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Hemlock woolly adelgid (HWA), Adelges tsugae Annand (Hemiptera: Adelgidae), has devastated eastern hemlock (Tsuga canadensis [L.] Carriere) in a major portion of its native range in eastern North America. Population dynamics of HWA in the absence of predators have been studied for decades. After many years and much effort directed towards rearing and releasing biological control agents to manage HWA, one of these agents, Laricobius nigrinus Fender (Coleoptera: Derodontidae), is now successfully established at significant densities at sites from the southern to the mid-Atlantic states of the eastern U.S. However, high densities of HWA still persist at many locations throughout the region and spread of HWA and associated damage to hemlock continues. Population models for HWA have suggested that even upwards of 90% predation on eggs laid by the overwintering sistens generation will have minimal effect in reducing the population densities of HWA, if HWA are at high density. In this study, we tested the ability of L. nigrinus to reduce HWA densities, and experimentally tested these model predictions to better understand what impact, if any, L. nigrinus has on HWA densities. By using predator exclusion cages at field sites with well-established populations of L. nigrinus, we were able to record HWA densities, fecundity, overwintering mortality, and predation by L. nigrinus, as well as the proportion of branch tips producing new growth on study trees. Using our field-collected data, we refitted the model in ways that allowed us to predict what population densities we could expect for the following summertime progrediens generation given previous HWA density and levels of L. nigrinus. In both years, we found that despite high rates (greater than 80% ovisac predation) of predation by L. nigrinus on uncaged branches compared to caged branches, there were no significant differences in subsequent densities of the HWA spring generation between caged and uncaged treatments, as predicted by our model. In 2018, our field-collected densities of the summer progrediens generation were lower than what was predicted by the model in both predator exclusion treatments, possibly due to the model not incorporating tree health and climatic factors. Simulation models of pest insect populations based on field-collected data such as fecundity, density, overwintering mortality, and predation, could prove to be important in informing researchers and managers about the role of the biological control agent in the population dynamics of the target host. USDA Forest Service, USA [14-CA-11420004-181] This work was funded by a cooperative agreement with the USDA Forest Service, USA (No. 14-CA-11420004-181). We thank R., Reardon, N. Schneeburger, R. Rhea of the Forest Service for arranging and administrating our funding, M. Meyer for helping us find field sites in NJ and R. Mays, A. Dechaine (Virginia Tech) and B. Mudder (USDA Forest Service) for helping collect samples from the field. We thank A. Roehrig, A. Langevin for processing samples. We thank H. Broadley, J. Anderson, and B. Padilla for help with statistical analysis and graphics. We thank J. Chandler, R. Van Driesche, and D. Orwig for reviewing earlier drafts of this MS. Public domain – authored by a U.S. government employee

The emerald ash borer (EAB), Agrilus planipennis Fairmaire, a buprestid beetle native to Asia, has become a serious pest of ash trees (Fraxinus spp.) in North America since the early 2000s. Due to the impracticality of applying insecticides in natural forests, biocontrol is the most viable method to manage EAB in natural ecosystems. Here, we report the first evidence for the establishment and impact of Spathius galinae Belokobylskij & Strazenac, a larval parasitoid first released in North America in 2016 and 2017 at six mixed-hardwood forest sites, in Connecticut, New York, and Massachusetts. We also report current levels of abundance and parasitism of another introduced larval EAB parasitoid, Tetrastichus planipennisi Yang (Hymenoptera: Eulophidae), released in 2015 and 2016 in these same sites. Spathius galinae was recovered at all release sites in 2018, and its density in sampled trees had increased 1.5- to 20-fold (relative to the first postrelease sample year), reaching a final density of 2.3–14.3 broods/m2 of phloem area and causing 13.1–49.2% marginal rate of parasitism at four of the six sites. In contrast, T. planipennisi was only recovered in 2018 at four of the six release sites, and both its density (0.1–2.3 broods/m2 of phloem area) and parasitism (0.1–5.6%) were lower than that of S. galinae throughout the study at the four sites where recoveries were made. Our data fill a critical gap in the development of a biocontrol-based EAB management plan to protect surviving ash trees capable of reaching maturity and producing replacement trees.

Hemlock woolly adelgid (HWA), Adelges tsugae Annand (Hemiptera: Adelgidae), is an invasive pest causing significant mortality to eastern and Carolina hemlock in eastern North America. Since 2003, management of HWA has included targeted release of the HWA predator Laricobius nigrinus Fender (Coleoptera: Derodontidae), native to western North America. Establishment of L. nigrinus at release sites is well documented, but investigations of its impact on HWA populations have been limited. A four-year (2014-2018), two-phase study using predator exclusion cages to assess the impact of L. nigrinus on HWA was conducted at nine previous release sites in the eastern United States. Significantly more HWA sistens ovisacs were disturbed on no-cage and open-cage branches than on caged branches where predators were excluded. Mean disturbance levels on cage, no-cage and open-cage branches was 8, 38, and 27 percent, respectively. Seven of nine sites had a mean HWA ovisac disturbance greater than 50% for at least one year. Winter temperatures were also a significant factor in overall mortality of the sistens generation with a mean of 46% on study branches. Six of nine sites had a mean overall mortality (winter mortality and predation) greater than 80% for at least one year. Larvae of Laricobius spp. were recovered at all sites during this study. Sequencing of the COI gene from recoveries in Phase One (2015 and 2016) indicated that 88% were L. nigrinus and 12% were L. rubidus LeConte. Microsatellite analysis performed during Phase Two (2017 and 2018) indicated that approximately 97% of larval recoveries were L. nigrinus, 2% were hybrids of L. nigrinus and L. rubidus, and 1% were L. rubidus. Results of this study suggest that L. nigrinus can significantly impact the HWA sistens generation ovisacs and continued investment in the use of this species as a biological control is recommended. USDA Forest Service cooperative agreementUnited States Department of Agriculture (USDA)United States Forest Service [14-CA-11420004-028] We gratefully acknowledge the assistance of Dr. Thomas Kuhar and Dr. Douglas Pfieffer (Virginia Tech) for review of this project, as well as Dr. Carlyle Brewster (Virginia Tech) for assistance with data analysis. We thank Dr. Nathan Havill (USDA Forest Service), Kari Stanley (Virginia Tech), and Melissa Fischer (Washington State Department of Natural Resources) for assistance with molecular protocols, as well as Mark Mayer (New Jersey Department of Agriculture), Scott Passwaters (James River State Park), Biff Thompson (Maryland Department of Agriculture), Jesse Webster (Great Smoky Mountains National Park), and the staff at Blackberry Farm (Tennessee) for site assistance. Thank you to Arthur Morgan School, Celo Community, and the Chattahoochee National Forest for permission to use sites in NC and GA. We are grateful to Natalie Morris, Kara Jeffries, Andy Dechaine, Ryan Mays, Ashley Toland, and James Wahls (Virginia Tech), Bryan Mudder and Andrew Tait (USDA Forest Service), David Bechtel, Philip Hensley, Elizabeth McCarty, Forest Palmer, Pat Parkman, and David Paulsen (University of Tennessee), and Peter Menzies, Marika Lapham, and Ivy Haas (University of North Carolina Asheville), for laboratory and field assistance. This work was funded under USDA Forest Service cooperative agreement 14-CA-11420004-028. Public domain – authored by a U.S. government employee