Scientists & Staff
Nathan P. Havill
Current Research
With expertise in the ecology and evolution of multi-trophic interactions, I work to provide information about invasive pests, their natural enemies, and the impacted trees. Understanding the interactions among these trophic levels is critical to making sound management decisions to reduce the impact of invasive pests. Facets of my research objectives include investigating: 1) systematics and population genetics of invasive pests, especially hemlock woolly adelgid, balsam woolly adelgid, gypsy moth, and winter moth; 2) systematics and population genetics of predators and parasitoids being evaluated as biological control agents of invasive species, especially of hemlock woolly adelgid and emerald ash borer; and 3) systematics and biogeography of North American tree species and their relatives worldwide.
My goal is to help reconstruct the evolutionary history of multiple trophic levels of a pest system to guide management and mitigate the impact of invasive insect pests on forested ecosystems of the northeastern United States. I am especially concerned with research related to classical biological control to develop practical, cost effective tools used to evaluate the safety and efficacy of predators or parasites introduced as biological control agents.
Why This Research is Important
The exotic invasive pests that are the subject of this research originated in either Asia or Europe. Invasive insects in North America are not often pests in their region of origin due to resistant tree species and natural enemies that co-evolved with the pest over millions of years. The insect becomes a pest following introduction outside its native ranges when these bottom-up and/or top-down regulating factors are no long effective. The hemlock woolly adelgid, Adelges tsugae (Adelgidae), originated in Japan and causes the death and decline of eastern North American hemlock trees in 17 eastern states. The balsam woolly adelgid, Adelges piceae (Adelgidae), originated in Europe and has been killing fir trees throughout North America for over 100 years. The gypsy moth, Lymantria dispar (Lymantriidae), also was also introduced from Europe over 100 years ago. It is controlled to some extent by a pathogenic fungus in areas where it is well established, so management in the U. S. currently focuses on slowing its spread and on preventing invasion by more virulent strains from Asia. The winter moth, Operophtera brumata (Geometridae), is also from Europe and defoliates a broad range of tree species, including oaks and birches in southern New England, and is hybridizing with the native Bruce spanworm, O. bruceata. Finally, the emerald ash borer, Agrilus planipennis (Buprestidae) originated in Asia, was first discovered in the U.S. in Michigan in 2002, and has killed tens of millions of ash trees in at least 15 eastern states. Together, these pest species represent a diverse group of insects with very different feeding strategies, life cycles, host ranges, and natural enemy complexes.
Education
- Yale University, Ph.D. Ecology & Evolutionary Biology, 2006
- Yale University, M.S. Ecology & Evolutionary Biology, 2003
- University of Wisconsin, Madison, M.S. Entomology, 1998
- College of William and Mary, B.S. Biology, 1996
Professional Organizations
- Society for the Study of Evolution (2012 - Current)
- Connecticut Entomological Society (2001 - Current)
- Entomological Society of America (1996 - Current)
Featured Publications & Products
- Havill, Nathan P.; Montgomery, Michael E.; Keena, Melody. 2011. Hemlock woolly adelgid and its hemlock hosts: A global perspective. In: Onken, B.; Reardon, R. eds. Implementation and status of biological control of the hemlock woolly adelgid. FHTET-2011-04. Morgantown, WV: U.S. Department of Agriculture, Forest Service Forest Health Technology Enterprise Team: 3-14. Chapter 1.
- Michael E., Montgomery; Shiyake, S.; Havill, Nathan P. 2011. A new species of Laricobius (Coleoptera: Derodontidae) from Japan with phylogeny and a key for native and introduced congeners in North America. Annals of the Entomological Society of America 104(3):389-401.
- Havill, Nathan P.; Salom, Scott; Davis, Gina; Fischer, Melissa; Mausel, David; Onken, Bradley. 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?. In: Onken, B.; Reardon, R. eds. Implementation and status of biological control of the hemlock woolly adelgid. FHTET-2011-04. Morgantown, WV: U.S. Department of Agriculture, Forest Service Forest Health Technology Enterprise Team: 212-221. Chapter 21.
Publications & Products
- Liebhold, Andrew M.; Campbell, Faith T.; Gordon, Doria R.; Guo, Qinfeng ; Havill, Nathan ; Kinder, Bradley ; MacKenzie, Richard ; Lance, David R.; Pearson, Dean E.; Sing, Sharlene E.; Warziniack, Travis ; Venette, Robert C.; Yemshanov, Denys. 2021. The Role of International Cooperation in Invasive Species Research. In: Poland, Therese M.; Patel-Weynand, Toral; Finch, Deborah M.; Ford Miniat, Chelcy; Hayes, Deborah C.; Lopez, Vanessa M., eds. Invasive Species in Forests and Rangelands of the United States: A Comprehensive Science Synthesis for the United States Forest Sector. Heidelberg, Germany: Springer International Publishing: 293 - 304. Chapter 13.
- Blackburn, Laura M.; Elkinton, Joseph S.; Havill, Nathan P.; Broadley, Hannah J.; Andersen, Jeremy C.; Liebhold, Andrew M. 2020. Predicting the invasion range for a highly polyphagous and widespread forest herbivore. NeoBiota
- Davis, Gina ; Lowrey, Laura ; Eckberg, Tom ; Gannon, Amy ; Malesky, Danielle ; Havill, Nathan. 2020. Notes on balsam woolly adelgid, Adelges piceae (Ratzeburg, 1844) (Hemiptera: Adelgidae), range expansion in Idaho, Montana and Utah. The Pan-Pacific Entomologist
- Gagné, Raymond J.; Havill, Nathan P. 2020. A New Species of Lestodiplosis (Diptera: Cecidomyiidae) Preying on Pineus (Hemiptera: Adelgidae), with a Redescription of Lestodiplosis juniperina (Felt). Proceedings of the Entomological Society of Washington
- Havill, Nathan P.; Griffin, Brian P.; Andersen, Jeremy C.; Foottit, Robert G.; Justesen, Mathias J.; Caccone, Adalgisa ; D'Amico, Vincent ; Elkinton, Joseph S. 2020. Species delimitation and invasion history of the balsam woolly adelgid, Adelges (Dreyfusia) piceae (Hemiptera: Aphidoidea: Adelgidae), species complex. Systematic Entomology
- Neidermeier, Alex N.; Ross, Darrell W.; Havill, Nathan P.; Wallin, Kimberly F. 2020. Temporal Asynchrony of Adult Emergence Between Leucopis argenticollis and Leucopis piniperda (Diptera: Chamaemyiidae), Predators of the Hemlock Woolly Adelgid (Hemiptera: Adelgidae), with Implications for Biological Control. Environmental Entomology
- Rose, Alexander ; Ross, Darrell W.; Havill, Nathan P.; Motley, Kyle ; Wallin, Kimberly F. 2020. Coexistence of three specialist predators of the hemlock woolly adelgid in the Pacific Northwest USA. Bulletin of Entomological Research
- 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
- 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
- Andersen, Jeremy C.; Havill, Nathan P.; Broadley, Hannah J.; Boettner, George H.; Caccone, Adalgisa ; Elkinton, Joseph S. 2019. Widespread hybridization among native and invasive species of Operophtera moths (Lepidoptera: Geometridae) in Europe and North America. Biological Invasions
- 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
- Bittner, Tonya D.; Havill, Nathan ; Caetano, Isis A.L.; Hajek, Ann E. 2019. Efficacy of Kamona strain Deladenus siricidicola nematodes for biological control of Sirex noctilio in North America and hybridisation with invasive conspecifics. NeoBiota
- Elkinton, Joseph S.; Bittner, Tonya D.; Pasquarella, Valerie J.; Boettner, George H.; Liebhold, Andrew M.; Gould, Juli R.; Faubert, Heather ; Tewksbury, Lisa ; Broadley, Hannah J.; Havill, Nathan P.; Hajek, Ann E. 2019. Relating Aerial Deposition of Entomophaga maimaiga Conidia (Zoopagomycota: Entomophthorales) to Mortality of Gypsy Moth (Lepidoptera: Erebidae) Larvae and Nearby Defoliation. Environmental Entomology
- Havill, Nathan ; Cognato, Anthony ; del-Val, Ek ; Rabaglia, Robert ; Garrick, Ryan C. 2019. New Molecular Tools for Dendroctonus frontalis (Coleoptera: Curculionidae: Scolytinae) Reveal an East-West Genetic Subdivision of Early Pleistocene Origin. Insect Systematics and Diversity
- Mech, Angela M.; Harper, Sherilyn J.; Havill, Nathan P.; von Dohlen, Carol D.; Burke, Gaelen R. 2019. Ecological factors influencing the beneficial endosymbionts of the hemlock woolly adelgid (Hemiptera: Adelgidae). Insect Science
- 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
- Salom, Scott M.; Davis, Gina ; Elkinton, Joseph ; Foley, Jeremiah ; Havill, Nathan ; Jubb, Carrie ; Mayfield, Albert ; McAvoy, Tom ; Rhea, Rusty ; Talbot Trotter, R. ; Whitmore, Mark. 2019. A response to "Media representation of hemlock woolly adelgid management risks: a case study of science communication and invasive species control," published in biological invasions online on September 18, 2018. Biological Invasions
- Wantuch, Holly A.; Havill, Nathan P.; Hoebeke, E. Richard; Kuhar, Thomas P.; Salom, Scott M. 2019. Predators associated with the pine bark adelgid (Hemiptera: Adelgidae), a native insect in Appalachian forests, United States of America, in its southern range. The Canadian Entomologist
- Darr, Molly ; Brooks, Rachel ; Havill, Nathan ; Hoebeke, E. ; Salom, Scott. 2018. Phenology and Synchrony of Scymnus coniferarum (Coleoptera: Coccinellidae) with Multiple Adelgid Species in the Puget Sound, WA, USA. Forests
- Darr, Molly; Salom, Scott; Brooks, Rachel K.; Foottit, Robert G.; Miller, Gary L.; Havill, Nathan P. 2018. First report of Pineus strobi (Hartig, 1839) (Hemiptera: Adelgidae) in western North America. Pan-Pacific Entomologist
- Havill, Nathan P.; Gaimari, Stephen D.; Caccone, Adalgisa. 2018. Cryptic east-west divergence and molecular diagnostics for two species of silver flies (Diptera: Chamaemyiidae: Leucopis ) from North America being evaluated for biological control of hemlock woolly adelgid. Biological Control. 121: 23-29.
- Picq, Sandrine; Keena, Melody; Havill, Nathan; Stewart, Don; Pouliot, Esther; Boyle, Brian; Levesque, Roger C.; Hamelin, Richard C.; Cusson, Michel. 2018. Assessing the potential of genotyping-by-sequencing-derived single nucleotide polymorphisms to identify the geographic origins of intercepted gypsy moth (Lymantria dispar) specimens: A proof-of-concept study. Evolutionary Applications
- Weglarz, Kathryn M.; Havill, Nathan P.; Burke, Gaelen R.; von Dohlen, Carol D. 2018. Partnering With a Pest: Genomes of Hemlock Woolly Adelgid Symbionts Reveal Atypical Nutritional Provisioning Patterns in Dual-Obligate Bacteria. Genome Biology and Evolution
- Andersen, Jeremy C.; Havill, Nathan P.; Caccone, Adalgisa; Elkinton, Joseph S. 2017. Postglacial recolonization shaped the genetic diversity of the winter moth (Operophtera brumata) in Europe. Ecology and Evolution
- Havill, N.P.; Elkinton, J.; Andersen, J.C.; Hagen, S.B.; Broadley, Hannah J.; Boettner, G.J.; Caccone, A. 2017. Asymmetric hybridization between non-native winter moth, Operophtera brumata (Lepidoptera: Geometridae), and native Bruce spanworm, Operophtera bruceata, in the Northeastern United States, assessed with novel microsatellites and SNPs. Bulletin of Entomological Research
- Holman, Garth; Del Tredici, Peter; Havill, Nathan; Lee, Nam Sook; Cronn, Richard; Mathews, Sarah; Raubeson, Linda; Campbell, Christopher S. 2017. A new species and introgression in eastern Asian hemlocks (Pinaceae: Tsuga). Systematic Botany. 42(4): 733-746.
- 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
- Havill, Nathan P. ; Shiyake, Shigehiko; Lamb Galloway, Ashley; Foottit, Robert G.; Yu, Guoyue; Paradis, Annie; Elkinton, Joseph; Montgomery, Michael E.; Sano, Masakazu; Caccone, Adalgisa. 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(9): 2065-2080.
- Arsenault, Arielle L.; Havill, Nathan P. ; Mayfield, Albert E.; Wallin, Kimberly F. 2015. Behavioral Responses of Laricobius spp. and Hybrids (Coleoptera: Derodontidae) to Hemlock Woolly Adelgid and Adelgid Host Tree Odors in an Olfactometer. Environmental Entomology. 44(6): 1562-1570.
- Favret, Colin; Havill, Nathan P. ; Miller, Gary L.; Sano, Masakazu; Victor, Benjamin. 2015. Catalog of the adelgids of the world (Hemiptera, Adelgidae). ZooKeys. 534(2): 35-54.
- Fischer, Melissa J.; Brewster, Carlyle C.; Havill, Nathan P.; Salom, Scott M.; Kok, Loke T. 2015. Assessment of the potential for hybridisation between Laricobius nigrinus (Coleoptera: Derodontidae) and Laricobius osakensis, predators of the hemlock woolly adelgid (Hemiptera: Adelgidae). Biocontrol Science and Technology. 25(12): 1467-1482.
- Fischer, Melissa J.; Havill, Nathan P.; Brewster, Carlyle C.; Davis, Gina A.; Salom, Scott M.; Kok, Loke T. 2015. Field assessment of hybridization between Laricobius nigrinus and L. rubidus, predators of Adelgidae. Biological Control. 82: 1-6.
- Fischer, Melissa J.; Havill, Nathan P.; Jubb, Carrie S.; Prosser, Sean W.; Opell, Brent D.; Salom, Scott M.; Kok, Loke T. 2014. Contamination delays the release of Laricobius osakensis for biological control of hemlock woolly adelgid: cryptic diversity in Japanese Laricobius spp. Southeastern Naturalist. 13(special issue 6): 178-191.
- Havill, Nathan P.; Vieira, Ligia C.; Salom, Scott M. 2014. Biology and control of hemlock woolly adelgid. FHTET-2014-05. Morgantown, WV: U.S. Department of Agriculture, Forest Service, Forest Health Technology Enterprise Team. 21 p.
- Havill, Nathan P.; Vieira, Ligia C.; Salom, Scott M. 2014. Hemlock woolly adelgid: a threat to eastern forests. National Woodlands. 37(2): 14-15.
- Jones, C.E.; Havill, Nathan P. ; Hanula, James; Braman, S.K. 2014. Post release recovery of hemlock woolly adelgid predators in the North Georgia mountains.. Journal of Entomological Science. 49(4): 383-400.
- Mayfield, Albert E.; Reynolds, Barbara C.; Coots, Carla I.; Havill, Nathan P.; Brownie, Cavell; Tait, Andrew R.; Hanula, James L.; Joseph, Shimat V.; Galloway, Ashley B. 2014. Establishment, hybridization and impact of Laricobius predators on insecticide-treated hemlocks: Exploring integrated management of the hemlock woolly adelgid. Forest Ecology and Management
- Cook, Stephen P.; Merickel, Frank ; Ward, Karen ; Havill, Nathan. 2013. First report of Adelges abietis (Linneaus) (Hemiptera: Adelgidae) in Idaho. The Pan-Pacific Entomologist
- Ravn, H.P.; Havill, N.P.; Akbulut, 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(6): 401-417.
- 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.
- Davis, G.A.; Havill, N.P.; Adelman, Z.N.; Caccone, A.; Kok, L.T.; Salom, S.M. 2011. DNA barcodes and molecular diagnostics to distinguish an introduced and native Laricobius (Coleoptera: Derodontidae) species in eastern North America. Biological Control. 58(1): 53-59.
- Havill, Nathan P.; Davis, Gina; Klein, Joanne; Caccone, Adalgisa; Salom, Scott. 2011. Hemlock woolly adelgid biological control: molecular methods to distinguish Laricobius nigrinus, L. rubidus, and their hybrids. 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: 25-28.
- Sano, Masakazu; Havill, Nathan P.; Ozaki, Kenichi. 2011. Taxonomic identity of a galling adelgid (Hemiptera: Adelgidae) from three spruce species in central Japan. Entomological Science. 14: 94-99.
- Yang, Zi-xiang; Chen, Xiao-ming; Havill, Nathan P.; Feng, Ying; Chen, Hang. 2010. Phylogeny of Rhus gall aphids (Hemiptera:Pemphigidae) based on combined molecular analysis of nuclear EF1α and mitochondrial COII genes. Entomological Science. 13(3): 351-357.
- Foottit, R.G.; Maw, H.E.L.; Havill, N.P.; Ahern, R.G.; Montgomery, M.E. 2009. DNA barcodes to identify species and explore diversity in the Adelgidae (Insecta: Hemiptera: Aphidoidea). Molecular Ecology Resources. 9 (Suppl. 1): 188-195.
- 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, Katherine A; Gottschalk, Kurt 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.
- Havill, Nathan P.; Montgomery, Michael; Shiyake, Shigehiko; Lamb, Ashley; Keena, Melody A.; Caccone, Adalgisa. 2009. Hemlock woolly adelgid population genetics. In: McManus, Katherine A; Gottschalk, Kurt 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.
- Havill, Nathan P.; Montgomery, Michael E. 2008. The role of arboreta in studying the evolution of host resistance to the hemlock woolly adelgid. Arnoldia. 65(3): 2-9.
- Butin, Elizabeth; Elkinton, Joseph; Havill, Nathan; Montgomery, Michael. 2003. Comparative impact of Scymnus ningshanensis and Pseudoscymnus tsugae (Coleoptera: Coccinellidae) on the hemlock woolly adelgid. In: Fosbroke, Sandra L.C.; Gottschalk, Kurt W., eds. Proceedings, U.S. Department of Agriculture interagency research forum on gypsy moth and other invasive species 2002; 2002 January 15-18; Annapolis, MD. Gen. Tech. Rep. NE-300. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northeastern Research Station: 14.
- Montgomery, Michael; Havill, Nathan; Cheah, Carole; McClure, Mark; Zilahi-Balogh, Gabriella; Lamb, Ashley; Salom, Scott. 2003. Hemlock woolly adelgid biological control research. In: Fosbroke, Sandra L. C.; Gottschalk, Kurt W., eds. 2002 January 15-18; Annapolis, MD. Gen. Tech. Rep. NE-300. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northeastern Research Station. p.74.
- Butin, Elizabeth; Montgomery, Montgomery; Havill, Nathan; Elkinton, Joseph. 2002. Pre-release host range assessment for classical biological controls: experience with predators for the hemlock woolly adelgid. In: Onken, B.; Reardon, R.; Lashomb, J., eds. The hemlock woolly adelgid in the eastern United States symposium; 2002 February 5-7; East Brunswick, NJ. East Brunswick, NJ: New Jersey Agricultural Experiment Station and Rutgers University: 205-214.
- Montgomery, Michael; Wang, Hongbin; Yao, Defu; Lu, Wenhau; Havill, Nathan; Li, Guangwu. 2002. Biology of Scymnus ningshanensis (Coleoptera: Coccinellidae): A predator of Adelges tsugae (Homoptera: Adelgidae). In: Onken, B.; Reardon, R.; Lashomb, J., eds. The hemlock woolly adelgid in the eastern United States symposium; 2002 February 5-7; East Brunswick, NJ. East Brunswick, NJ: New Jersey Agricultural Experiment Station and Rutgers University: 181-188.
Research Datasets
- Havill, Nathan P. 2019. Microsatellite genotypes for southern pine beetles, Dendroctonus frontalis, from the U.S. and Mexico. Fort Collins, CO: Forest Service Research Data Archive. https://doi.org/10.2737/RDS-2019-0002.
National Research Highlights
Silver flies show promise as potential biological controls of hemlock woolly adelgid
Year: 2017
Forest Service science is identifying potential biological control of hemlock wooly adelgid, an invasive insect that is devastating hemlock trees in the Northeast. Two species of silver flies from the Pacific Northwest are showing promise as potential biological controls of hemlock woolly adelgid in the East. The research has demonstrated that these predators can feed and develop on hemlock pest adelgids in the eastern U.S., and they are able to tolerate environmental conditions at the southern and northern ends of the area invaded by the Japanese adelgids. Efforts are underway to release these flies so they will establish in the eastern U.S. and evaluate their ability to control this invasive pest.
Non-native Insect Hybridization Provides Opportunity for Research
Year: 2016
A recent outbreak in New England of the non-native winter moth from Europe provides an unprecedented opportunity to examine the effects of hybridization because Forest Service scientists discovered that it is hybridizing with the native Bruce spanworm. They developed two new sets of genetic markers that confirmed the presence of multi-generation asymmetric hybridization.
First Release of a New Biological Control of Hemlock Woolly Adelgid
Year: 2015
Forest Service scientists and their research partners tested and released two species of silver flies from the western United States for biological control of hemlock woolly adelgid in Tennessee and New York.