. Potential roles of biotic and abiotic factors in gypsy moth egg mortality in 1996-97 in northern and southern Michigan counties. In: Proceedings, 1998 Gypsy Moth Research Review. January, Annapolis, Maryland. USDA Forest Service, Northeastern For. Exp. Stn.
• McCullough, D.G., R.A. Werner and D. Neumann. 1998. Fire and insects in northern and boreal forests of North America. Annual Review of Entomology 43:107-127.
• McCullough, D.G. 1998. Summary of research on pine shoot beetle at Michigan State University. In: Proc. 1998 Pine shoot beetle workshop. February 17, 1998, Toronto, Ontario.
• Conway, B.E., D. McCullough and L. Leefers. 1998. Computer simulation of jack pine budworm impact in Michigan' Upper Peninsula. In: J.M. Vasievich, J. Fried and L. Leefers (eds.). Proceedings of the Seventh Symposium on Systems Analysis in Forest Resources; Bellaire, Michigan, May 28-31, 1997. USDA Forest Service North Central Research Station General Technical Report. (In press).

Progress 01/01/97 to 12/31/97

Outputs
Number of Graduate Students = 1 (T.T. Work, M.S. in 1996) Objectives of this research were to examine the abundance, diversity and community structure of native forest insects in sugar maple ecosystems and red oak ecosystems. Half of the red oak ecosystems were severely defoliated by gypsy moth in at least 1 year of the study. Gypsy moth defoliation was quantified using a ceptometer. Abundance of some groups of specialized native oak-feeding Lepidoptera declined after gypsy moth outbreaks, but most other groups recovered within 1 year. Carabid beetle populations increased after severe defoliation, possibly due to altered microclimate or increased prey abundance. Diversity and abundance of Lepidoptera and carabids was variable among the 3 study years, but differed significantly between ecosystems in all years.

Impacts
(N/A)

Publications

• No publications reported this period

Progress 01/01/96 to 12/30/96

Outputs
Objectives of this research were to examine the abundance, diversity and community structure of native forest insects in sugar maple/basswood ecosystems and less productive red oak ecosystems. Sugar maple ecosystems were not defoliated by gypsy moth. Half of the oak sites had high gypsy moth populations in 1993, while the other half were not yet infested. In 1994 and 1995, the original outbreak sites sustained little defoliation while the other sites were severely defoliated. A ceptometer was useful for quantifying light penetration and defoliation severity in oak stands. Abundance of some adult Lepidopteran species was reduced the year after severe defoliation in oak sites, but most species recovered the following year. Oak-feeding noctuids were reduced for at least two years after defoliation. Lepidopteran communities significantly varied between oak and sugar maple ecosystems. Some carabid beetle species increased the year after severe defoliation, possibly due to altered prey availability or microclimate. Carabid beetle communities also differed between the two ecosystems. Community structure was assessed by evaluating abundance of guilds of native insects. Abundance differed between the two ecosystems but was not affected by gypsy moth. Graduate Students: 3.

Impacts
(N/A)

Publications

• MCCULLOUGH, D. G., WORK, T.T. 1996. Eff of gypsy moth invasion on native insect comm. In: The gypsy moth, a changing horizon. Proc. Natl Gyp Moth Rev. Traverse City, MI. Nov 1995. Natl Gypsy Moth Mgmt Board. p. 108-109.

Progress 01/01/95 to 12/30/95

Outputs
Objectives of this research were to examine the abundance, diversity and community structure of native forest insects in sugar maple/basswood ecosystems and less productive red oak ecosystems. Red oak ecosystems were very susceptible to gypsy moth defoliation; sugar maple ecosystems were much less susceptible. Half of the sites in each ecosystem had high gypsy moth populations in 1993, while the other half had not yet been heavily infested. Gypsy moth populations changed substantially from 1993 to 1994, with previous outbreak-level populations collapsing. In 1995, previously undefoliated red oak sites experienced heavy defoliation. Lepidopteran species and families of other insects associated with canopy, shrub and ground-level strata were determined for 1993 and 1994. Preliminary results indicate substantial differences between the two ecosystems in abundance of some insect families. Heavy gypsy moth defoliation in two red oak sites was associated with low abundance or diversity of some lepidopteran species in 1993. Lepidopteran species apparently recovered in 1994 and no differences between sites with heavy and low defoliation were observed. Identification of insects collected in 1995 is in progress. Carabid beetles collected from 1993-1995 are also being identified to species level. Analysis of effects of ecosystem type and gypsy moth defoliation on abundance and diversity of feeding guilds is in progress.

Impacts
(N/A)

Publications

• NO PUBLICATIONS REPORTED THIS PERIOD.

Progress 01/01/94 to 12/30/94

Outputs
Objectives of this research are to examine the abundance, diversity and community structure of native forest insects in productive sugar maple ecosystems and less productive red oak ecosystems. Half of the sites in each ecosystem had high gypsy moth populations, while the other half had low populations. Gypsy moth populations changed substantially from 1993 to 1994, with previous outbreak-level populations collapsing. Preliminary results indicate substantial differences between the two ecosystems in abundance of number of families of some groups. Ground inhabiting beetles, Nocutid moths, and shrub-inhabiting insects were most strongly affected by ecosystem. Gypsy moth populations were high in two of the red oak ecosystems, and some families of foliage-feeding lepidoptera were affected. Analysis of 1993 data and identification of insects collected in 1994 is in progress.

Impacts
(N/A)

Publications

• NO PUBLICATIONS REPORTED THIS PERIOD.

Progress 01/01/93 to 12/30/93

Outputs
Objectives of this research are to examine the abundance, diversity and community structure of native forest insects in sugar maple and red oak ecosystems, and in areas with high and low populations of gypsy moth, an exotic defoliator. Eight sites were located in the Manistee National Forest; four sites in each of the two ecosystems. Half of the sites in each ecosystem have high gypsy moth populations and half have low gypsy moth populations. Samples of insects were collected at four times corresponding to aerial application of microbial insecticides to suppress gypsy moth populations, mid-way through gypsy moth development, peak defoiation and after trees refoliated. Ground, shrub, lower canopy and upper canopy strata were sampled using pitfall traps, sweepnets, Monarch nets, Malaise traps, clipped branches and modified black light traps. A specimen collection was developed and identification of all insects to the family level was completed. Identification of adult lepidopteran species is continuing. Indices of family levels diversity is in progress. Each family will be assigned to a feeding guild and ratios among guilds (e.g. predator:herbivore) will be calculated to assess community structure. Graduate Students = 1.

Impacts
(N/A)

Publications

• NO PUBLICATIONS REPORTED THIS PERIOD.