Understanding and quantifying invasive species biology, ecology, interactions, and impacts - USDA

UNDERSTANDING AND QUANTIFYING INVASIVE SPECIES BIOLOGY, ECOLOGY, INTERACTIONS, AND IMPACTS

Sponsoring Institution

Forest Service/USDA

Project Status

EXTENDED

Funding Source

USDA INHOUSE

Reporting Frequency

Annual

Accession No.

0413624

Grant No.

(N/A)

Cumulative Award Amt.

(N/A)

Proposal No.

(N/A)

Multistate No.

(N/A)

Project Start Date

Oct 1, 2008

Project End Date

Sep 30, 2021

Grant Year

(N/A)

Program Code

[(N/A)]- (N/A)

Recipient Organization
USDA - Forest Service
NORTHERN RESEARCH STATION
NEWTOWN SQUARE,PA 19073

Performing Department
(N/A)

Non Technical Summary
This research problem area will develop basic biological and ecological information about invasive insects, diseases, and plants (collectively pests) that are currently in the United States. This change of knowledge will allow scientists, managers, and citizens to develop control treatments and management tactics for these pests. We will also develop information on how these pests damage our native forests and what those damages cost us. This information will allow managers and policymakers to change conditions by funding management programs to reduce the costs of those damages.

Animal Health Component

35%

Research Effort Categories

Basic

60%

Applied

35%

Developmental

Classification

Knowledge Area (KA)	Subject of Investigation (SOI)	Field of Science (FOS)	Percent
123	0699	1070	20%
211	0613	1130	20%
211	0620	1130	40%
212	0620	1070	10%
213	0620	1070	10%

Knowledge Area
212 - Pathogens and Nematodes Affecting Plants; 211 - Insects, Mites, and Other Arthropods Affecting Plants; 213 - Weeds Affecting Plants; 123 - Management and Sustainability of Forest Resources;

Subject Of Investigation
0620 - Broadleaf forests of the North; 0699 - Trees, forests, and forest products, general; 0613 - Mixed conifer-broadleaf forests;

Field Of Science
1130 - Entomology and acarology; 1070 - Ecology;

Keywords

asian longhorned beetle

beech bark disease

biological invasions

biology

ecological disturbances

hemlock woolly adelgid

Goals / Objectives
This problem area focuses on research on invasive insects, pathogens, and plants that are already established in the U.S. Lack of knowledge of biology and ecology of these established species is a major barrier to their control and management. Quantifying the ecological effects and socio-economic impacts of invasive species is often critical for obtaining sufficient support to implement management programs. The unit has existing partnerships with university, state, local, and federal agencies, international governments and organizations, non-governmental organizations, and private sector cooperators to develop the needed information. The research will provide the basic biological and ecological information needed to develop control and management actions. The development of ecological effects and socio-economic impacts will provide information to managers and policymakers to enable the development and funding of management programs.

Project Methods
Our research approach uses a combination of laboratory (including quarantine), field and theoretical studies, landscape and regional scale impact studies, and modeling efforts to improve understanding of invasive species biology, life history, host range, ecology, reproductive behavior, seasonal phenology, genetic architecture, mechanisms of invasion, and population dynamics. Successional dynamics and short and long term changes in biodiversity and species composition of invaded forests provide ecological information on which social and economic effects can be evaluated. In particular, studies in our quarantine facility or overseas in an organismâ¿¿s native range, conducted with international cooperators, are critical components of the research approach. Key research subjects are invasive insect species (Asian longhorned beetle, emerald ash borer, gypsy moth, hemlock woolly adelgid, Sirex woodwasp), pathogens (beech bark disease, butternut canker, chestnut blight, and sudden oak death), and plants (tree of heaven and Japanese stiltgrass).

Progress 10/01/19 to 09/30/20

Outputs
OUTPUTS: Research in Problem 1 resulted in the publication of 23 papers, including 22 refereed articles in scientific journals and 1 Forest Insect and Disease Leaflet. Much of the research in this problem focused the biology and ecology of particular invasive species including European and Asian gypsy moth (Lymantria dispar), emerald ash borer (Agrilus planipennis) and winter moth (Operophtera brumata). Papers were published on biology and management, range expansion, and geographic-based development and survival of European gypsy moth, the potential distribution of Asian gypsy moth, spread and population dynamics of emerald ash borer, and on genetics and historical survival of winter moth. There was also considerable research on the general ecology, dynamics, drivers, and impacts of invasions. Papers were published on factors that drive invasions of plants, bark and ambrosia beetles, impacts of climate on insect outbreaks, underlying mechanisms of outbreak synchrony in foliage feeding lepidoptera, and generalized and customized spread models for non-native insects. PARTICIPANTS: Individuals who worked on Problem 1 included the investigators, Robert Haack, Nathan Havill, Melody Keena, Andrew Liebhold, Talbot Trotter, and Kimberly Wallin. Partners and collaborators included scientists from many universities, federal and state agencies, and international organizations including China, Europe, and New Zealand. TARGET AUDIENCES: The target audiences include other scientists, federal and state regulatory agencies, resource managers, policy makers, timber nursery and tree care industries, and communities. PROJECT MODIFICATIONS: No modifications occurred to this problem.

Impacts
Exotic forest pests threaten North American forests and natural resources. Native trees lack co-evolved defense mechanisms and exotic pests often invade without their associated natural enemies. Interactions of invasive pests with native ecosystems are unknown. A greater understanding of the drivers, mechanisms, and impacts of invasions are critical for developing strategies to predict, prevent, and mitigate invasions. Understanding the biology of invasive species is critical for protecting native forests and natural resources. Research in Problem 1 addresses many aspects of invasive species ecology including their biology, behavior, ecosystem interactions, economic and environmental impacts, and how quickly and how far they will spread. It also seeks to understand global patterns and factors that drive biological invasions. Results provide regulatory agencies the information necessary to make scientifically-based decisions for biosecurity surveillance and regulating movement of host material. The research is also important to resource managers, policy makers, industries, and communities for implementing regulatory and management strategies. The benefits of the research include enhanced abilities to create resilient forest ecosystems and protect them from the impacts of invasive species.

Publications

Mech, Angela M.; Thomas, Kathryn A.; Marsico, Travis D.; Herms, Daniel A.; Allen, Craig R.; Ayres, Matthew P.; Gandhi, Kamal J. K.; Gurevitch, Jessica; Havill, Nathan P.; Hufbauer, Ruth A.; Liebhold, Andrew M.; Raffa, Kenneth F.; Schulz, Ashley N.; Uden, Daniel R.; Tobin, Patrick C. 2019. Evolutionary history predicts high⿐impact invasions by herbivorous insects. Ecology and Evolution. 9(21): 12216-12230. https://doi.org/10.1002/ece3.5709.
Rutledge, Claire E; Keena, Melody A. 2019. Mating Behavior and Reproductive Biology of Emerald Ash Borer (Coleoptera: Buprestidae) and Two of Its Native Congeners, the Twolined Chestnut Borer and the Bronze Birch Borer. Journal of Economic Entomology. 112(6): 2620-2631. https://doi.org/10.1093/jee/toz182.
Schulz, Ashley N.; Mech, Angela M.; Allen, Craig R.; Ayres, Matthew P.; Gandhi, Kamal J. K.; Gurevitch, Jessica; Havill, Nathan P.; Herms, Daniel A.; Hufbauer, Ruth A.; Liebhold, Andrew M.; Raffa, Kenneth F.; Raupp, Michael J.; Thomas, Kathryn A.; Tobin, Patrick C.; Marsico, Travis D. 2020. The impact is in the details: evaluating a standardized protocol and scale for determining non-native insect impact. NeoBiota. 55: 61-83. https://doi.org/10.3897/neobiota.55.38981.
Srivastava, Vivek; Griess, Verena C.; Keena, Melody A. 2020. Assessing the Potential Distribution of Asian Gypsy Moth in Canada: A Comparison of Two Methodological Approaches. Scientific Reports. 10: 22. 10 p. https://doi.org/10.1038/s41598-019-57020-7
Liu, Xiaofei; Zhang, Liyun; Haack, Robert A.; Liu, Jiang; Ye, Hui. 2019. A noteworthy step on a vast continent: new expansion records of the guava fruit fly, Bactrocera correcta (Bezzi, 1916) (Diptera: Tephritidae), in mainland China. BioInvasions Records. 8(3): 530-539. https://doi.org/10.3391/bir.2019.8.3.08
Srivastava, Vivek; Liang, Wanwan; Keena, Melody A.; Roe, Amanda D.; Hamelin, Richard C.; Griess, Verena C. 2020. Assessing Niche Shifts and Conservatism by Comparing the Native and Post-Invasion Niches of Major Forest Invasive Species. Insects. 11(8): 479. 19 p. https://doi.org/10.3390/insects11080479.
Streifel, Marissa A.; Tobin, Patrick C.; Kees, Aubree M.; Aukema, Brian H. 2019. Range expansion of Lymantria dispar dispar (L.) (Lepidoptera: Erebidae) along its north-western margin in North America despite low predicted climatic suitability. Journal of Biogeography. 46(1): 58-69. https://doi.org/10.1111/jbi.13474.
Tejedor, Ernesto; Serrano⿐Notivoli, Roberto; Luis, Martin; Saz, Miguel Angel; Hartl, Claudia; St. George, Scott; Büntgen, Ulf; Liebhold, Andrew M.; Vuille, Mathias; Esper, Jan. 2020. A global perspective on the climate⿐driven growth synchrony of neighbouring trees. Global Ecology and Biogeography. 29(7): 1114-1125. https://doi.org/10.1111/geb.13090.
Trotter, R Talbot; Limbu, Samita; Hoover, Kelli; Nadel, Hannah; Keena, Melody A. 2020. Comparing Asian Gypsy Moth [Lymantria dispar asiatica (Lepidoptera: Erebidae) and L. dispar japonica] Trap Data From East Asian Ports With Lab Parameterized Phenology Models: New Tools and Questions. Annals of the Entomological Society of America. 113(2): 125-138. https://doi.org/10.1093/aesa/saz037.
Ward, Samuel F.; Fei, Songlin; Liebhold, Andrew M. 2020. Temporal dynamics and drivers of landscape⿐level spread by emerald ash borer. Journal of Applied Ecology. 57(6): 1020-1030. https://doi.org/10.1111/1365-2664.13613.
Ward, Samuel F.; Liebhold, Andrew M.; Morin, Randall S.; Fei, Songlin. 2021. Population dynamics of ash across the eastern USA following invasion by emerald ash borer. Forest Ecology and Management. 479: 118574. 8 p. https://doi.org/10.1016/j.foreco.2020.118574.
Büntgen, Ulf; Liebhold, Andrew; Nievergelt, Daniel; Wermelinger, Beat; Roques, Alain; Reinig, Frederick; Krusic, Paul J.; Piermattei, Alma; Egli, Simon; Cherubini, Paolo; Esper, Jan. 2020. Return of the moth: rethinking the effect of climate on insect outbreaks. Oecologia. 192(2): 543-552. https://doi.org/10.1007/s00442-019-04585-9.
Coleman, Kimberly J.; Perry, Elizabeth E.; Thom, Dominik; Gladkikh, Tatiana M.; Keeton, William S.; Clark, Peter W.; Tursini, Ralph E.; Wallin, Kimberly F. 2020. The Woods around the Ivory Tower: A Systematic Review Examining the Value and Relevance of School Forests in the United States. Sustainability. 12(2): 531. 16 p. https://doi.org/10.3390/su12020531.
Coleman, Tom W.; Haavik, Laurel J.; Foelker, Chris; Liebhold, Andrew M. 2020. Gypsy moth. Forest Insect and Disease Leaflet 162. U.S. Department of Agriculture, Forest Service. 20 p
Essl, Franz; Lenzner, Bernd; Bacher, Sven; Bailey, Sarah; Capinha, Cesar; Daehler, Curtis; Dullinger, Stefan; Genovesi, Piero; Hui, Cang; Hulme, Philip E.; Jeschke, Jonathan M.; Katsanevakis, Stelios; Kühn, Ingolf; Leung, Brian; Liebhold, Andrew; Liu, Chunlong; MacIsaac, Hugh J.; Meyerson, Laura A.; Nuñez, Martin A.; Pauchard, Aníbal; Pyšek, Petr; Rabitsch, Wolfgang; Richardson, David M.; Roy, Helen E.; Ruiz, Gregory M.; Russell, James C.; Sanders, Nathan J.; Sax, Dov F.; Scalera, Riccardo; Seebens, Hanno; Springborn, Michael; Turbelin, Anna; Kleunen, Mark; Holle, Betsy; Winter, Marten; Zenni, Rafael D.; Mattsson, Brady J.; Roura⿐Pascual, Nuria. 2020. Drivers of future alien species impacts: An expert⿐based assessment. Global Change Biology. 26(9): 4880-4893. https://doi.org/10.1111/gcb.15199.
Golivets, Marina; Woodall, Christopher W.; Wallin, Kimberly F. 2019. Functional form and interactions of the drivers of understory non⿐native plant invasions in northern US forests. Journal of Applied Ecology. 56(12): 2596-2608. https://doi.org/10.1111/1365-2664.13504.
Hudgins, Emma J.; Liebhold, Andrew M.; Leung, Brian. 2020. Comparing generalized and customized spread models for nonnative forest pests. Ecological Applications. 30(1): e01988. 13 p. https://doi.org/10.1002/eap.1988.
Andersen, Jeremy C.; Havill, Nathan P.; Mannai, Yaussra; Ezzine, Olfa; Dhahri, Samir; Ben Jamâa, Mohamed Lahbib; Caccone, Adalgisa; Elkinton, Joseph S. 2019. Identification of winter moth (Operophtera brumata) refugia in North Africa and the Italian Peninsula during the last glacial maximum. Ecology and Evolution. 9(24): 13931-13941. https://doi.org/10.1002/ece3.5830.
Blackburn, Gwylim S; Bilodeau, Pierre; Cooke, Tracey; Cui, Mingming; Cusson, Michel; Hamelin, Richard C; Keena, Melody A; Picq, Sandrine; Roe, Amanda D; Shi, Juan; Wu, Yunke; Porth, Ilga. 2020. An Applied Empirical Framework for Invasion Science: Confronting Biological Invasion Through Collaborative Research Aimed at Tool Production. Annals of the Entomological Society of America. 113(4): 230-245. https://doi.org/10.1093/aesa/saz072.
Kawatsu, Kazutaka; Yamanaka, Takehiko; Patoèka, Jan; Liebhold, Andrew M. 2020. Nonlinear time series analysis unravels underlying mechanisms of interspecific synchrony among foliage⿐feeding forest Lepidoptera species. Population Ecology. 62(1): 5-14. https://doi.org/10.1002/1438-390X.12025.
Keena, Melody A.; Richards, Jessica Y. 2020. Comparison of Survival and Development of Gypsy Moth Lymantria dispar L. (Lepidoptera: Erebidae) Populations from Different Geographic Areas on North American Conifers. Insects. 11(4): 260. 17 p. https://doi.org/10.3390/insects11040260.
Lantschner, M. Victoria; Corley, Juan C.; Liebhold, Andrew M. 2020. Drivers of global Scolytinae invasion patterns. Ecological Applications. 30(5): e02103. 12 p. https://doi.org/10.1002/eap.2103.
Liebhold, Andrew M.; Björkman, Christer; Roques, Alain; Bjørnstad, Ottar N.; Klapwijk, Maartje J. 2020. Outbreaking forest insect drives phase synchrony among sympatric folivores: Exploring potential mechanisms. Population Ecology. 13 p. https://doi.org/10.1002/1438-390X.12060.

Progress 10/01/18 to 09/30/19

Outputs
OUTPUTS: Outputs: 19 Publications. Research in Problem 1 resulted in the publication of 19 papers, of which 14 were in refereed journals and 1 was a scientific proceedings. Much of the research in this problem focused on understanding the global drivers, dynamics, and ecological-scale impacts of invasions. Research evaluated biomass losses resulting from insect and disease invasions in US Forests, spatio-temporal variation in spread of invasions, impacts of tree diversity in regulating pest invasions, drivers of spread of vector-borne tree disease, and how canopy gaps drive ground dwelling invertebrate community structure and function. Our research also investigated the biology and ecology of particular invasive species including European and Asian gypsy moth (Lymantria dispar), Asian longhorned beetle (Anoplophora glabripennis), hemlock woolly adelgid (Adelges tsugae), oak gall wasp (Zapatella davisae), and emerald ash borer (Agrilus planipennis) and other related Agrilus species. Papers were published on genomics of oak gall wasp, life history and mortality factors of apple buprestid, temperature models for Gypsy moth development and phenology models for egg hatch, influence of birds in dispersal of hemlock woolly adelgid, and factors impacting historical increases in two-line chestnut borer and bronze birch borer populations. PARTICIPANTS: Individuals who worked on Problem 1 included the investigators, Robert Haack, Nathan Havill, Melody Keena, Andrew Liebhold, Therese Poland, Toby Petrice, and Kimberly Wallin. Partners and collaborators included scientists from many universities, federal and state agencies, and international organizations including China, Europe, and New Zealand. TARGET AUDIENCES: The target audiences other scientists, federal and state regulatory agencies, resource managers, policy makers, timber nursery and tree care industries, and communities. PROJECT MODIFICATIONS: No modifications occurred to this problem

Impacts
Exotic forest pests threaten North American forests and natural resources. Native trees lack co-evolved defense mechanisms and exotic pests often invade without their associated natural enemies. Interactions of invasive pests with native ecosystems are unknown. A greater understanding of the drivers, mechanisms, and impacts of invasions are critical for developing strategies to predict, prevent, and mitigate invasions. Understanding the biology of invasive species is critical for protecting native forests and natural resources. Research in Problem 1 addresses exploration of invasion ecology as well as identification and description of new invasive species, understanding their biology, behavior, ecology ecosystem interactions, economic and environmental impacts, and how quickly and how far they will spread. Results provide regulatory agencies the information necessary to make scientifically-based decisions about implementing quarantine regulations. The research is also important to resource managers, policy makers, industries, and communities. The benefits of the research include enhanced abilities to create resilient forest ecosystems and protect them from the impacts of invasive species.

Publications

Guo, Qinfeng; Fei, Songlin; Potter, Kevin M.; Liebhold, Andrew M.; Wen, Jun. 2019. Tree diversity regulates forest pest invasion. Proceedings of the National Academy of Sciences. 116(15): 7382-7386. https://doi.org/10.1073/PNAS.1821039116.
Haack, Robert A.; Petrice, Toby R. 2019. Historical Population Increases and Related Inciting Factors of Agrilus anxius, Agrilus bilineatus, and Agrilus granulatus liragus (Coleoptera: Buprestidae) in the Lake States (Michigan, Minnesota, and Wisconsin). The Great Lakes Entomologist. 52: 21-33.⿿
Haynes, Kyle J.; Walter, Jonathan A.; Liebhold, Andrew M. 2019. Population spatial synchrony enhanced by periodicity and low detuning with environmental forcing. Proceedings of the Royal Society B: Biological Sciences. 286(1903): 20182828-. 10 p. https://doi.org/10.1098/rspb.2018.2828.
Keena, Melody A; Shi, Juan. 2019. Effects of Temperature on First Instar Lymantria (Lepidoptera: Erebidae) Survival and Development With and Without Food. Environmental Entomology. 48(3): 655-666. https://doi.org/10.1093/ee/nvz028.
Liebhold, Andrew M. 2019. Air pollution as an experimental probe of insect population dynamics. Journal of Animal Ecology. 88(5): 662-664. https://doi.org/10.1111/1365-2656.12992.
Machta, Jonathan; Blackwood, Julie C.; Noble, Andrew; Liebhold, Andrew M.; Hastings, Alan. 2019. A Hybrid Model for the Population Dynamics of Periodical Cicadas. Bulletin of Mathematical Biology. 81(4): 1122-1142. https://doi.org/10.1007/s11538-018-00554-0.
McManus, Katherine A., compiler. 2018. Proceedings, 29th USDA Interagency Research Forum on Invasive Species. Annapolis, MD; January 9-12, 2018. U.S. Department of Agriculture, Forest Service, Forest Health Technology Enterprise Team FHTET-2018-01.
Osada, Yutaka; Yamakita, Takehisa; Shoda-Kagaya, Etsuko; Liebhold, Andrew M.; Yamanaka, Takehiko. 2018. Disentangling the drivers of invasion spread in a vector-borne tree disease. Journal of Animal Ecology. 87(6): 1512-1524. https://doi.org/10.1111/1365-2656.12884.
Perry, Kayla I.; Wallin, Kimberly F.; Wenzel, John W.; Herms, Daniel A. 2018. Forest disturbance and arthropods: Small-scale canopy gaps drive invertebrate community structure and composition. Ecosphere. 9(10): e02463. 19 p. https://doi.org/10.1002/ecs2.2463.
Redwood, Mame E.; Matlack, Glenn R.; Huebner, Cynthia D. 2019. Seed longevity and dormancy state in an invasive tree species: Ailanthus altissima (Simaroubaceae). The Journal of the Torrey Botanical Society. 146(2): 79-86. https://doi.org/10.3159/Torrey-d-18-00038.1.
Russo, Nicholas J.; Elphick, Chris S.; Havill, Nathan P.; Tingley, Morgan W. 2019. Spring bird migration as a dispersal mechanism for the hemlock woolly adelgid. Biological Invasions. 21(5): 1585-1599. https://doi.org/10.1007/s10530-019-01918-w.
Yamaguchi, Ryo; Yamanaka, Takehiko; Liebhold, Andrew M. 2019. Consequences of hybridization during invasion on establishment success. Theoretical Ecology. 12(2): 197-205. https://doi.org/10.1007/s12080-019-0415-6.
Andersen, J.C.; Camp, C.P.; Davis, M.J.; Havill, N.P.; Elkinton, J.S. 2019. Development of microsatellite markers for an outbreaking species of oak gall wasp, Zapatella davisae (Hymenoptera: Cynipidae), in the northeastern United States. Bulletin of Entomological Research. 109(4): 510-517. https://doi.org/10.1017/S0007485318000858.
Blackwood, Julie C.; Machta, Jonathan; Meyer, Alexander D.; Noble, Andrew E.; Hastings, Alan; Liebhold, Andrew M. 2018. Competition and Stragglers as Mediators of Developmental Synchrony in Periodical Cicadas. The American Naturalist. 192(4): 479-489. https://doi.org/10.1086/699255.
Cui, Zhi-Jun; Zhang, Yan-Long; Zhang, Xin; Luo, Zhao-Hui; Zhang, Ping; Golec, Julian; Poland, Therese M.; Zalucki, Myron P.; Han, Peng; Lu, Zhao-Zhi. 2019. Life history and mortality factors of Agrilus mali Matsumura (Coleoptera: Buprestidae) in wild apples in Northwestern China . Agricultural and Forest Entomology. 21(3): 309-317. https://doi.org/10.1111/afe.12333.
Fei, Songlin; Morin, Randall S.; Oswalt, Christopher M.; Liebhold, Andrew M. 2019. Biomass losses resulting from insect and disease invasions in US forests. Proceedings of the National Academy of Sciences: 201820601. 6 p. https://doi.org/10.1073/pnas.1820601116.
Gippet, Jérôme M.W.; Liebhold, Andrew M.; Fenn-Moltu, Gyda; Bertelsmeier, Cleo. 2019. Human-mediated dispersal in insects. Current Opinion in Insect Science. 35: 96-102. https://doi.org/10.1016/j.cois.2019.07.005.
Goldstein, Joshua; Park, Jaewoo; Haran, Murali; Liebhold, Andrew; Bjørnstad, Ottar N. 2019. Quantifying spatio-temporal variation of invasion spread. Proceedings of the Royal Society B: Biological Sciences. 286(1894): 20182294-. https://doi.org/10.1098/rspb.2018.2294.
Gray, David R; Keena, Melody A. 2019. A Phenology Model for Asian Gypsy Moth Egg Hatch. Environmental Entomology. 48(4): 903-910. https://doi.org/10.1093/ee/nvz051.

Progress 10/01/16 to 09/30/17

Outputs
OUTPUTS: Research in Problem 1 resulted in the publication of 18 papers, of which 11 were in refereed journals, 4 were chapters in books, and 3 were in scientific proceedings or general technical reports. Much of the research in this problem focused the biology and ecology of particular invasive species including European and Asian gypsy moth (Lymantria dispar), Asian longhorned beetle (Anoplophora glabripennis), emerald ash borer (Agrilus planipennis) and hemlock woolly adelgid (Adelges tsugae). Papers were published on thermal requirements for gypsy moth egg hatch and larval development; on Asian longhorned beetle trail following behavior and temperature driven phenology models; on emerald ash borer impacts on eastern forests, woodpeckers, and nuthatches; and on the impacts of cold on hemlock woolly adelgid populations. Book chapters were written on general biology, feeding ecology, laboratory rearing, and urban and forest tree impacts of longhorned beetles. There was also considerable research on the general ecology, dynamics, and patterns of invasions. Papers were published on human history impacts on invasions and the geography of spatial synchrony of outbreaks. PARTICIPANTS: Individuals who worked on Problem 1 included the investigators Robert Haack, Nathan Havill, Melody Keena, Andrew Liebhold, and Talbot Trotter. Partners and collaborators included scientists from many universities, federal and state agencies, and international organizations including China, Europe, and New Zealand TARGET AUDIENCES: The target audience includes other scientists, federal and state regulatory agencies, resource managers, policy makers, timber nursery and tree care industries, and communities. PROJECT MODIFICATIONS: No modifications occurred to this problem

Impacts
Exotic forest pests threaten North American forests and natural resources. Native trees lack co-evolved defense mechanisms and exotic pests often invade without their associated natural enemies. Interactions of invasive pests with native ecosystems are unknown. A greater understanding of the biology of invasive species is critical for protecting native forests and natural resources. Research in Problem 1 addresses the identification and description of new invasive species, understanding their biology, behavior, ecology ecosystem interactions, economic and environmental impact, and how quickly and how far they will spread. Results provide regulatory agencies the information necessary to make scientifically-based decisions about implementing quarantine regulations. The research is also important to resource managers, policy makers, industries, and communities. The benefits of the research include enhanced abilities to create resilient forest ecosystems and protect them from the impacts of invasive species

Publications

Bertelsmeier, Cleo; Ollier, Sébastien; Liebhold, Andrew; Keller, Laurent. 2017. Recent human history governs global ant invasion dynamics. Nature Ecology & Evolution. 1(7): 0184-. 8 p. https://doi.org/10.1038/s41559-017-0184.
Büntgen, Ulf; Greuter, Lucie; Bollmann, Kurt; Jenny, Hannes; Liebhold, Andrew; Galván, J. Diego; Stenseth, Nils C.; Andrew, Carrie; Mysterud, Atle 2017. Elevational range shifts in four mountain ungulate species from the Swiss Alps. Ecosphere. 8(4): e01761. 15 p. https://doi.org/10.1002/ecs2.1761.
Graves, Fern; Baker, Thomas C.; Zhang, Aijun; Keena, Melody; Hoover, Kelli 2016. Sensory aspects of trail-following behaviors in the Asian longhorned beetle, Anoplophora glabripennis. Journal of Insect Behavior. 29(6): 615-628.
Haack, Robert A. 2017. Cerambycid pests in forests and urban trees. Chapter 11. In: Wang, Q. Cerambycidae of the world: biology and pest management. Boca Raton, FL: CRC Press: 352-384.
Haack, Robert A. 2017. Feeding biology of cerambycids. Chapter 3. In: Wang, Qiao, ed. Cerambycidae of the world; biology and pest management. Boca Raton, FLP CRC Press: 105-124.
Haack, Robert A.; Keena, Melody A.; Eyre, Dominic. 2017. Life history and population dynamics of Cerambycidae. Chapter 2. In: Wang, Q. Cerambycidae of the world: biology and pest management. Boca Raton, FL: CRC Press: 71-103.
Kappel, Alexander P.; Trotter, R. Talbot; Keena, Melody A.; Rogan, John; Williams, Christopher A. 2017. Mapping of the Asian longhorned beetle's time to maturity and risk to invasion at contiguous United States extent. Biological Invasions. 19(7): 1999-2013. https://doi.org/10.1007/s10530-017-1398-0.
Keena, M.A. 2016. Inheritance and world variation in thermal requirements for egg hatch in Lymantria dispar (Lepidoptera: Erebidae). Environmental Entomology 45(1): 1-10.
Keena, Melody A. 2017. Laboratory rearing and handling of cerambycids. In: Wang, Qiao, ed. Cerambycidae of the world: biology and pest management. Boca Raton, FL: CRC Press: 253-289. Chapter 7.
Koenig, Walter D.; Liebhold, Andrew M. 2017. A decade of emerald ash borer effects on regional woodpecker and nuthatch populations. Biological Invasions. 19(7): 2029-2037. https://doi.org/10.1007/s10530-017-1411-7.
Limbu, Samita; Keena, Melody; Chen, Fang; Cook, Gericke; Nadel, Hannah; Hoover, Kelli 2017. Effects of temperature on development of Lymantria dispar asiatica and Lymantria dispar japonica (Lepidoptera: Erebidae). Environmental Entomology. 46(4): 1012-1023. https://doi.org/10.1093/ee/nvx111.
McManus, Katherine A., comp. 2016. 27th USDA interagency research forum on Invasive species; 2016 January 12-15; Annapolis, MD. FHTET-2016-09. Morgantown, WV: U.S. Department of Agriculture, Forest Service, Forest Health Technology Enterprise Team. 107 p.
Morin, Randall S.; Liebhold, Andrew M.; Pugh, Scott A.; Crocker, Susan J. 2017. Regional assessment of emerald ash borer, Agrilus planipennis, impacts in forests of the Eastern United States. Biological Invasions. 19: 703-711. doi:10.1007/s10530-016-1296-x
Tobin, Patrick C.; Turcotte, Richard M.; Blackburn, Laura M.; Juracko, John A.; Simpson, Brian T. 2017. The big chill: quantifying the effect of the 2014 North American cold wave on hemlock woolly adelgid populations in the central Appalachian Mountains. Population Ecology. 59(3): 251-258. https://doi.org/10.1007/s10144-017-0589-y.
Trotter, R. Talbot; Keena, Melody A. 2016. A variable-instar climate-driven individual beetle-based phenology model for the invasive Asian longhorned beetle (Coleoptera: Cerambycidae). Environmental Entomology. 45(6): 1360-1370.
Walter, Jonathan A.; Sheppard, Lawrence W.; Anderson, Thomas L.; Kastens, Jude H.; Bjørnstad, Ottar N.; Liebhold, Andrew M.; Reuman, Daniel C.; Blasius, Bernd. 2017. The geography of spatial synchrony. Ecology Letters. 20(7): 801-814. https://doi.org/10.1111/ele.12782.
von Dohlen, Carol D.; Spaulding, Usha; Patch, Kistie B.; Weglarz, Kathryn M.; Foottit, Robert G.; Havill, Nathan P.; Burke, Gaelen R. 2017. Dynamic Acquisition and Loss of Dual-Obligate Symbionts in the Plant-Sap-Feeding Adelgidae (Hemiptera: Sternorrhyncha: Aphidoidea). Frontiers in Microbiology. 8: 439-. 15 p. https://doi.org/10.3389/fmicb.2017.01037.
⿿Kauffman, Bruce W.; Clatterbuck, Wayne K.; Liebhold, Andrew M.; Coyle, David R. 2017. Gypsy moth in the southeastern U.S.: Biology, ecology, and forest management strategies. SREF-FH-008. Athens, GA: Southern Regional Extension Forestry. 10 p.

Progress 10/01/15 to 09/30/16

Outputs
OUTPUT: Research in Problem 1 resulted in the publication of 19 papers, of which 16 were in refereed journals, 2 scientific proceedings, and 1 in a scientific newsletter. Much of the research in this problem focused the biology and ecology of particular invasive species including gypsy moth (Lymantria dispar), Asian longhorned beetle (Anoplophora glabripennis), and hemlock woolly adelgid (Adelges tsugae). Papers were published on genetic variation in gypsy moth populations, thermal requirements for gypsy moth egg hatch, and development of gypsy moth and defoliation on radiata pine in Spain; on Asian longhorned beetle dispersal in North America and on its life history, development, and natural enemy abundance at two sites in China; on hemlock woolly adelgid impacts on forest decline and ecological proceses, and on the life history patterns of longhorned beetles and outbreak dynamics of bark beetles. There was also considerable research on the general ecology, dynamics, and patterns of invasions, Papers were published on climate and weather effects spatial distribution and population dynamics of invasives, and on temporal and interspecific variation in rates of spread of invasive species and speed of invasions. PARTICIPANTS: Individuals who worked on Problem 1 included the investigators, Leah Bauer, Mary Ann Fajvan, Robert Haack, Nathan Havill, Cynthia Huebner, Melody Keena, Andrew Liebhold, Therese Poland, and Talbot Trotter. Partners and collaborators included scientists from many universities, federal and state agencies, and international organizations including China, Europe, and New Zealand. TARGET AUDIENCES: The target audiences other scientists, federal and state regulatory agencies, resource managers, policy makers, timber nursery and tree care industries, and communities. PROJECT MODIFICATIONS: No modifications occurred to this problem.

Impacts
Exotic forest pests threaten North American forests and natural resources. Native trees lack co-evolved defense mechanisms and exotic pests often invade without their associated natural enemies. Interactions of invasive pests with native ecosystems are unknown. A greater understanding of the biology of invasive species is critical for protecting native forests and natural resources. Research in Problem 1 addresses the identification and description of new invasive species, understanding their biology, behavior, ecology ecosystem interactions, economic and environmental impact, and how quickly and how far they will spread. Results provide regulatory agencies the information necessary to make scientifically-based decisions about implementing quarantine regulations. The research is also important to resource managers, policy makers, industries, and communities. The benefits of the research include enhanced abilities to create resilient forest ecosystems and protect them from the impacts of invasive species.

Publications

Tisseuil, Clement; Gryspeirt, Aiko; Lancelot, Renaud; Pioz, Maryline; Liebhold, Andrew; Gilbert, Marius. 2016. Evaluating methods to quantify spatial variation in the velocity of biological invasions. Ecography. 39(5): 409-418.
Trotter, R. Talbot; Hull-Sanders, Helen M. 2015. Quantifying Dispersal of the Asian longhorned beetle (Anoplophora glabripennis, Coleoptera) with incomplete data and behavioral knowledge. Biological Invasions. 17(12): 3359-3369.
van Driesche, R.G.; Pratt, P.D.; Center, T.D.; Rayamajhi, M.B.; Tipping, P.W.; Purcell, M.; Fowler, S.; Causton, C.; Hoddle, M.S.; Kaufman, L.; Messing, R.H.; Mongtomery, M.E.; van Klinken, R.; Duan, J.J.; Meyer, J-Y. 2016. 10 cases of biological control restoring natural systems. p. 208-246, Chp. 10, in van Driesche, R.G.; Simberloff, D.; Blossey, B.; Causton, C.; Hoddle, M.; Marks, C.O.; Heinz, K.M.; Wagner, D.L.; Warner, K.D., eds. Integrating biological control into conservation practices. Wiley-Blackwell. 360 p.
Keena, M.A. 2016. Inheritance and world variation in thermal requirements for egg hatch in Lymantria dispar (Lepidoptera: Erebidae). Environmental Entomology 45(1): 1-10
Koenig, Walter D.; Liebhold, Andrew M. 2016. Temporally increasing spatial synchrony of North American temperature and bird populations. Nature Climate Change, Vol. 6(6): 4 pages.: 614-617.
Kſivan, V., Lewis, M., Bentz, B.J., Bewick, S., Lenhart, S.M. and Liebhold, A., 2016. A dynamical model for bark beetle outbreaks. Journal of Theoretical Biology, 407: 25-37.
Lovett, G.M.; Weiss, M.; Liebhold, A.M.; Holmes, T.P.; Leung, B.; Lambert, K.F.; Orwig, D.A.; Campbell, F.T.; Rosenthal, J.; McCullough, D.G.; Wildova, R.; Ayres, M.P.; Canham, C.D.; Foster, D.R.; LaDeau, S.L.; Weldy, T. 2016. Nonnative forest insects and pathogens in the United States: Impacts and policy options. Ecological Applications 26(5): 1437-1455
Morin, R.S. and A.M. Liebhold. 2016. Invasive forest defoliator contributes to the impending downward trend of oak dominance in eastern North America. Forestry 89: 284-289. doi: 10.1093/forestry/cpv053
Redwood, Mame E.; Matlack, Glenn R.; Huebner, Cynthia D. 2016. Seed longevity and dormancy state in a disturbance-dependent forest herb, Ageratina altissima. Seed Science Research 26, 148⿿152.
Roques, Alain; Auger-Rozenberg, Marie-Anne; Blackburn, Tim M.; Garnas, Jeff; Pysek, Petr; Rabitsch, Wolfgang; Richardson, David M.; Wingfield, Michael J.; Liebhold, Andrew M.; Duncan, Richard P. 2016. Temporal and interspecific variation in rates of spread for insect species invading Europe during the last 200 years. Biological Invasions. 18(4): 907-920.
Allstadt, A.J., A. M. Liebhold, D. M. Johnson, R. E. Davis, and K. J. Haynes. 2015. Temporal variation in the synchrony of weather and its consequences for spatiotemporal population dynamics. Ecology, 96: 2935⿿2946.
Castedo-Dorado, F., G. Lago-Parra M.J. Lombardero, A.M. Liebhold and M.F. Álvarez-Taboada. 2016. European gypsy moth (Lymantria dispar dispar L.) completes development and defoliates exotic radiata pine plantations in Spain. New Zealand Journal of Forestry Science 46:18.
Chen, F.; Shi, J.; Keena, M. 2016. Evaluation of the effects of light intensity and time interval after the start of scotophase on the female flight propensity of Asian gypsy moth (Lepidoptera: Erebidae). Environmental Entomology 45(2): 404-409.
Culley, Theresa M.; Huebner, Cynthia D.; Novy, Ari. 2016. Regional and Local Genetic Variation in Japanese Stiltgrass (Microstegium vimineum). Invasive Plant Science and Management, 9(2):96-111
Fajvan, M.A., R.S. Morin and T. Lister. 2015. Assessment of the ecohydrological role of eastern hemlock (Tsuga canadensis) and its pending decline in Chesapeake Bay watershed. IN: Proceedings of the Geological Society of America Annual Meeting, Baltimore MD, November 1-4, 2015. Paper No.288-16. (Abstract) https://gsa.confex.com/gse/2015AM/webprogram/Paper266088.html In: Potter, Kevin M.; Conkling, Barbara L., eds. 2012. Forest health monitoring: 2009 national technical report. Gen. Tech. Rep. SRS-167. Asheville, NC: U.S. Department of Agriculture Forest Service, Southern Research Station. 211-217.
Fajvan, M.A., and R.S. Morin. 2016. Monitoring the effects of an invasive insect (Adelges tsugae) on forest decline and landscape-level hydrologic processes. IN: Proceedings of the Geological Society of America Annual Meeting, Denver, CO September 24-28, 2016. Paper No.310-10. (Abstract) https://gsa.confex.com/gsa/2016AM/webprogram/Paper283319.html
Favret, C.; Havill, N.P.; Miller, G.L.; Sano, M.; Victor, B. 2015. Catalog of the adelgids of the world (Hemiptera, Adelgidae). ZooKeys 534: 35-54. doi: 10.3897/zookeys.534.6456
Haack, R.A. 2015. Life history patterns of longhorned beetles (Cerambycidae): A worldwide perspective. Newsletter of the Michigan Entomological Society 60(3&4): 31.
Havill, N.P.; Shiyake, S.; Galloway, A.L.; Foottit, R.G.; Yu, G.; Paradis, A.; Elkinton, J.; Montgomery, M.E.; Sano, M.; Caccone, A. 2016. Ancient and modern colonization of North America by hemlock woolly adelgid, Adelges tsugae (Hemiptera: Adelgidae), an invasive insect from East Asia. Molecular Ecology 25: 2065-2080.

Progress 10/01/14 to 09/30/15

Outputs
OUTPUTS: Research in Problem 1 resulted in the publication of 42 papers in refereed journals, as chapters in books, or scientific proceedings in GTRs. Much of the research in this problem focused on the biology and ecology of particular invasive species including gypsy moth (Lymantria dispar), emerald ash borer (Agrilus planipennis), Asian longhorned beetle (Anoplophora glabripennis), hemlock woolly adelgid (Adelges tsugae), and several others. Papers were published on genetic variation in gypsy moth populations, gypsy moth spread rates, critical temperature limits, population synchronization, and wing morphometrics; on emerald ash borer biology and invasion history, antennal genes and gene knockdown, phenology and development on artificial diet, life history, mating behavior and host selection; on Asian longhorned beetle biology, dispersal, and landscape risk; and on hemlock woolly adelgid biology, ancient and modern colonization, stylet morphology, and enzymes. There was also considerable research on ecological processes and drivers of invasive species establishment, spread, and impact. Papers were published on Allee effects and invasive species establishment, drivers of macroscale forest invasions, impacts of invasive insects and diseases on regional forest dynamics, how invasion alters regional forest species composition and successional trajectories, and responses of an invasive plant to fire. Other papers considered global issues such as the impacts of globalization on forest health, comparison of insect invasions in North America and Japan, intercontinental ecological comparisons as a global tactic for managing invasives, and studies on a woodborer from the Bahamas and a bark beetle in China. PARTICIPANTS: Individuals who worked on Problem 1 included the investigators, Leah Bauer, Kurt Gottschalk, Robert Haack, Nathan Havill, Cynthia Huebner, Melody Keena, Andrew Liebhold, Kathy McManus, Toby Petrice, Therese Poland, Patrick Tobin, Talbot Trotter, and Kimberly Wallin. Partners and collaborators included scientists from many universities, federal and state agencies, and international organizations including Russia, New Zealand, and China. TARGET AUDIENCES: The target audiences include other scientists, federal and state regulatory agencies, resource managers, policy makers, timber nursery and tree care industries, and communities. PROJECT MODIFICATIONS: No modifications occurred to this problem

Impacts
Exotic forest pests threaten North American forests and natural resources. Native trees lack co-evolved defense mechanisms and exotic pests often invade without their associated natural enemies. Interactions of invasive pests with native ecosystems are unknown. A greater understanding of the biology of invasive species is critical for protecting native forests and natural resources. Research in Problem 1 addresses the identification and description of new invasive species, understanding their biology, behavior, ecology ecosystem interactions, economic and environmental impact, and how quickly and how far they will spread. Results provide regulatory agencies the information necessary to make scientifically-based decisions about implementing quarantine regulations. The research is also important to resource managers, policy makers, industries, and communities. The benefits of the research include enhanced abilities to identify high risk exotic pests and pathways.

Publications

Zhao, Chaoyang; Alvarez Gonzales, Miguel A.; Poland, Therese M.; Mittapalli, Omprakash. 2015. Core RNAi machinery and gene knockdown in the emerald ash borer (Agrilus planipennis). Journal of Insect Physiology. 72: 70-78.
Johnson, D.; Dattelbaum, K.; Tobin, P. 2014. Mechanisms of rapid synchronization in gypsy moth populations. in Parrotta, John A.; Moser, Cynthia F., Scherzer, Amy J.; Koerth, Nancy E.; Lederle, Daryl R., eds. , Sustaining forests, sustaining people: The role of research. XXIV IUFRO World Congress; 2014 Oct 5-11; Salt Lake City, UT. The International Forestry Review 16(5): 347. Abstract
Chase, K.; Brockerhoff, E.; Liebhold, A.; Kelly, D. 2014. Allee effects and the establishment of exotic invasive bark beetles. 2014, in Parrotta, John A.; Moser, Cynthia F., Scherzer, Amy J.; Koerth, Nancy E.; Lederle, Daryl R., eds. , Sustaining forests, sustaining people: The role of research. XXIV IUFRO World Congress; 2014 Oct 5-11; Salt Lake City, UT. The International Forestry Review 16(5): 354. Abstract
Chen, Fang; Keena, Melody; Shi, Juan. 2015. Genetic variation within gypsy moths from China and relatedness to gypsy moths from other world areas. In McManus, K.A. and Gottschalk, K.W., compilers, Proceedings, 26th USDA interagency research forum on invasive species; 2015 . 2015 Jan. 13-16; Annapolis, MD. FHTET-2015-09. Ft. Collins CO: U.S. Department of Agriculture, Forest Health Technology Enterprise Team: 60. Poster Abstract
Chen, Peng; Lu, Jun; Haack, Robert A.; Ye, Hui. 2015. Attack pattern and reproductive ecology of Tomicus brevipilosus (Coleoptera: Curculionidae) on Pinus yunnanensis in Southwestern China. Journal of Insect Science 15: 1-8.
Dattelbaum, K.; Parry, D.; Tobin, P.; Agosta, S.; Johnson, D. 2014. Critical thermal limits for the southern spread of the gypsy moth. in Parrotta, John A.; Moser, Cynthia F., Scherzer, Amy J.; Koerth, Nancy E.; Lederle, Daryl R., eds. , Sustaining forests, sustaining people: The role of research. XXIV IUFRO World Congress; 2014 Oct 5-11; Salt Lake City, UT. The International Forestry Review 16(5): 478. Abstract
Elkinton, J., G. Boettner, A. Liebhold, and R. Gwiazdowski. 2014. Biology, Spread and Biological Control of winter moth in the eastern United States. FHTET-2014-07.
Elkinton, Joseph S.; Liebhold, Andrew; Boettner, George H.; Sremac, Marinko.2014. Invasion spread of Operophtera brumata in northeastern United States and hybridization with O. bruceata. Biological Invasions (2014) 16:2263⿿2272.
Huebner, Cynthia D.; Karriker, Kent. 2015. Response of smooth rock skullcap (Scutellaria saxatilis), a globally rare plant, to fire. Res. Pap. NRS-28. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northern Research Station. 16 p.
Kappel, Alexander; Keena, Melody; Williams, Christopher; Trotter III, R. Talbot. 2015. Phenology/host based landscape risk for the Asian longhorned beetle in the conterminous U.S. In McManus, K.A. and Gottschalk, K.W., compilers, Proceedings, 26th USDA interagency research forum on invasive species; 2015 . 2015 Jan. 13-16; Annapolis, MD. FHTET-2015-09. Ft. Collins CO: U.S. Department of Agriculture, Forest Health Technology Enterprise Team: 75. Poster Abstract
Keena, Melody A.; Nadel, Hannah; Gould, Juli. 2015. Survival and phenology of Agrilus planipennis (Coleoptera: Buprestidae) reared on a newly developed artificial diet free of host material. Great Lakes Entomologist. 48(1-2): 9-33.
Keena, Melody A.; Shi, Juan. 2014. Effects of temperature on neonate Lymantria survival with and without food. McManus, Katherine A; Gottschalk, Kurt W., comps. Proceedings, 25th USDA interagency research forum on invasive species; 2014 . 2014 Jan. 7-10; Annapolis, MD. FHTET-2014-01. Ft. Collins CO: U.S. Department of Agriculture, Forest Health Technology Enterprise Team:: 70. Poster Abstract
Keena, Melody; Shi, Juan; Chen, Fang. 2015. Differences in wing morphometrics of Lymantria dispar between populations that vary in female flight capability. In McManus, K.A. and Gottschalk, K.W., compilers, Proceedings, 26th USDA interagency research forum on invasive species; 2015 . 2015 Jan. 13-16; Annapolis, MD. FHTET-2015-09. Ft. Collins CO: U.S. Department of Agriculture, Forest Health Technology Enterprise Team: 78. Poster Abstract
Haack, R. 2014. Invasive longhorned beetles (Cerambycidae) in North America: who, where, and what is being done? in Parrotta, John A.; Moser, Cynthia F., Scherzer, Amy J.; Koerth, Nancy E.; Lederle, Daryl R., eds. , Sustaining forests, sustaining people: The role of research. XXIV IUFRO World Congress; 2014 Oct 5-11; Salt Lake City, UT. The International Forestry Review 16(5): 331. Abstract
Haack, R.A.; Baranchikov, Y.; Bauer, L. S.; Poland, T.M. 2015. Emerald ash borer biology and invasion history. Pages 1-13 in ⿿Biology and Control of Emerald Ash Borer,⿝ edited by Roy G. Van Driesche and Richard C. Reardon. USDA Forest Service, Forest Health Technology Enterprise Team, Morgantown, WV, FHTET-2014-09.
Haack, Robert A.; Baranchikov, Yuri; Bauer, Leah S.; Poland, Therese M. 2015. Emerald ash borer biology and invasion history. In: Van Driesche, R.G.; Reardon, R.C., eds. Biology and control of emerald ash borer. FHTET-2014-09. Morgantown, WV: U.S. Department of Agriculture, Forest Service, Forest Health Technology Enterprise Team: 1-13. Chapter 1.
Haack, Robert A.; Sikken, Ariën; Petrice, Toby R.;Passoa, Steven C. 2014. A befuddling Bahamian borer. Newsletter of the Michigan Entomological Society 59(3-4): 31-32.
Havill, Nathan; Shiyake, Shigehiko; Galloway, Ashley; Foottit, Robert; Yu, Guoyue; Caccone, Adalgisa. 2015. Ancient and modern colonization by hemlock adelgids. In McManus, K.A. and Gottschalk, K.W., compilers, Proceedings, 26th USDA interagency research forum on invasive species; 2015 . 2015 Jan. 13-16; Annapolis, MD. FHTET-2015-09. Ft. Collins CO: U.S. Department of Agriculture, Forest Health Technology Enterprise Team: 69. Poster Abstract
Konter, Oliver; Esper, Jan; Liebhold, Andrew; Kyncl, Tomas; Schneider, Lea; Düthorn, Elisabeth; Buntgen, Ulf. 2015. Tree-ring evidence for the historical absence of cyclic larch budmoth outbreaks in the Tatra Mountains. Trees. 29(3): 809-814.
Krivak-Tetley, F.; Ayres, M. 2014. Intercontinental ecological comparisons as a global tactic for managing invasives. in Parrotta, John A.; Moser, Cynthia F., Scherzer, Amy J.; Koerth, Nancy E.; Lederle, Daryl R., eds. , Sustaining forests, sustaining people: The role of research. XXIV IUFRO World Congress; 2014 Oct 5-11; Salt Lake City, UT. The International Forestry Review 16(5): 332. Abstract
Liebhold, A. 2014. Contrasting geographical variation in invisibility to non-native forest pests with their impacts. in Parrotta, John A.; Moser, Cynthia F., Scherzer, Amy J.; Koerth, Nancy E.; Lederle, Daryl R., eds. , Sustaining forests, sustaining people: The role of research. XXIV IUFRO World
Liebhold, Andrew M. 2014. Responses to ⿿Clear, present, significant, & imminent danger: Questions for the California Light Brown Apple Moth (Epiphyas postvittana) Technical Working Group⿝. American Entomologist, Winter 2014 244-248
Liebhold, Andrew; Wingfield, Michael. 2014. Globalization and its implications for forest health. In: Nikolakis, William; Innes, John, eds. Forests and Globalization. Challenges and opportunities for sustainable development. New York, NY: Routledge: 36-47.
Liebhold, Andrew; Wingfield, Michael. 2014. Globalization and its implications to forest health. Chp. 4, p. 36-47 in Nikolakis, W. and Innes, John, eds. Forests and Globalization, Challenges and opportunities for sustainable development. Routledge.
McManus, Katherine A.; Gottschalk, Kurt W., comps. 2014. 25th U.S. Department of Agriculture interagency research forum on invasive species [proceedings]; 2014 Jan. 8-10; Annapolis MD. FHTET-2014-06. Ft. Collins CO: U.S. Department of Agriculture, Forest Health Technology Enterprise Team. 101 p.
McManus, Katherine A.; Gottschalk, Kurt W., comps. 2015. 26th USDA interagency research forum on invasive species; 2015 January 13-16; Annapolis, MD. FHTET-2015-09. Morgantown, WV: U.S. Department of Agriculture, Forest Service, Forest Health Technology Enterprise Team. 104 p.
Mech, A.; Gandhi, K.; Tobin, P.; Teskey, R.; Rhea, J. 2014. Influence of increasing summer temperatures on range dynamics of an invasive forest pest and its host tree. in Parrotta, John A.; Moser, Cynthia F., Scherzer, Amy J.; Koerth, Nancy E.; Lederle, Daryl R., eds. , Sustaining forests, sustaining people: The role of research. XXIV IUFRO World Congress; 2014 Oct 5-11; Salt Lake City, UT. The International Forestry Review 16(5): 352. Abstract
Morin, Randall S.; Liebhold, Andrew M. 2015. Invasions by two non-native insects alter regional forest species composition and successional trajectories. Forest Ecology and Management. 341: 67-74.
Meng, P.S.; Hoover, K.; Keena, M.A. 2015. Asian longhorned beetle (Coleoptera: Cerambycidae), an introduced pest of maple and other hardwood trees in North America and Europe. Journal of Integrated Pest Management. 6(1): 4-4.
Morin, R.; Liebhold, A. 2014. Invasions of forest insects and diseases and their impacts on regional forest dynamics in North America. in Parrotta, John A.; Moser, Cynthia F., Scherzer, Amy J.; Koerth, Nancy E.; Lederle, Daryl R., eds. , Sustaining forests, sustaining people: The role of research. XXIV IUFRO World Congress; 2014 Oct 5-11; Salt Lake City, UT. The International Forestry Review 16(5): 343. Abstract
Morin, Randall S.; Liebhold, Andrew M. 2015. Invasions by two non-native insects alter regional forest species composition and successional trajectories. In McManus, K.A. and Gottschalk, K.W., compilers, Proceedings, 26th USDA interagency research forum on invasive species; 2015 . 2015 Jan. 13-16; Annapolis, MD. FHTET-2015-09. Ft. Collins CO: U.S. Department of Agriculture, Forest Health Technology Enterprise Team: 39. Abstract
Oten, Kelly L.F.; Cohen, Allen C.; Hain, Fred P. 2014. Stylet bundle morphology and trophically related enzymes of the hemlock woolly adelgid (Hemiptera: Adelgidae). Annals of Entomological Society of America. 107(3): 680-690.
Perry, Kayla; Wallin, Kimberly F.; Herms, Daniel A. 2015. Effects of forest disturbance on ground-dwelling invertebrate dispersal. In McManus, K.A. and Gottschalk, K.W., compilers, Proceedings, 26th USDA interagency research forum on invasive species; 2015 . 2015 Jan. 13-16; Annapolis, MD. FHTET-2015-09. Ft. Collins CO: U.S. Department of Agriculture, Forest Health Technology Enterprise Team: 79. Poster Abstract
Petrice, Toby R.; Haack, Robert A. 2015. Studies on the European oak borer and other Agrilus in Michigan. 2015. In McManus, K.A. and Gottschalk, K.W., compilers, Proceedings, 26th USDA interagency research forum on invasive species; 2015 . 2015 Jan. 13-16; Annapolis, MD. FHTET-2015-09. Ft. Collins CO: U.S. Department of Agriculture, Forest Health Technology Enterprise Team: 87. Poster Abstract
Poland, Therese M.; Chen, Yigen; Koch, Jennifer; Pureswaran, Deepa. 2015. Review of the emerald ash borer (Coleoptera: Buprestidae), life history, mating behaviours, host plant selection, and host resistance. The Canadian Entomologist. 147(03): 252-262.
Shi, Juan; Chen, Fang; Keena, Melody A. 2015. Differences in wing morphometrics of Lymantria dispar (Lepidoptera: Erebidae) between populations that vary in female flight capability. Annals of the Entomological Society of America. 108(4): 528-535.
Tobin, P. Gray, D.; Liebhold, A. 2014. Supraoptimal temperatures restrict and retract the distribution range of the gypsy moth. in Parrotta, John A.; Moser, Cynthia F., Scherzer, Amy J.; Koerth, Nancy E.; Lederle, Daryl R., eds. , Sustaining forests, sustaining people: The role of research. XXIV IUFRO World Congress; 2014 Oct 5-11; Salt Lake City, UT. The International Forestry Review 16(5): 352. Abstract
Tobin, Patrick C.; Liebhold, Andrew M.; Roberts, E. Anderson; Blackburn, Laura M. 2015. Estimating spread rate of non-native species: The gypsy moth as a case study. Chp. 9, pp. 131-144 in Venette, R.C., ed. Pest Risk Modelling and Mapping for Invasive Alien Species. CAB International and U.S. Department of Agriculture.
Trotter III, R. Talbot; Hull-Sanders, Helen. 2015. Asian longhorned beetle dispersal in invaded landscapes. 2015. In McManus, K.A. and Gottschalk, K.W., compilers, Proceedings, 26th USDA interagency research forum on invasive species; 2015 . 2015 Jan. 13-16; Annapolis, MD. FHTET-2015-09. Ft. Collins CO: U.S. Department of Agriculture, Forest Health Technology Enterprise Team: 94. Poster Abstract
Yamanaka, Takehiko; Morimoto, Nobuo; Nishida, Gordon M.; Kirtani, Keizi; Moriya, Seiichi; Liebhold, Andrew M. 2015. Comparison of insect invasions in North America, Japan and their Islands. Biological Invasions 17: 3049-3061.

Progress 10/01/13 to 09/30/14

Outputs
OUTPUTS: The fungal pathogen Entomophaga maimaiga is prevalent in gypsy moth [Lymantria dispar (L.)] populations throughout North America. Fungal mortality was assessed using both field-collected larvae and laboratory-reared larvae caged on the forest floor. We found significant positive effects of moisture-related variables (rainfall, soil moisture, and relative humidity) on mortality due to fungal infection in both data sets, and significant negative effects of temperature on the mortality of field-collected larvae. These relationships may be helpful in understanding how gypsy moth dynamics vary across space and time, and in forecasting how the gypsy moth and fungus will interact as they move into warmer or drier areas, or new weather conditions occur due to climate change. We documented gypsy moth (GM) range expansion, stasis, and retraction across a fairly narrow latitudinal region which showed differences in the amount of exposure above the optimal temperature for GM larval and pupal development. Temperature regimes in the Coastal Plain and Piedmont regions of Virginia, where the GM range has retracted or remained static, were warmer than those in the Appalachian Mountains of Virginia and West Virginia, where GM has expanded its range. Our analyses at a smaller spatial scale confirmed a statistically negative association between exposure time above the optimal temperature for GM larvae and pupae, and the rate of GM invasion spread over the 20-year period. We isolated four chemicals from the trail of Asian longhorned beetle (ALB) virgin and mated females that were not present in trails of mature males. Males responded to the full pheromone blend, regardless of mating status, but virgin females chose the control over the pheromone, suggesting that they may use it as a spacing pheromone to avoid intraspecific competition and maximize resources. These results indicate that all four chemicals are components of the trail pheromone. Over two field seasons, a total of 160 flight intercept panel traps were deployed in Harbin, China, which trapped a total of 65 beetles. In 2012, traps using lures with a 1:1 ratio of the male-produced pheromone components designed to release at a rate of 1 or 4 milligram per day per component in conjunction with plant volatiles caught significantly more ALB females than other pheromone release rates, other pheromone ratios, plant volatiles only, and no lure controls. Males were caught primarily in traps baited with plant volatiles only. Significantly more females were trapped using the pheromone with the 10-fold higher three or four plant volatile release rates compared with the plant volatiles only, low four plant volatile pheromone, and control. Our findings show that the male-produced pheromone in combination with plant volatiles can be used to detect ALB. PARTICIPANTS: National Center for Ecological Analysis and Synthesis (NCEAS), Michigan State University, University of Massachusetts, Cornell University, Agriculture and AgriFood Canada, Natural Resources Canada ⿿ Canadian Forest Service, USDA Animal and Plant Health Inspection Service (APHIS), USDA Agricultural Research Service (ARS), Chinese Academy of Sciences, Connecticut Agricultural Experiment Station, Virginia Polytechnic University, USFS State & Private Forestry Forest Health Protection, Cornell University, University of Tennessee, Pennsylvania State University, Osaka Museum of Natural History, University of Copenhagen, Duzce University (Turkey), Abant I˿zzet Baysal University (Turkey), CABI Europe, Yunnan University (China), Nanjing Forestry University (China), Chinese Academy of Forestry, University of Idaho, University of Virginia, Universidad Nacional de Salta (Argentina), INTA EEA (Bariloche, Argentina), University of Padova (Italty), North Carolina State University, Swiss Federal Institute for Forest, Snow, and Landscape Research, University of Bern (Switzerland), University of Oslo (Norway), State University of New York College of Env.Sci.Forestry, University of Minnesota, TARGET AUDIENCES: Policymakers, scientists, pest managers, quarantine/trade managers, forest and land managers PROJECT MODIFICATIONS: none

Impacts
The impacts of invasive species on forests and urban trees are a major reason for managers' concern over their introduction. Our research into basic biological factors of invasive plants, insects, and diseases provides the information needed by other scientists and managers to develop new control and management treatments and tactics.

Publications

Battisti, Andrea; Benvegnu, I.; Colombari, F.; Haack, R.A. 2014. Invasion by the chestnut gall wasp in Italy causes significant yield loss in Castanea sativa nut production. Agricultural and Forest Entomology 16: 75-79.
Bigsby,Kevin M.; Ambrose, Mark J.; Tobin, Patrick C.; Sills, Erin O. 2014. The cost of gypsy moth sex in the city. Urban Forestry and Urban Greening 13(3):459-468.
Büntgen,Ulf; Liebhold, A.M.; Jenny, H.; Mysterud,A.; Egli,S.; Nievergelt, D.; Stenseth,N.C.; Bollman, K. 2014. European springtime temperature synchronizes ibex horn growth across the eastern Swiss Alps. Ecology Letters 17: 303-313.
Büntgen,Ulf; Tegel,W.; Kaplan,J.O.; Schaub,M.;Hagedorn, F.; Bürgi, M.;Brázdil, R.; Helle, G.; Carrer, M.; Heussner, K-U.; Hoffman,J.; Kontic, R.; Kyncl, T.; Kyncl, J.; Camarero, J.J.; Tinner, W.; Esper, J.; Liebhold, A. 2014. Placing unprecedented recent fir growth in a European-wide and Holocene-long context. Frontiers in Ecology 12: 100-106.
Harrison, Robert L.; Keena, Melody A.; Rowley, Daniel L. 2014. Classification, genetic variation and pathogenicity of Lymantria dispar nucleopolyhedrovirus isolates from Asia, Europe, and North America. Journal of Invertebrate Pathology. 116: 27-35.
Havill, Nathan P.; Vieira, Ligia C.; Salom, Scott M. 2014. Hemlock woolly adelgid: a threat to eastern forests. National Woodlands. 37(2): 14-15.
Petrice, Toby R.; Haack, Robert A. 2014. Biology of the European oak borer in Michigan, United States of America, with comparisons to the native twolined chestnut borer. The Canadian Entomologist. 146(01): 36-51.
Reilly, James R.; Hajek, Ann E.; Liebhold, Andrew M.; Plymale, Ruth. 2014. Impact of Entomophaga maimaiga (Entomophthorales: Entomophthoraceae) on outbreak gypsy moth populations (Lepidoptera: Erebidae): The role of weather. Environmental Entomology 43(3): 632-641.
Hoover, Kelli; Keena, Melody; Nehme, Maya; Wang, Shifa; Meng, Peter; Zhang, Aijun. 2014. Sex-specific trail pheromone mediates complex mate finding behavior in Anoplophora glabripennis. Journal of Chemical Ecology. 40(2): 169-180.
Huebner, Cynthia D.; Steinman, Jim; Hutchinson, Todd F.; Ristau, Todd E.; Royo, Alejandro A. 2014. The distribution of a non-native (Rosa multiflora) and native (Kalmia latifolia) shrub in mature closed-canopy forests across soil fertility gradients. Plant and Soil. 377(1-2): 259-276.
Meng, P.S.; Trotter, R.T.; Keena, M.A.; Baker, T.C.; Yan, S.; Schwartzberg, E.G.; Hoover, K. 2014. Effects of pheromone and plant volatile release rates and ratios on trapping Anoplophora glabripennis (Coleoptera: Cerambycidae) in China. Environmental Entomology. 43(5): 1379-1388.
Morin, Randall; Liebhold, Andrew. 2014. Impacts of Forest Pests on Regional Forest Dynamics in the Eastern US. Journal of Forestry January 2014, 2013 Society of American Foresters National Convention, 117. Presentation Abstract
Siegert, Nathan W.; McCullough, Deborah G.; Liebhold, Andrew M.; Telewski, Frank W. 2014. Dendrochronological reconstruction of the epicenter and early spread of emerald ash borer in North America. Diversity and Distributions 20: 847-858.
Tobin, P.C.; Bolyard, J.L.; Onufrieva, K.S.; Hickman, A.D. 2014. The effect of male and female age on Lymantria dispar (Lepidoptera: Lymantriidae) fecundity. Journal of Economic Entomology 107: 1076-1083.
Tobin, P.C.; Parry, D.; Aukema, B.H. 2014. The influence of climate change on insect invasions in temperate forest ecosystems. Book chapter, Challenges and Opportunities for the World⿿s Forests in the 21st Century. T. Fleming (ed.) Springer, pp 267-296.
Tobin, Patrick C.; Gray, David R.; Liebhold, Andrew M. 2014. Supraoptimal temperatures influence the range dynamics of a non-native insect. Diversity and Distributions 20: 813-823.

Progress 10/01/12 to 09/30/13

Outputs
OUTPUTS: Population cycles in forest Lepidoptera often result from recurring density dependent epizootics of entomopathogens. While these systems are typically dominated by a single pathogen species, insects are often infected by multiple pathogens, yet little is known how pathogens interact to affect host dynamics. The apparent invasion of northeastern North America by Entomophaga maimaiga some time prior to 1989 provides a unique opportunity to evaluate such interactions. Prior to the arrival of E. maimaga, the oscillatory dynamics of host gypsy moth, Lymantria dispar, populations were apparently driven by epizootics of a nucleopolyhedrovirus. Subsequent to its emergence, E. maimaiga has caused extensive mortality in host populations, but little is known about how it has altered multigenerational dynamics of the gypsy moth and its virus. Here we compared demographic data collected in gypsy moth populations prior to vs. after E. maimaiga⿿s invasion. We found that the recently invading fungal pathogen virtually always causes greater levels of mortality in hosts than does the virus, but fungal mortality is largely density independent. Moreover, the presence of the fungus has apparently not altered the gypsy moth⿿virus density-dependent interactions that were shown to drive periodic oscillations in hosts before the arrival of the fungus. To understand what determines outbreaks dynamics in the woodwasp, Sirex noctilio, we developed an individual-based model for a wasp population developing within a pine plantation. We show that outbreaks may be the result of the insect⿿s life history. Specifically we show that limited dispersal may not only increase population persistence but also create the conditions for eruptive dynamics. When the probability of long distance dispersal is greater than zero, but relatively small (PLDD= 0.1) large outbreaks are the norm, with all of the suitable trees dead at the end of the simulation. For PLDD= 0 (only local dispersal allowed) outbreaks are smaller in size, and in some cases not well defined and spread over longer periods. For PLDD= 1 (only long distance dispersal allowed), the frequency of local population extinction (without outbreaks) increases significantly. Aggregated attacks may induce physiological changes in the trees which could allow other wasps to detect them. These changes may in turn trigger an outbreak. In contrast, healthy, vigorous trees are not suitable for wasp oviposition. In our model the density of suitable trees (healthy trees but yet suitable for oviposition) are a key factor determining population persistence before outbreaks. From an applied perspective, our results emphasize the importance of adequate plantation management in preventing woodwasp infestation. PARTICIPANTS: National Center for Ecological Analysis and Synthesis (NCEAS), Michigan State University, University of Massachusetts, Cornell University, Agriculture and AgriFood Canada, USDA Animal and Plant Health Inspection Service (APHIS), USDA Agricultural Research Service (ARS), Chinese Academy of Sciences, Connecticut Agricultural Experiment Station, Virginia Polytechnic University, USFS State & Private Forestry Forest Health Protection, Cornell University, University of Tennessee, Osaka Museum of Natural History, University of Copenhagen, Duzce University (Turkey), Abant I˿zzet Baysal University (Turkey), CABI Europe, Yunnan University (China), Chinese Academy of Forestry, University of Idaho, University of Virginia, Universidad Nacional de Salta (Argentina), INTA EEA (Bariloche, Argentina) TARGET AUDIENCES: Policymakers, scientists, pest managers, quarantine/trade managers, forest and land managers

Impacts
The impacts of invasive species on forests and urban trees are a major reason for managers' concern over their introduction. Our research into basic biological factors of invasive plants, insects, and diseases provides the information needed by other scientists and managers to develop new control and management treatments and tactics.

Publications

Allstadt, A.J.; Haynes, K.J.; Liebhold, A.M.; ⿢Johnson, D.M. 2013. Long-term shifts in the cyclicity of outbreaks of a forest-defoliating insect. Oecologia 172: 141-151.
Aparicio, J.P.; Corley, J.C.; Rabinovich, J.E. 2013. Life history traits of Sirex noctilio F. (Hymenoptera: Siricidae) can explain outbreaks independently of environmental factors. Mathematical Biosciences and Engineering 10 1265-1279.
Britton, Kerry O.; Liebhold, Andrew M. 2013. One world, many pathogens! New Phytologist 197: 9-10. Commentary.
Chamorro,M. L.; Volkovitsh, M.G.; Haack, R.A.; Poland, T.M.; Lingafelter, S.W. 2013. An illustrated guide to the larva of Agrilus planipennis Fairmaire (emerald ash borer) (Coleoptera: Buprestidae). In McManus, K.A.; Gottschalk, K.W., eds. Proceedings. 23rd U.S. Department of Agriculture interagency research forum on invasive species 2012; 2012 January 10-13; Annapolis, MD. Gen. Tech. Rep. NRS-P-114. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northern Research Station: 74. Abstract
Chen, Yigen; Poland, Therese; Whitehill, Justin; Bonello, Pierluigi. 2011. Changes in ash foliar chemistry in response to EAB adult and larval feeding. p. 17-18 in Parra, Gregory; Lance, David; Mastro, Victor; Reardon, Richard; Benedict, Chuck, compilers. 2011 Emerald Ash Borer National Research and Technology Development Meeting: 2011 October 12-13; Wooster, OH. FHTET-2011-06. Abstract.
Cook, Stephen P.; Merickel, Frank; Ward, Karen; Havill, Nathan. 2012. First report of Adelges abietis (Linnaeus) (Hemiptera: Adelgidae) in Idaho. The Pan-Pacific Entomologist 88: 435-436. Scientific Note.
Cuddington, K.; Fortin, M.-J.; Gerber, L.R.; Hastings, A.; Liebhold, A.; OConnor, M.; Ray, C. 2013. Process-based models are required to manage ecological systems in a changing world. Ecosphere. 4(2). dx.doi.org/10.1890/ES12-00178.1
Donovan, Geoffrey H.; Butry, David T.; Michael, Yvonne L.; Prestemon, Jeffrey P.; Liebhold, Andrew M; Gatziolis, D.; Mao, M.Y. 2013. The relationship between trees and human health. American Journal of Preventive Medicine 44: 139-145.
Elkinton, J.S.; Liebhold, A.M.; Boettner, G.; Luzader, E. 2013. Spread of winter moth in the northeastern United States. In McManus, K.A.; Gottschalk, K.W. compilers. 2013. Proceedings. 24th U.S. Department of Agriculture interagency research forum on invasive species 2013; 2013 January 8-11. Annapolis, MD. FHTET-13-01. Fort Collins, CO: U.S. Department of Agriculture, Forest Service: 67. Abstract
Fischer, M.J.; Havill, N.P.; Davis, G.A. 2013. Evaluation of hybridization among three Laricobius spp. predators of hemlock woolly adelgid (Adelgidae): Preliminary results. In McManus, K.A.; Gottschalk, K.W., eds. 2012 January 10-13; Annapolis, MD. Gen. Tech. Rep. NRS-P-114. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northern Research Station. 19-20. Abstract
D⿿Amico, V.; Havill, N.P. 2013. The life cycle of hemlock woolly adelgid: a case study for the graphical explanation of data. In McManus, K.A.; Gottschalk, K.W., eds. Proceedings. 23rd U.S. Department of Agriculture interagency research forum on invasive species 2012; 2012 January 10-13; Annapolis, MD. Gen. Tech. Rep. NRS-P-114. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northern Research Station: 79-80. Abstract
Frank, K.L.; Tobin, P.C.; Thistle, Jr., H.W.; Kalkstein, Laurence S. 2013. Interpretation of gypsy moth frontal advance using meteorology in a conditional algorithm. International Journal of Biometeorology 57: 459-473.
Gwiazdowski, R.A.; Elkinton, J.S.; Dewaard, J.R.; Sremac, M. 2013. Phylogeographic diversity of the winter moths Operophtera brumata and O. bruceata (Lepidoptera: Geometridae) in Europe and North America. Annals, Entomological Society of America 106(2): 143-151.
Haack, Robert A. 2012. Seasonality of oak twig pruner shoot fall: a long-term dog walking study. Newsletter of the Michigan Entomological Society 57: 24
Haack, Robert A.; Rabaglia, Robert J. 2013. Exotic bark and ambrosia beetles in the USA: Potential and current invaders. Chp. 3, p. 48-74 in J. Peña, ed. Potential invasive pests of agricultural crops. CAB International.
Havill, N.P.; Davis, Gina; Mausel, David; Klein, Joanne; McDonald, Richard; Jones, Cera; Fischer, Melissa; Salom, Scott; Caccone, Adelgisa. 2012. Hybridization between a native and introduced predator of Adelgidae: An unintended result of classical biological control. Biological Control 63: 359-369.
Havill, N.P.; Elkinton, J.S.; Keena, M.A.; Gloria-Soria, A.; Caccone, A. 2013. Population genomics of admixture in non-native forest pests: winter moth and gypsy moth. In McManus, K.A.; Gottschalk, K.W. compilers. 2013. Proceedings. 24th U.S. Department of Agriculture interagency research forum on invasive species 2013; 2013 January 8-11. Annapolis, MD. FHTET-13-01. Fort Collins, CO: U.S. Department of Agriculture, Forest Service: 77. Abstract
Haynes, K.J.; Bjørnstad, O.N.; Allstadt, A.J.; Liebhold, A.M. 2012. Geographical variation in the spatial synchrony of a forest-defoliating insect: isolation of environmental and spatial drivers. Proceedings of the Royal Society, B 280: 20122373
Hu, Shao-ji; Ning, Tiao; Fu, Da-ying; Haack, Robert A.; Zhang, Z.; Chen, D; Ma, X.; Ye, H. 2013. Dispersal of the Japanese pine sawyer, Monochamus alternatus (Coleoptera: Cerambycidae), in Mainland China as inferred from molecular data and associations to indices of human activity. PLOS ONE 8: e57568
Huebner, Cynthia D.; Nowak, David J.; Pouyat, Richard V.; Bodine, Allison R. 2012. Nonnative invasive plants: Maintaining biotic and soceioeconomic integrity along the urban-rural-natural gradient. In: Laband, D.N.; Lockaby, B.G.; Zipperer, W., eds. Urban-rural interfaces: linking people and nature. Madison, WI: American Society of Agronomy, Soil Science Society of America: 71-98.
Keena, M.A.; Bonello, P.; Nadel, H. 2013. An artificial diet that does not contain host material for rearing the emerald ash borer (Agrilus planipennis) larvae. In McManus, K.A.; Gottschalk, K.W., eds. Proceedings. 23rd U.S. Department of Agriculture interagency research forum on invasive species 2012; 2012 January 10-13; Annapolis, MD. Gen. Tech. Rep. NRS-P-114. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northern Research Station: 93. Abstract
Keena, Melody A.; Bonello, Pierluigi (Enrico); Nadel, Hannah. 2011. Improving the artificial diet that does not contain host material and methods developed for rearing the emerald ash borer. p. 68-69 in Parra, Gregory; Lance, David; Mastro, Victor; Reardon, Richard; Benedict, Chuck, compilers. 2011 Emerald Ash Borer National Research and Technology Development Meeting: 2011 October 12-13; Wooster, OH. FHTET-2011-06. Abstract.
Keena, Melody A.; Trotter III, R. Talbot; Cheah, Carole; Montgomery, Michael E. 2012. Effects of temperature and photoperiod on the aestivo-hibernal egg diapause of Scymnus camptodromus (Coleoptera: Coccinellidae). Environmental Entomology. 41(6): 1662-1671.
Keever, Carson C.; Nieman, Christal; Ramsay, Larissa; Ritland, Carol E.; Bauer, Leah S.; Lyons, D. Barry; Cory, Jenny S. 2013. Microsatellite population genetics of the emerald ash borer (Agrilus planipennis Fairmaire): comparisons between Asian and North American populations. Biological Invasions 15: 1537-1559.
Koenig, Walter D.; Liebhold, Andrew M.; Bonter, David N.; Hochachka, Wesley M.; Dickinson, Janis L. 2013. Effects of the emerald ash borer invasion on four species of birds. Biological Invasions. 15: 2095-2103.
Koenig, Walter E.; Liebhold, Andrew M. 2013. Avian predation pressure as a potential driver of periodical cicada cycle length. The American Naturalist. 181: 145-149.
LI Li, YANG Jia-Ni, CUI Kai, R. Talbot Trotter, LI Zheng-Hong, LI Gen-Qian, and LIAO Sheng-Xi. 2013. Strobili and seed characteristics of Tsuga dumosa and its relationship with environmental factors. Chinese Journal of Plant Ecology 37:820-829
Petrice, Toby R.; Haack, Robert A.; Chen, Peng. 2012. Host range studies on Agrilus and other wood borers in Southwest China. Newsletter of the Michigan Entomological Society 57: 25. Abstract
Liebhold, A.M.; Plymale, R.; Elkinton, J.S.; Hajek, A.E. 2013. How has the emergence of Entomophaga maimaiga altered the dynamics of North American gypsy moth populations? In McManus, K.A.; Gottschalk, K.W. compilers. 2013. Proceedings. 24th U.S. Department of Agriculture interagency research forum on invasive species 2013; 2013 January 8-11. Annapolis, MD. FHTET-13-01. Fort Collins, CO: U.S. Department of Agriculture, Forest Service: 35. Abstract
Liebhold, Andrew M.; Plymale, Ruth; Elkinton, Joseph S., Hajek, Ann E. 2013. Emergent fungal entomopathogen does not alter density dependence in a viral competitor. Ecology 94: 1217-1222.
Ravn, H.P.; Havill, N.P.; Akubulut, S.; Foottit, R.G.; Serin, M.; Erdem,M.; Mutun, S.; Kenis, M. 2013. Dreyfusia nordmannianae in Northern and Central Europe: potential for biological control and comments on its taxonomy. Journal of Applied Entomology 137: 401-417.
Reilly, J.R.; Liebhold, A.M.; Plymale, R.; Hajek, A.E. 2013. The impact of Entomphaga maimaiga on outbreak gypsy moth populations: the role of weather. In McManus, K.A.; Gottschalk, K.W. compilers. 2013. Proceedings. 24th U.S. Department of Agriculture interagency research forum on invasive species 2013; 2013 January 8-11. Annapolis, MD. FHTET-13-01. Fort Collins, CO: U.S. Department of Agriculture, Forest Service: 34. Abstract
Rutledge, Claire E.; Keena, Melody A. 2012. Mating frequency and fecundity in Agrilus anxius (Coleoptera: Buprestidae). Annals of the Entomological Society of America. 105(6): 852-858.
Sanchez, V.; Keena, M.A. 2013. Development of the teneral adult Anoplophora glabripennis (Coleoptera: Cerambycidae): time to initiate and completely bore out of maple wood. Environmental Entomology. 42(1): 1-6. dx.doi.org/10.1603/EN12225.
Tobin, P.C.; Turcotte, R.M.; Snider, D. 2013. Tobin PC, Turcotte RM, Snider DA. (2013) When one is not necessarily a lonely number: initial colonization dynamics of Adelges tsugae on eastern hemlock, Tsuga canadensis. Biological Invasions 15:1925-193
Tobin, Patrick C.; Hajek, Ann E. 2012. Release, establishment, and initial spread of the fungal pathogen Entomophaga maimaiga in island populations of Lymantria dispar. Biological Control 63: 31-39.
Tobin, Patrick C.; Onufrieva, Ksenia S.; Thorpe, Kevin W. 2012. The relationship between male moth density and female mating success in invading populations of Lymantria dispar. Entomologia Experimentalis et Applicata 146: 103-111.
Trotter, Talbot. 2013. Relationships among climate, forests, and insects in North America: Examples of the importance of long-term and broad-scale perspectives. In: Camp, Ann E.; Irland, Lloyd C.; Carroll, Charles J.W. Long-term silvicultural and ecological studies. Results for science and management: volume 2.GISF Res. Pap. 013. New Haven, CT: Yale University, School of Forestry and Environmental Studies, Global Institute of Sustainable Forestry: 161-177.
Von Dohlen, C. D.; Usha Spaulding, U.; Shields, K.; Havill, N.P.; Rosa, C; Hoover, K. 2013. Diversity of proteobacterial endosymbionts in hemlock woolly adelgid (Adelges tsugae) (Hemiptera: Adelgidae) from its native and introduced range. Environmental Microbiology 15(7), 2043⿿2062
Walter, J.A.; Meixler, M.S.; Mueller, T.; Fagan, W.; Tobin, P.C.; Haynes, K.J. 2013. Topography influences allee effects at the invasion front of the gypsy moth. In McManus, K.A.; Gottschalk, K.W. compilers. 2013. Proceedings. 24th U.S. Department of Agriculture interagency research forum on invasive species 2013; 2013 January 8-11. Annapolis, MD. FHTET-13-01. Fort Collins, CO: U.S. Department of Agriculture, Forest Service: 36. Abstract

Progress 10/01/11 to 09/30/12

Outputs
OUTPUTS: The emerald ash borer (EAB) (Agrilus planipennis Fairmaire) is a nonnative phloem-feeding beetle that was accidentally introduced near Detroit, Michigan, two to three decades ago. North American ash (Fraxinus spp.) exhibit little or no resistance, and as this insect species expands its range, extensive mortality results. Previous studies of the impacts of EAB, typical of most insect and disease impact studies, utilized data acquired from sites with known infestations and cannot be used to make regional estimates of change on forest land. By contrast, this study investigated the regional impacts of EAB on the affected resource using information from a large-scale forest inventory (Forest Inventory and Analysis program of the US Department of Agriculture, Forest Service) previously implemented to estimate regional forest resources. Results indicate that since 1980, ash has been increasing throughout many of the Great Lakes States but EAB is reversing this trend in recently invaded areas. Within 50 km of the epicenter of the EAB invasion, a major decline was observed after 2004. For growing stock (trees at least 12.7 cm diameter at breast height), average ash volume decreased from 12.7 to 3.2 m3⿢ha⿿1 and mortality increased from 0.1 to 1.4 m3⿢ha⿿1⿢year⿿1 on timberland between the 2004 and 2009 inventories. The consequences of single versus multiple mating on the longevity, fecundity, and fertility of female emerald ash borers were examined. In the first treatment, dissections of the common oviduct showed that 43 of 52 singly-mated females had received spermatophores. In the next two treatments, females were observed to mate one time, then housed either alone (observed separate) or with their mate (observed together). In the fourth treatment, females were paired with a randomly chosen male (unobserved together). Weight (0.0428 SE 0.0008 g) and longevity (50.5 SE 1.6 d) of female beetles did not differ among treatments. Fecundity, but not fertility, had a significant positive correlation with longevity in all treatments. Almost all of the females ⿿observed together⿿ laid eggs (87%, N = 31), while significantly fewer females ⿿unobserved together⿿ (61%, N = 31) and ⿿observed separate⿿ (54%, N = 31) did. The fecundity of females that did lay eggs did not differ among treatments. Based on our results a single mating may be sufficient to ensure maximal fecundity for females, but there is potential for failure of any one mating, and no apparent cost to multiple mating. Thus, multiple mating is likely the best strategy for female emerald ash borers to maximize fecundity. The implications of results for laboratory rearing, and potential population level effects are discussed. PARTICIPANTS: NCEAS MSU Agri Food Canada, APHIS, ARS, Chinese Academy of Sciences, CT Agricultural Experiment Station, VA Tech, USFS-FHP, Cornell University, University of Tennessee, Osaka Museum of Natural History, TARGET AUDIENCES: Policymakers, scientists, pest managers, quarantine/trade managers, forest and land managers

Impacts
The impacts of invasive species on forests and urban trees are a major reason for managers⿿ concern over their introduction. Recent work has expanded on the ecological and socio-economic effects of emerald ash borer and invasive species in general. Our research into basic biological factors of invasive plants, insects, and diseases provides the information needed by other scientists and managers to develop new control and management treatments and tactics.

Publications

Bray, Alicia M.; Bauer, Leah S.; Poland, Therese M.; Haack, Robert A.; Cognato, Anthony I.; Smith, James J. 2011. Genetic analysis of emerald ash borer (Agrilus planipennis Fairmaire) populations in Asia and North America. Biological Invasions. 13: 2869-2887.
Chamorro, M. Lourdes; Lingafelter, Steven W.; Haack, Robert A.; Poland, Therese M.; Volkovitsh, Mark G.; Zhang, Runzhi. 2011. Systematics and biology of Agrilus planipennis Fairmaire (emerald ash borer) and its relatives: A new USDA RS-FS international initiative. In McManus, Katherine A.; Gottschalk, Kurt W., eds. Proceedings, 22nd U.S. Department of Agriculture, Interagency Research Forum on Invasive Species 2011; 2011 January 11-14; Annapolis, MD. Gen. Tech. Rep. NRS-P-92. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northern Research Station: 68. Abstract
Chamorro, M. Lourdes; Volkovitsh, Mark G.; Poland, Therese M.; Haack, Robert A.; Lingafelter, Steven W. 2012. Preimaginal stages of the emerald ash borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae): An invasive species of ash trees (Fraxinus). PLoS ONE. 7(3): e33185. doi:10.1371/journal.pone.0033185. 12 p.
Dean, Kimberly M; Vandenberg, John D.; Griggs, Michael H.; Bauer, Leah S.; Fierke, Melissa K. 2012. Susceptibility of two hymenopteran parasitoids of Agrilus planipennis (Coleoptera: Buprestidae) to the entomopathogenic fungus Beauveria bassiana (Ascomycota: Hypocreales). Journal of Invertebrate Pathology. 109: 303-306.
Duan, Jian J.; Oppel, Craig B.; Ulyshen, Michael D.; Bauer, Leah S.; LeLito, Jonathan. 2011. Biology and life history of Tetrastichus planipennisi (Hymenoptera: Eulophidae) a larval endoparasitoid of the emerald ash borer (Coleoptera:Buprestidae). Florida Entomologist. 94(4): 933-940.
Grubin, Sarah M.; Ross, Darrell W.; Wallin, Kimberly F. 2011. Prey suitability and phenology of Leucopis spp (Diptera: Chamaemyiidae) associated with hemlock woolly adelgid (Hemiptera: Adelgidae) in the Pacific Northwest. Environmental Entomology. 40(6): 1410-1416.
Haack, Robert A. 2011. An entomological collectible: Beetle wing artwork. Newsletter of the Michigan Entomological Society 56(3&4): 40.
Haack, Robert A.; Fulbright, Dennis W.; Battisti, Andrea. 2011. The Asian chestnut gall wasp: A threat to Michigan⿿s chestnut industry and worldwide. Newsletter of the Michigan Entomological Society 56(3&4): 31.
Hajek, Ann E.; Tobin, Patrick C. 2011. Introduced pathogens follow the invasion front of a spreading alien host. Journal of Animal Ecology. 80: 1217-1226.
Havill, Nathan; Davis, Gina; Fischer, Melissa; Salom, Scott; Mausel, Dave; Onken, Brad. 2011. The introduction of Laricobius nigrinus as a biological control agent for the hemlock woolly adelgid: Is there a threat to the native congener, L. rubidus? Chp. 21, p.212-221 in Onken, Brad; Reardon Richard, eds. Implementation and status of biological control of the hemlock woolly adelgid. FHTET-2011-4.
Havill, Nathan; Montgomery, Michael E.; Keena, Melody A. 2011. Hemlock woolly adelgid and its hemlock hosts: A global perspective. Chp. 1, p. 3-14 in Onken, Brad; Reardon Richard, eds. Implementation and status of biological control of the hemlock woolly adelgid. FHTET-2011-4.
Haynes, Kyle J.; Liebhold, Andrew M.; Johnson, Derek M. 2012. Elevational gradient in the cyclicity of a forest-defoliating insect. Population Ecology. 54: 239-250.
Keena, Melody A.; Trotter, R. Talbot; Cheah, Carole; Montgomery, Michael. 2011. Temperature requirements to break the egg diapause of Scymnus camptodromus (Coleoptera: Coccinellidae). In McManus, Katherine A.; Gottschalk, Kurt W., eds. Proceedings, 22nd U.S. Department of Agriculture, Interagency Research Forum on Invasive Species 2011; 2011 January 11-14; Annapolis, MD. Gen. Tech. Rep. NRS-P-92. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northern Research Station: 75. Abstract
Lamb, Ashley; Montgomery, Michael E.; Viera, Ligia Cota; Shiyake, Shigehiko; Salom, Scott. 2011. Laricobius osakensis, a hemlock wooly adelgid predator from Japan. Chp. 7, p. 90-96 in Onken, Brad; Reardon Richard, eds. Implementation and status of biological control of the hemlock woolly adelgid. FHTET-2011-4.
Lee, Jung-Su; Haack, Robert A.; Choi, Won Il. 2011. Attack pattern of Platypus koryoensis (Coleoptera: Curculionidae: Platypodidae) in relation to crown dieback of Mongolian oak in Korea. Environmental Entomology. 40: 1363-1369.
Liebhold, A.M., K.J. Haynes and O.N. Bjørnstad. 2012. Spatial Synchrony of Insect Outbreaks. pages 113⿿125 in ⿿Insect Outbreaks Revisited⿝ (P. Barbosa, D.K. Letourneau, A.A. Agrawal, eds.). Wiley, New York. 465 pp.
Liebhold, Andrew M. 2012. Synchrony, Spatial, pp. 734-737 in Hastings, A; Gross, L.J., eds. Encyclopedia of Theoretical Ecology. University of California Press.
McManus, Katherine A; Gottschalk, Kurt W., eds. 2011. Proceedings 22nd US Department of Agriculture interagency research forum on invasive species 2011; 2011 January 11-14; Annapolis, MD. Gen. Tech. Rep. NRS-P-92. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northern Research Station. 105 p.
Montgomery, Michael E.; Keena, Melody A. 2011. Scymnus (Neopullus) lady beetles from China. Chp. 5, p. 53-76 in Onken, Brad; Reardon Richard, eds. Implementation and status of biological control of the hemlock woolly adelgid. FHTET-2011-4.
Montgomery, Michael E.; McAvoy, Thomas J.; Salom, Scott M. 2011. Other species considered. Chp. 10, p. 116-122 in Onken, Brad; Reardon Richard, eds. Implementation and status of biological control of the hemlock woolly adelgid. FHTET-2011-4.
Ott, Daniel S.; Yanchuk, Alvin D.; Huber, Dezene P.W.; Wallin, Kimberly F. 2011. Genetic variation of lodgepole pine, Pinus contorta var. latifolia, chemical and physical defenses that affect mountain pine beetle, Dendoctronus ponderosae, attack and tree mortality. Journal of Chemical Ecology 37: 1002-1012.
Pugh, Scott A.; Liebhold, Andrew M.; Morin, Randall S. 2011. Changes in ash tree demography associated with emerald ash borer invasion, indicated by regional forest inventory data from the Great Lakes States. Canadian Journal of Forest Research 41: 2165-2175.
Ross, D.W., Gaimari, S.D., Kohler, G.R., Wallin, K.F., Grubin, S.M. 2012 Chamaemyiid predators of the hemlock woolly adelgid from the Pacific Northwest. Chp. 8, pp. 97-106. in Onken, Brad; Reardon Richard, eds. Implementation and status of biological control of the hemlock woolly adelgid. FHTET-2011-4.
Rutledge, Claire E.; Keena, Melody A. 2012. Mating frequency and fecundity in the emerald ash borer Agrilus planipennis (Coleoptera: Buprestidae). Annals of the Entomological Society of America. 105(1): 66-72.
Shiyake, Shigehiko; Montgomery, Michael E.; Leschen, Richard A.B. 2012. Three species of Laricobius (Coleoptera: Derondontidae) recorded from Japan in 2011. Sayabane 5: 11-15
Ulyshen, Michael D., Duan, Jian J., Bauer, Leah S., Gould, Juli R., Taylor, Phil, Bean, Dick, Holko, Carol, Van Driesche, Roy. 2011. Field-cage methodology for evaluating climatic suitability for introduced wood-borer parasitoids: Preliminary results from the emerald ash borer system. Journal of Insect Science, 11(141), 10 p.
Wallin, K.F. 2012. Integrating the early steps of host selection behavior into biological control of HWA, Chp. 20, pp. 202-211. in Onken, Brad; Reardon Richard, eds. Implementation and status of biological control of the hemlock woolly adelgid. FHTET-2011-4.
Wallin, Kimberly F.; Latty, Tanya M.; Ross, Darrell W. 2011. Orientation behavior of the predator Laricobius nigrinus (Coleoptera: Derodontidae) to hemlock woolly adelgid and host tree odors in a multi-chambered olfactometer. Environmental Entomology. 40(4): 864-872.

Progress 10/01/10 to 09/30/11

Outputs
OUTPUTS: Reliable estimates of the impacts and costs of biological invasions are critical to developing credible management, trade and regulatory policies. We developed a novel modeling approach that maximizes the use of available data, accounts for multiple sources of uncertainty, and provides cost estimates for three major feeding guilds of non-native forest insects. For each guild, we calculated the economic damages for five cost categories and we estimated the probability of future introductions of damaging pests. We found that costs are largely borne by homeowners and municipal governments. Wood- and phloemboring insects are anticipated to cause the largest economic impacts by annually inducing nearly $1.7 billion in local government expenditures and approximately $830 million in lost residential property values. Our damage estimates provide a crucial but previously missing component of cost-benefit analyses to evaluate policies and management options intended to reduce species introductions. The flight variability in the woodwasp Sirex noctilio was described using tethered females in a flight mill device. The maximum distance recorded was 49km. All wasps lost weight during flight (mean weight loss of 10.0% of initial body mass). Flight patterns for each individual were significantly dependent on wasp body mass, suggesting a relationship with the resources used in flight and their availability. Variability in size and behavior can have significant consequences on population dynamics by determining local and regional dispersal. The success of Japanese stiltgrass as an invader may be due to its ability to respond to stochastic events and to maintain a beneficial genetic make-up when conditions are stable. This paper evaluates the importance of Japanese stiltgrass seed type in terms of seed mass, viability, and germination across variable moisture regimes (three regions in West Virginia) and at two light levels (roadside and forest interior). Seeds from the drier sites were smaller in mass than seeds from the more mesic sites. Japanese stiltgrass' longer-lived and larger seeds from the roadsides may ensure population survival over the long term. Cohorts of emerald ash borer larvae were experimentally established in July of 2008 on healthy green ash trees in wooded plots by caging gravid emerald ash borer females or placing laboratory-reared eggs on trunks of selected trees. Host tree defense was the most important mortality factor, causing 32.0 to 41.1% mortality in the experimental cohorts and 17.5 to 21.5% in wild emerald ash borer stages by spring 2009, and 16.1 to 29% for the remaining experimental cohorts, and 9.9 to 11.8% for wild immature emerald ash borer stages by fall 2009. Woodpecker predation was the second most important factor, inflicting no mortality in the experimental cohorts but causing 5.0 to 5.6% mortality to associated wild emerald ash borer stages by spring 2009 and 9.2 to 12.8% and 3.2 to 17.7%, respectively, for experimental cohorts and wild emerald ash borer stages by fall 2009. Mortality from disease and parasites in both the experimental and wild cohorts was low in both the spring and fall sample periods. PARTICIPANTS: Agriculture and Agri-Food Canada, APHIS, ARS, Arizona State University, BOKU-University of Natural Resources and Applied Life Sciences-Vienna, Benedictine University, Canadian Forest Service, Chinese Academy of Sciences, Cornell University, Dartmouth College, Laboratorio de Ecología de Insectos INTA Bariloche Argentina, National Center for Ecological Analysis and Synthesis, Michigan State University, McGill University, North Carolina State University, Paul Smith's College, The Pennsylvania State University, Russian Academy of Science, S&PF Forest Health Protection, Slovenian Forestry Institute, Swiss Federal Research Institute, Systematic Entomology Laboratory, University of California-Santa Barbara, University of Central Florida, University of Louisiana, University of Massachusetts, University of Minnesota, Virginia Tech, West Virginia University TARGET AUDIENCES: Policymakers, scientists, pest managers, quarantine/trade managers, forest and land managers

Impacts
The impacts of invasive species on forests and urban trees are a major reason for managers' concern over their introduction. Research on quantifying those economic impacts has moved forward significantly with the work conducted by the National Center for Ecological Analysis and Synthesis in which Forest Service R&D participated. This new information has great utility for policymakers and managers alike and has already been cited in Congressional hearings and in the text supporting new bills introduced into Congress. Our research into basic biological factors of invasive plants, insects, and diseases provides the information needed by other scientists and managers to develop new control and management treatments and tactics.

Publications

Aldrich, Preston R.; Bruguglio, Joseph S.; Kapadia, Shyam N., Morker, Minesh U.; Rawal, Ankit; Kalra, Preeti; Huebner Cynthia; Greer, Gary K. 2010. Genetic structure of the invasive tree Ailanthus altissima in eastern United States cities. Journal of Botany doi:10.1155/2010/795735. 9 pages
Aukema, J.E., D.G. McCullough, B. Von Holle, A.M. Liebhold, K. Britton and S.J. Frankel. 2010. Historical Accumulation of Nonindigenous Forest Pests in the Continental US. Bioscience 60: 886-897.
Aukema, J.E.; Leung, B.; Kovacs, Kent; Chivers, Corey; Britton, Kerry O.; Englin, Jeffrey; Frankel, Susan J.; Haight, Robert G.; Holmes, Thomas P.; Liebhold, Andrew M.; McCullough, Deborah G.; Von Holle, Betsy. 2011 Economic Impacts of Non-Native Forest Insects in the Continental United States. PLoS ONE 6(9): e24587. doi:10.1371/journal.pone.0024587
Bigsby, Kevin M.; Tobin, Patrick C.; Sills, Erin O. 2011. Anthropogenic drivers of gypsy moth spread. Biological Invasions. 13: 2077-2090.
Bray, Alicia M.; Bauer, Leah S.; Poland, Therese M.; Haack, Bob A.; Smith, James J. 2011. Invasion genetics of emerald ash borer (Agrilus planipennis Fairmaire). In: McManus, Katherine A; Gottschalk, Kurt W., eds. 2010. Proceedings. 21st U.S. Department of Agriculture interagency research forum on invasive species 2010; 2010 January 12-15; Annapolis, MD. Gen. Tech. Rep. NRS-P-75. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northern Research Station: 5.
Bruzzone, Octavio; Villacide, José M.; Bernstein, Carlos; Corley, Juan C. 2009. Flight variability in the woodwasp, Sirex noctilio (Hymenoptera: Siricidae): an analysis of flight data using wavelets. The Journal of Experimental Biology 212: 731-737.
Chen, Shi; Fleischer, Shelby J.; Tobin, Patrick C.; Saunders, Michael C. 2011. Projecting insect voltinism under high and low greenhouse gas emission conditions. Environmental Entomology. 40(3): 505-515.
Chen, Yigen; Ciaramitaro, Tina; Poland, Therese M. 2011. Moisture content and nutrition as selection forces for emerald ash borer larval feeding behaviour. Ecological Entomology. 36: 344-354.
Chen, Yigen; Poland, Therese. 2010. Wood-boring emerald ash borer larval feeding changes black ash foliar chemistry. Newsletter of the Michigan Entomological Society 55: 29
Corley, Juan; Villacide, José. 2009. Dispersal and dynamics of the woodwasp, Sirex noctilio, in Argentina. In McManus, Katherine A.; Gottschalk, Kurt W.; eds. Proceedings, 19th U.S. Department of Agriculture Interagency Research Forum on Invasive Species 2008; 2008 January 8-11; Annapolis, MD. Gen. Tech. Rep. NRS-P-36. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northern Research Station: 13-15.Abstract.
Duan, Jian J.; Ulyshen, Michael D.; Bauer, Leah S.; Gould, Juli, Van Driesche, Roy. 2010. Measuring the impact of biotic factors on populations of immature emerald ash borers (Coleoptera: Buprestidae). Environmental Entomology 39(5): 1513-1522.
Garnas, Jeffrey R.; Ayres, Matthew P.; Liebhold, Andrew M.; Evans, Celia. 2011. Subcontinental impacts of an invasive tree disease on forest structure and dynamics. Journal of Ecology 99: 532-541.
Haack, Robert A.; Vasily Grebennikov, V.; Jendek, Eduard; Petrice, Toby R.;Zablotny, James E. 2011. The European oak borer, Agrilus sulcicollis (Coleoptera: Buprestidae): new to North America. In: McManus, Katherine A; Gottschalk, Kurt W., eds. 2010. Proceedings. 21st U.S. Department of Agriculture interagency research forum on invasive species 2010; 2010 January 12-15; Annapolis, MD. Gen. Tech. Rep. NRS-P-75. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northern Research Station: 90.
Hansen, Jason A.; Petrice, Toby R.; Haack, Robert A. 2011. New state distribution and host records of North American Buprestidae (Coleoptera). Great Lakes Entomologist 44 (1-2): 74-77.
Havill, Nathan; Davis, Gina; Salom, Scott; Klein, Joanne; Caccone, Adalgisa. 2010. Population genetics of Laricobius nigrinus and L. rubidus, predators or adelgids in North America. In Onken, B.; and Reardon, R., compilers. Fifth Symposium on Hemlock Woolly Adelgid in the Eastern United States; 2010 August 17-19; Asheville, NC. FHTET-2010-7, Morgantown, WV: U.S. Department of Agriculture, Forest Service, Forest Health Technology Enterprise Team: 5-6. Abstract.
Huang, Dingcheng; Haack, Robert A.; Zhang, Runzhi. 2011. Does global warning increase establishment rates of invasive alien species A centurial time series analysis. PLoS One. 6(9): e24733.
Huebner, C.D. 2011. Seed Mass, Viability, and Germination of Japanese Stiltgrass (Microstegium vimineum) under Variable Light and Moisture Conditions. Invasive Plant Science and Management 4:274-283.
Huebner, Cynthia D. 2011. Establishment and spread of Microstegium vimineum (Japanese stiltgrass) in closed-canopy forests. In: McManus, Katherine A; Gottschalk, Kurt W., eds. 2010. Proceedings. 21st U.S. Department of Agriculture interagency research forum on invasive species 2010; 2010 January 12-15; Annapolis, MD. Gen. Tech. Rep. NRS-P-75. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northern Research Station: 29.
Johnson, Derek M.; Büntgen, Ulf; Frank, David C.; Kausrud, Kyrre; Haynes, Kyle J.; Liebhold, Andrew M.; Esper, Jan; Stenseth, Nils Chr. 2010. Climatic warming disrupts recurrent Alpine insect outbreaks. Proceedings, National Academy of Science 107: 20576-20581.
Koenig, Walter D.; Ries, Leslie; Olsen, V. Beth K.; Liebhold, Andrew M. 2011. Avian predators are less abundant during periodical cicada emergences, but why? Ecology 91: 784-790.
Koenig, Walter D.; Walters, Eric L.; Liebhold, Andrew M. 2011. Effects of gypsy moth outbreaks on North American woodpeckers. The Condor. 113(2): 352-361.
Liebhold, Andrew M.; McCullough, Deborah G. 2010. Forest Insects, pp. 238-241. In Simberloff, Daniel and Rejmánek, eds. Encyclopedia of Biological Invasions. University of California Press. Refereed
McManus, Katherine A; Gottschalk, Kurt W., eds. 2010. Proceedings, 21st U.S. Department of Agriculture interagency research forum on invasive species 2010; 2010 January 12-15; Annapolis, MD. Gen. Tech. Rep. NRS-P-75. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northern Research Station. 156 p.
Petrice, Toby; Haack, Robert A. 2010. Studies on the biology of the European oak borer, Agrilus sulcicollis (Coleoptera: Buprestidae), in the United States. Newsletter of the Michigan Entomological Society 55: 29.
Schwartzberg, Lora; Montgomery, Michael E. 2010. Relationships of nonstructural carbohydrates and the hemlock woolly adelgid. 2010. In Onken, B.; and Reardon, R., compilers. Fifth Symposium on Hemlock Woolly Adelgid in the Eastern United States; 2010 August 17-19; Asheville, NC. FHTET-2010-7, Morgantown, WV: U.S. Department of Agriculture, Forest Service, Forest Health Technology Enterprise Team: 198. Abstract.
Timer, Jody; Tobin, Patrick C.; Saunders, Michael C. 2010. Geographic variation in diapause induction: The grape berry moth (Lepidoptera: Tortricidae). Environmental Entomology. 39(6): 1751-1755.
Tobin, Patrick C.; Liebhold, Andrew M. 2011. Gypsy moth, pp. 298-304. In Simberloff, Daniel and Rejmánek, eds. Encyclopedia of Biological Invasions. University of California Press. Refereed
Tobin, Patrick C.; Van Stappen, Julie; Blackburn, Laura M. 2011. Visitation rates to the Apostle Islands National Lakeshore and the introduction of the non-native species Lymantria dispar (L). In: McManus, Katherine A; Gottschalk, Kurt W., eds. 2010. Proceedings. 21st U.S. Department of Agriculture interagency research forum on invasive species 2010; 2010 January 12-15; Annapolis, MD. Gen. Tech. Rep. NRS-P-75. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northern Research Station: 134.
Trotter, R. Talbot III. 2010. Long-term weather variability and shifting distribution limits of the invasive hemlock woolly adelgid (Adelges tsugae Annand). In: McManus, Katherine A; Gottschalk, Kurt W., eds. 2010. Proceedings. 21st U.S. Department of Agriculture interagency research forum on invasive species 2010; 2010 January 12-15; Annapolis, MD. Gen. Tech. Rep. NRS-P-75. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northern Research Station: 136-137.
Vercken, E.; Kramer, A.M.; Tobin, P.C.; Drake, J.M. 2011. Critical patch size generated by Allee effect in gypsy moth, Lymantria dispar (L.). Ecology Letters. 14: 179-186.
Whitney, Alexandra N.; Keena, Melody A. 2011. Effects of host wood moisture on the life cycle development of the Asian longhorned beetle. In: McManus, Katherine A; Gottschalk, Kurt W., eds. 2010. Proceedings. 21st U.S. Department of Agriculture interagency research forum on invasive species 2010; 2010 January 12-15; Annapolis, MD. Gen. Tech. Rep. NRS-P-75. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northern Research Station: 142.

Progress 10/01/09 to 09/30/10

Outputs
OUTPUTS: Recent analyses on the dynamics and spread of the emerald ash borer (EAB) has revealed a 2 yr life cycle in the Great Lakes area, and that newly-colonized trees were found up to 638 and 540 m from infested source populations after 1 and 3 yr, respectively. Studies also indicated that physical or phytochemical changes in host trees induced by light exposure could account for EAB preference for host trees grown in the sun. Economic analysis based upon the discounted cost of ash treatment, removal, and replacement, coupled with simulations of EAB spread and infestation over the next decade, predicted an expanding EAB infestation over 25 states leading to the replacement of more than 17 million ash trees at a discounted cost of $10.7 billion. Recent studies on the initial establishment of invasive species have shown that many new colonies go extinct in the absence of management because these new colonies lack a sufficient density to sustain it. Some of this work builds upon complementary work in conservation biology where low-density populations are threatened with extinction. The results suggest that many non-native invasive insects require a minimum number of individuals when invading new areas to ensure reproductive success or successfully colonize new host trees or habitats. Insect forest outbreaks remain an important component of forest dynamics. Recent work has shown that the regional weather patterns acting directly on gypsy moth outbreaks and its natural enemies had little effect on the outbreak patterns of this insect, but that patterns in lower trophic levels, such as mast seeding, can propagate across higher trophic levels, thus explaining concurrent gypsy moth outbreaks across several states. Studies on the larch budmoth have highlighted the complexity of climate change and its effect in forest ecosystems by revealing how increasing temperatures can lead to a reduction in outbreak intensity. The invasion dynamics of non-native invasive plants was studied to address the patterns of invasion with the development of lands for human housing. Using housing data from the New England and data on plant invasions, it was shown that housing was positively associated with invasive plant species richness and consequently, that the number of invasive non-native plants will likely increase as a result of future housing growth. Other studies on the invasion patterns of Microstegium vimineum, a non-native weed, confirmed that this plant is more successful in invading roadside habitats than forest interiors, and that greater canopy openings in forests was positively associated with its presence. PARTICIPANTS: Agriculture and Agri-Food Canada, APHIS, ARS, BOKU-University of Natural Resources and Applied Life Sciences-Vienna, Canadian Forest Service, Chinese Academy of Sciences, Cornell University, Michigan State University, Mississippi State University, The Pennsylvania State University, Russian Academy of Science, S&PF Forest Health Protection, Slovenian Forestry Institute, Swiss Federal Research Institute, Systematic Entomology Laboratory, University of Arkansas, University of Louisiana, University of Nevada, University of Padua-Italy, University of Virginia, University of Wisconsin, Virginia Department of Game and Inland Fisheries, Virginia Tech, West Virginia University TARGET AUDIENCES: Policymakers, scientists, pest managers, quarantine/trade managers, forest and land managers

Impacts
Information about EAB dispersal, and feeding preferences, is critically needed to contend with new infestations. Understanding these dynamics can be used to develop appropriate protocols for EAB detection and regulatory activities. The estimates of costs suggest that a substantial investment might be efficiently spent to slow EAB spread and postpone the ultimate costs of ash treatment, removal, and replacement. Information on invasion success and its link to initial population density is a critical component when allocating resources for management. Because many invaders require a minimum number to successful establish, there is an opportunity to considering the population dynamics of an invader when constructing management guidelines. Drivers of forest insect outbreaks across large regions are critical because large scale outbreaks can overwhelm the budgetary and logistical abilities of agencies to suppress populations. Understanding these dynamics, and the potential influence of climate change on outbreaks, facilitates the development of science-based management protocols to manage outbreaks. Due to the positive association between housing and the presence of non-native invasive plants, housing development should be considered in plans to manage and monitor invasive plants. Also, because of past work indicating that half of non-native plant species in North America are shade tolerant, which suggests our forests are susceptible to invasion, identifying plants that are more likely to invade forests will help prioritize resources in management strategies.

Publications

Huebner, Cynthia D. 2010. Establishment of an invasive grass in closed-canopy deciduous forests across local and regional environmental gradients. Biological Invasions. 12: 2069-2080.
Kovacs, Kent F.; Haight, Robert G.; McCullough, Deborah G.; Mercader, Rodrigo J.; Siegert, Nathan W.; Liebhold, Andrew M. 2010. Cost of potential emerald ash borer damage in U.S. communities, 2009-2019. Ecological Economics 69: 569-578.
Liebhold A.; McCullough, D.; Aukema, J.; et al. 2010. Estimating the economic impact of alien forest insects in the USA. John A. Parrotta and Mary A. Carr, eds. Forests for the Future: Sustaining Society and the Environment, XXIII IUFRO World Congress, 23-28 August 2010, Seoul, Republic of Korea. Abstracts. The International Forestry Review 12(5): 366. Abstract.
Limback, Chenin K.; McCullough, Deborah G.; Chen, Yigen; et al. 2010. Host vigor and emerald ash borer larvae on green and white ash. In: Lance, David; Buck, James; Binion, Denise; Reardon, Richard; Mastro, Victor; compilers. Emerald ash borer research and technology development meeting: 20-21 October 2009; Pittsburgh, PA. FHTET-2010-01, Morgantown, WV: U.S. Department of Agriculture, Forest Health Technology Enterprise Team: 44-46. Abstract.
McManus, Michael L.; Gottschalk, Kurt W. 2009. Evolution of a meeting: 20+ years of research communication and coordination. In McManus, K.A.; Gottschalk K.W.; eds. Proceedings, 20th U.S. Department of Agriculture Interagency Research Forum on Invasive Species 2009; 2009 January 13-16; Annapolis, MD. Gen. Tech. Rep. NRS-P-51. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northern Research Station: 1-4.
Petrice, T.R.; Haack, R.A.; Strazanac, J.S.; Lelito, J.P. 2009. Biology and larval morphology of Agrilus suncinctus (Coleoptera: Buprestidae), with comparisons to the emerald ash borer, Agrilus planipennis. The Great Lakes Entomologist 42: 173-184.
Contarini, Mario; Onufrieva, Ksenia S.; Thorpe, Kevin W.; et al. 2009. Mate-finding failure as an important cause of Allee effects along the leading edge of an invading insect population. Entomologia experimentalis et applicata 133: 307-314.
Fajvan, Mary Ann; Bohall Wood, Petra. 2010. Maintenance of eastern hemlock forests: Factors associated with hemlock vulnerability to hemlock woolly adelgid. In: Rentch, James S.; Schuler, Thomas M., eds. 2010. Proceedings from the conference on the ecology and management of high-elevation forests in the central and southern Appalachian Mountains; 2009 May 14-15; Slatyfork, WV. Gen. Tech. Rep. NRS-P-64. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northern Research Station: 31-38.
Gavier-Pizarro,Gregorio I ; Radeloff, Volker C.; Stewart, Susan I.; Huebner,Cynthia D.;Keuler, Nicholas S. 2010. Housing is positively associated with invasive exotic plant species richness in New England. Ecological Applications. 20(7): 1913-1925
Gottschalk, K.W.; Long, R.P.; Wargo, P.M. 2010. Oak (Quercus) decline around the world: A review. In: XXIII IUFRO World Congress; John A. Parrotta and Mary A. Carr, eds. Forests for the Future: Sustaining Society and the Environment, XXIII IUFRO World Congress, 23-28 August 2010, Seoul, Republic of Korea. Abstracts. The International Forestry Review 12(5): 373. Abstract.
Haack, Robert A.; Petrice, Toby R.; Zablotny, James E. 2009. First report of the European oak borer, Agrilus sulcicollis (Coleoptera: Buprestidae), in the United States. Great Lakes Entomologist. 42: 1-7.
Haack, Robert A.; Hérard, Franck; Sun, Jinghua; Turgeon, Jean J. 2010. Managing invasive populations of Asian longhorned beetle and citrus longhorned beetle: a worldwide perspective. Annual Review of Entomology 55: 521-546.
Haack, Robert A.; Petrice, Toby R.; Zablotny, James E. 2009. The European oak borer, Agrilus sulcicollis (Coleoptera: Buprestidae): New to Michigan and the United States. Newsletter of the Michigan Entomological Society 54: 25.
Haight, Robert G.; Kovacs, Kent; Liebhold, Andrew M.; McCullough, Deborah G. 2009. Economic assessment of potential emerald ash borer damage in urban areas in the United States. In McManus, K.A.; Gottschalk K.W.; eds. Proceedings, 20th U.S. Department of Agriculture Interagency Research Forum on Invasive Species 2009; 2009 January 13-16; Annapolis, MD. Gen. Tech. Rep. NRS-P-51. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northern Research Station: 32-33. Abstract.
Hajek Ann E.; Tobin, Patrick C. 2009. A space-time odyssey: movement of gypsy moth and its pathogens. In McManus, K.A.; Gottschalk K.W.; eds. Proceedings, 20th U.S. Department of Agriculture Interagency Research Forum on Invasive Species 2009; 2009 January 13-16; Annapolis, MD. Gen. Tech. Rep. NRS-P-51. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northern Research Station: 33. Abstract.
Havill, Nathan P.; Klein, Joanne; Keena, Melody A.; Caccone, Adalgisa. 2009. Characterization of microsatellite loci for Laricobius nigrinus and L rubidus, predators of adelgids in North America. In McManus, K.A.; Gottschalk K.W.; eds. Proceedings, 20th U.S. Department of Agriculture Interagency Research Forum on Invasive Species 2009; 2009 January 13-16; Annapolis, MD. Gen. Tech. Rep. NRS-P-51. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northern Research Station: 76. Abstract.
Havill, Nathan P.; Montgomery, Michael; Shiyake, Shigehiko; Lamb, Ashley; Keena, Melody A.; Caccone, Adalgisa. 2009. Hemlock woolly adelgid population genetics. In McManus, K.A.; Gottschalk K.W.; eds. Proceedings, 20th U.S. Department of Agriculture Interagency Research Forum on Invasive Species 2009; 2009 January 13-16; Annapolis, MD. Gen. Tech. Rep. NRS-P-51. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northern Research Station: 75. Abstract.
Haynes, Kyle J.; Liebhold, Andrew M.; Fearer, Todd M.; et al. 2009. Spatial synchrony propagates through a forest food web via consumer-resource interactions. Ecology 90: 2974-2983.
Huber, John T.; Gibson, Gary A.P.; Bauer, Leah S.; Liu, Houping; Gates, Michael. 2008. The genus Mymaromella (Hymenoptera: Mymarommatidae) in North America, with a key to described extant species. Journal of Hymenoptera Research. 17(2): 175-194.
Plymale, Ruth C.; Liebhold, Andrew M.; Hajek, Ann E. 2009. Spatial population dynamics and heterogeneity of an insect/pathogen interaction. In McManus, K.A.; Gottschalk K.W.; eds. Proceedings, 20th U.S. Department of Agriculture Interagency Research Forum on Invasive Species 2009; 2009 January 13-16; Annapolis, MD. Gen. Tech. Rep. NRS-P-51. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northern Research Station: 92. Abstract.
Sieg, Carolyn Hull; Denslow, Julie S.; Huebner, Cynthia D.; Miller, James H. 2010. The role of the Forest Service in nonnative invasive plant research. In: Dix, Mary Ellen; Britton, Kerry, editors. A dynamic invasive species research vision: Opportunities and priorities 2009-29. Gen. Tech. Rep. WO-79/83. Washington, DC: U.S. Department of Agriculture, Forest Service, Research and Development. p. 35-41.
Siegert, N.W.; McCullough, D.G.; Williams, D.W.; Fraser, I.; Poland, T.M.; Pierce, S.J. 2010. Dispersal of Agrilus planipennis (Coleoptera: Buprestidae) from discrete epicenters in two outlier sites. Environmental Entomology 39: 253-265.
Siegert, Nathan W.; McCullough, Deborah G.; Poland, Therese M.; et al. 2010. Effects of cluster of girdled trees on the spread of emerald ash borer in low-density infestations. In: Lance, David; Buck, James; Binion, Denise; Reardon, Richard; Mastro, Victor; compilers. Emerald ash borer research and technology development meeting: 20-21 October 2009; Pittsburgh, PA. FHTET-2010-01, Morgantown, WV: U.S. Department of Agriculture, Forest Health Technology Enterprise Team: 23. Abstract.
Liebhold, A. 2010. The gypsy moth in North America: current status of an alien oak defoliator. John A. Parrotta and Mary A. Carr, eds. Forests for the Future: Sustaining Society and the Environment, XXIII IUFRO World Congress, 23-28 August 2010, Seoul, Republic of Korea. Abstracts. The International Forestry Review 12(5): 13-14. Abstract.
Liebhold, A.M. 2010. Invasions of forest insects: agents of global change. John A. Parrotta and Mary A. Carr, eds. Forests for the Future: Sustaining Society and the Environment, XXIII IUFRO World Congress, 23-28 August 2010, Seoul, Republic of Korea. Abstracts. The International Forestry Review 12(5): 376. Abstract.
Liebhold, Andrew M.; Tobin, Patrick C. 2010. Exploiting the Achilles heels of pest invasions: allee effects, stratified dispersal and management of forest insect establishment and spread. New Zealand Journal of Forestry Science 40 suppl.: S25-S33.
Tluczek, A.R.; McCullough, Deborah G.; Poland, T.M.; et al. 2010. The fast and the furious: influence of host vigor on development and mortality of emerald ash borer larvae. In: Lance, David; Buck, James; Binion, Denise; Reardon, Richard; Mastro, Victor; compilers. Emerald ash borer research and technology development meeting: 20-21 October 2009; Pittsburgh, PA. FHTET-2010-01, Morgantown, WV: U.S. Department of Agriculture, Forest Health Technology Enterprise Team: 47-48. Abstract.
Tobin, Patrick C.; Klein, Kenneth T.; Leonard, Donna S. 2009. Gypsy moth (Lepidoptera: Lymantriidae) flight behavior and phenology based on field-deployed automated pheromone-baited traps. Environmental Entomology 38: 1555-1562.
Abdoullaye, Doukary; Acevedo, I.; Adebayo, Abisola A.; et al. 2010. Permanent genetic resources added to the Molecular Ecology Resources Database 1 August 2009-30 September 2009. Molecular Ecology Resources 10: 232-236.
Aurelle, D.; Baker, A.J.; Bottin, L.; Brouat, C.; Caccone, A.; et al. 2010. Permanent genetic resources added to the Molecular Ecology Resources Database 1 February 2010-31 March 2010. Molecular Ecology Resources 10: 751-754.
Baranchikov, Yu.N.; Montgomery, M.E. 2009. Comparable evaluation of polyphagy of gypsy moth populations from different continents. Transactions of St. Petersburg Forest Engineering Academy 183: 40-46. [in Russian, English abstract]
Buntgen, Ulf; Frank, David; Liebhold, Andrew; Johnson, Derek; Carrer, Marco; Urbinati, Carlo; Grabner, Michael; Nicolussi, Kurt; Levanic, Tom; Esper. 2009. Three centuries of insect outbreaks across the European Alps. New Phytologist 182: 929-941.
Chen, Yigen; Poland, Therese M. 2009. Biotic and abiotic factors affect green ash volatile production and emerald ash borer adult feeding preference. Environmental Entomology 38: 1756-1764.

Progress 10/01/08 to 09/30/09

Outputs
OUTPUTS: Recent surveys of hemlock woolly adelgid (HWA) mortality across the eastern United States were used to show a significant positive relationship between minimum winter temperatures and winter survival at the landscape scale. The strength and nature of this relationship however, varies through time. Using landscape estimates of minimum winter temperature, two simple methods of estimating landscape-scale adelgid survival rates suggest much of the range of Tsuga canadensis in the US and the entire range of T. caroliniana fall in areas where winter temperatures will not impose critical limits on HWA populations. The relationship between eastern hemlock (T. canadensis) crown condition and changes in radial growth associated with infestation by HWA was evaluated. Tree-ring chronologies of eastern hemlock were used to develop a binomial decline index based on three consecutive years of below average growth. Radial growth decline was modeled as a function of an extensive array of tree, crown, and site variables that were collected over an 11 year period in Delaware Water Gap National Recreation Area. Some site-related variables such as site-location and aspect were significantly related to decline probabilities individually. However, they explained only a small proportion of response variance, and the only site variable included in the final model was mean plot-level HWA infestation level. For every 1% increase in mean percent HWA infestation per plot, there was an 8% increase in the likelihood that a tree would be classified as being in decline. Intensive Forest Inventory and Analysis (FIA) plot data collected in the Allegheny National Forest (ANF), Pennsylvania, between 1999 and 2006 were evaluated for their ability to predict ANF⿿s vulnerability to invasion by exotic plants. Approximately, 11% of the 449 species documented in these plots were exotic, which is higher than has been found in other northeastern forested plots. Only1%of the ANF flora was invasive exotic plants and these were at low abundance, confirming that most invasions are still at an early stage of establishment. Sites richer in native or non-invasive exotic plants and with more alkaline soils were more likely to be invaded. Younger forests, forests with non-forest patches present, and forests rich in native species were more likely to be colonized by exotic plants. Seven different Phytophthora species were used to test the foliar, root, and stem susceptibility of common eastern US oak species and understory plants to Phytophthora infection. Young foliage of Quercus rubra was the most susceptible to infection followed by Castanea dentata for both wounded and non-wounded inoculations. Mature foliage of Hamamelis virginiana, Kalmia latifolia, and Q. alba were the most susceptible to wound and non-wound inoculations. All Phytophthora species were pathogenic to fine and taproots of at least one oak species with fine root damage ranging from 9 to 55% greater than controls. Roots were more susceptible during the fall inoculation period than the summer. All but two species of Phytophthora were pathogenic to oak stems with Quercus montana and Q. rubra the most susceptible. PARTICIPANTS: West Virginia University, Michigan State University, University of Wisconsin, Cornell University, Wisconsin DNR, University of California-Berkeley, University of Canterbury, Chequamegon-Nicolet NF, Allegheny NF, University of Georgia, University of Idaho, Chinese Academy of Forestry, APHIS, USDOI-National Park Service, S&PF Forest Health Protection, France INRA, Canadian Forest Service, Pennsylvania State University, University of Puerto Rico-Mayaquez, University of Louisiana, Japanese Institute for Agro-Environmental Sciences TARGET AUDIENCES: Policymakers, scientists, pest managers, quarantine/trade managers, forest and land managers

Impacts
Knowledge of the effects of winter temperatures on HWA survival can be used by pest and forests managers to determine areas where treatments for HWA can be minimized due to the reduced winter survival. Treatment efforts can then be reassigned to areas where winter mortality will not be a critical component of the system. Tree crown variables such as live crown ratio, crown density, and the modified ZBadj index, a combination of foliage transparency and branch dieback, had the most explanatory power for predicting hemlock growth loss. These crown variables were relatively accurate predictors of the degree of hemlock growth decline during HWA infestation. Use of the crown variables is faster and easier than coring trees to collect growth decline information and can be used as a surrogate for that information. Increased use of intensive sampling for FIA in the U.S.A. and similar monitoring programs in other countries would increase the ability to find invasive plants when their infestations are still small, but suggest adding a step to the plot selection phase that would allow forest-wide or regional stratified sampling of typically coarse-scale variables, such as historic or predicted defoliation or fire events, and forest or land type. A more accurate picture of the importance of disturbance variables in defining forest vulnerability to plant invasion may be achieved which could then guide management decisions. Information on the susceptibility and resistance of common eastern forest species to Phytophthora species will better inform risk models and provide information to pest and forest managers that will allow the evaluation of potential impacts and alternatives for management to reduce those impacts.

Publications

Balci, Y.; Balci, S.; MacDonald, W.L.; and Gottschalk, K.W. 2008. Foliar susceptibility of eastern oak species to Phytophthora infection. For. Path. 38:320⿿331. DOI: 10.1111/j.1439-0329.2008.00548.x.
Balci, Y.; Balci, S.; MacDonald, W.L.; and Gottschalk, K.W. 2008. Relative susceptibility of oaks to seven species of Phytophthora isolated from oak forest soils. Forest Pathology 38, 394⿿409. DOI: 10.1111/j.1439-0329.2008.00559.x.
Gavier, Gregorio; Stewart, Susan; Huebner, Cynthia D.; Radeloff, Volker C. 2009. Is housing a factor of invasive plants distribution at coarse and fine scales?. In: McManus, Katherine A; Gottschalk, Kurt W., eds. Proceedings. 19th U.S. Department of Agriculture interagency research forum on invasive species 2008; 2008 January 8-11; Annapolis, MD. Gen. Tech. Rep. NRS-P-36. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northern Research Station: 27-29.
Gray, Rebecca Hoffman; Lorimer, Craig G.; Tobin, Patrick C.; Raffa, Kenneth F. 2008. Preoutbreak dynamics of a recently established invasive herbivore: roles of natural enemies and habitat structure in stage-specific performance of gypsy moth (Lepidoptera: Lymantriidae) populations in northeastern Wisconsin. Environmental Entomology. 37(5): 1174-1184.
Hajek, Ann E.; Tobin, Patrick C.; Blackburn, Laura; Hannam, Joshua J.; Diss-Torrance, Andrea; Raffa, Kenneth; Nielsen, Charlotte 2009. Dispersal of gypsy moth pathogens to newly established host populations. In: McManus, Katherine A; Gottschalk, Kurt W., eds. Proceedings. 19th U.S. Department of Agriculture interagency research forum on invasive species 2008; 2008 January 8-11; Annapolis, MD. Gen. Tech. Rep. NRS-P-36. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northern Research Station: 32.
Huebner, Cynthia D.; Morin, Randall S.; Zurbriggen, Ann; White, Robert L. 2009. Patterns of exotic plant invasions in Pennsylvania''s Allegheny National Forest using intensive Forest Inventory and Analysis plots. Forest Ecology and Management. 257: 258-270.
Keena, Melody A. 2009. Phenology of the Asian longhorned beetle under simulated annual environmental fluctuations. In: McManus, Katherine A; Gottschalk, Kurt W., eds. Proceedings. 19th U.S. Department of Agriculture interagency research forum on invasive species 2008; 2008 January 8-11; Annapolis, MD. Gen. Tech. Rep. NRS-P-36. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northern Research Station: 37.
Kelly, Dave; Koenig, Walter D.; Liebhold, Andrew M. 2008. An intercontinental comparison of the dynamic behavior of mast seeding communities. Population Ecology 50: 329-342.
Kramer, Andrew M.; Dennis, Brian; Liebhold, Andrew M.; Drake, John M. 2009. The evidence for Allee effects. Population Ecology. 51: 341-354.
Li, Li; Montgomery, Michael E.; Yu, Guoyue. 2009. Functional responses of Tetraphleps galchanoides preying on different stages of hemlock woolly adelgid. Chinese Bulletin of Entomology 46(1): 72-76. [in Chinese with English abstract]
McManus, Katherine A.; Gottschalk, Kurt W., eds. 2009. Proceedings 19th U.S. Department of Agriculture Interagency Research Forum on Invasive Species 2008. Gen. Tech. Rep. NRS-P-36. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northern Research Station. 100 p.
Perkowski, Matthew P.; Brooks, John R.; Gottschalk, Kurt W. 2008. Preliminary results of the FVS gypsy moth event monitor using remeasurement plot data from Northern West Virginia. In: Havis, Robert N.; Crookston, Nicholas L., comps. 2008. Third Forest Vegetation Simulator Conference; 2007 February 13⿿15; Fort Collins, CO. Proceedings RMRS-P-54. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station. p. 81-86
Petrice, Toby R.; Haack, Robert A.; Strazanac, John S.; Lelito, Jonathan P. 2008. The biology of Agrilus subcinctus (Coleoptera: Bupresdtida): our native ash Agrilus species. Newsletter of the Michigan Entomological Society 53(3 & 4): 41.
Rentch, James; Fajvan, Mary Ann; Evans, Richard A.; Onken, Bradley. 2009. Using dendrochronology to model hemlock woolly adelgid effects on eastern hemlock growth and vulnerability. Biological Invasions 11:551-563
Robinet, Christelle; Liebhold, Andrew M. 2009. Dispersal polymorphism in an invasive forest pest affects its ability to establish. Ecological Applications 19(7): 1935-1943.
Roden, D.B.; Haack, R.A.; Keena, M.A.; McKenney, D.W.; Beall, F.D.; Roden, P.M. 2009. Potential northern distribution of Asian longhorned beetle in North America. In: McManus, Katherine A; Gottschalk, Kurt W., eds. Proceedings. 19th U.S. Department of Agriculture interagency research forum on invasive species 2008; 2008 January 8-11; Annapolis, MD. Gen. Tech. Rep. NRS-P-36. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northern Research Station: 65-67.
Schwartzberg, Lora; Montgomery, Michael E.; McDonald, Richard. 2009. Preliminary study of the biology of Scymnus (Pullus) coniferarum Crotch 1874 (Coleoptera: Coccinellidae), an adelgid predator native to the western United States and a potential biocontrol for Adelges tsugae. Odum Conference 2009 ⿿ Understanding and managing biological invasions as dynamic processes: integrating information across space and time: proceedings of the meeting; 2009 April 30-May 1, 2009, Rensselaerville, NY: 2. Abstract. New York Invasive Species Institute, Ithaca, NY. Informally reviewed
Siegert, Nathan W.; McCullough, Deborah G.; Liebhold, Andrew M.; Telewski, Frank W. 2009. Reconstruction of the establishment and spread of emerald ash borer through dendrochronological analysis. In: McManus, Katherine A; Gottschalk, Kurt W., eds. Proceedings. 19th U.S. Department of Agriculture interagency research forum on invasive species 2008; 2008 January 8-11; Annapolis, MD. Gen. Tech. Rep. NRS-P-36. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northern Research Station: 70.
Strazanac, John S.; Gould, Juli R.; Haack, Robert A.; Fraser, Ivich 2009. Field release of Spathius agrili Yang (braconidae): monitoring non-target wood borers. In: McManus, Katherine A; Gottschalk, Kurt W., eds. Proceedings. 19th U.S. Department of Agriculture interagency research forum on invasive species 2008; 2008 January 8-11; Annapolis, MD. Gen. Tech. Rep. NRS-P-36. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northern Research Station: 78.
Sánchez, V.; Keena, M.A. 2009. Allozyme variation in hemlock woolly adelgid (Hemiptera: Adelgidae) from the United States and China. Annals, Entomological Society of America 103(3): 539-546.
Trotter, R. Talbot, III; Shields, Kathleen S. 2009. Variation in winter survival of the invasive hemlock woolly adelgid (Hemiptera: Adelgidae) across the eastern United States. Environmental Entomology. 38(3): 557-587.
Tobin, Patrick C.; Frazier, James L. 2009. A slide down a slippery slope: ethical guidelines in the dissemination of computer-based presentations. Bulletin of the Ecological Society of America 39-42.
Tobin, Patrick C.; Hajek, Ann E.; Blackburn, Laura M.; Hannam, Joshua J.; Diss-Torrance, Andrea; Raffa, Kenneth F.; Nielson, Charlotte 2009. Space-time interactions between gypsy moth and associated entomopathogens in newly established populations. In: McManus, Katherine A; Gottschalk, Kurt W., eds. Proceedings. 19th U.S. Department of Agriculture interagency research forum on invasive species 2008; 2008 January 8-11; Annapolis, MD. Gen. Tech. Rep. NRS-P-36. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northern Research Station: 81.
Tobin, Patrick C.; Robinet, Christelle; Johnson, Derek M.; Whitmire, Stefanie L.; Bjornstad, Ottar N.; Liebhold, Andrew M. 2009. The role of Allee effects in gypsy moth, Lymantria dispar (L.), invasions. Population Ecology. 51: 373-384.
Trotter, R. Talbot; Shields, Kathleen S. 2009. Assessing assumptions about adelgid attributes: landscape patterns of an invasive insect. In: McManus, Katherine A; Gottschalk, Kurt W., eds. Proceedings. 19th U.S. Department of Agriculture interagency research forum on invasive species 2008; 2008 January 8-11; Annapolis, MD. Gen. Tech. Rep. NRS-P-36. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northern Research Station: 82.
Yamanaka, Takehiko; Liebhold, Andrew M. 2009. Mate-location failure, the Allee effect, and the establishment of invading populations. Population Ecology. 51: 337-340.