Top-down regulation of hemlock woolly adelgid (Adelges tsugae) in its native range in the Pacific Northwest of North America.

The density of insect herbivores is regulated by top-down factors (e.g., natural enemies), bottom-up effects (e.g., plant defenses against herbivory), or a combination of both. As such, understanding the relative importance of these factors can have important implications for the establishment of effective management options for invasive species. Here, we compared the relative importance of top-down and bottom-up factors on the abundance of hemlock woolly adelgid (HWA), Adelges tsugae. HWA is invasive in eastern North America, but its native range includes the Pacific Northwest of North America where it has co-evolved with western hemlock, Tsuga heterophylla. Eastern hemlock, Tsuga canadensis, can also be found planted in city and park settings in the Pacific Northwest and the presence of both host species allowed us to directly compare the importance of predators (top-down) and host plant resistance (bottom-up) on HWA abundance by placing mesh exclusion bags on branches of both species and monitoring HWA abundance over two years. We found no evidence for bottom-up control of HWA on western hemlock (a native host). HWA established more readily on that species than on eastern hemlock on which it is a major pest in eastern North America. We found strong evidence for top-down control in that both summer and winter-active predators significantly reduced HWA densities on the branches of both tree species where predators were allowed access. These findings support the validity of the biological control program for HWA, the goal of which is to reduce outbreak populations of HWA in eastern North America.

Establishment and postrelease recovery of Laricobius nigrinus and Laricobius osakensis (Coleoptera: Derodontidae), released for biological control of Adelges tsugae (Hemiptera: Adelgidae), in the Northeastern United States.

Hemlock woolly adelgid (HWA), Adelges tsugae Annand, is a major forest pest in the eastern United States responsible for killing millions of eastern hemlock, Tsuga canadensis (L.) Carrière and Carolina hemlock, T. caroliniana Engelmann. The US biological control program for HWA has largely invested in the rearing and release of Laricobius nigrinus Fender and more recently L. osakensis Montgomery and Shiyake. Though the establishment of L. nigrinus has been well-documented in the southern, mid-Atlantic, and coastal portions of the northeastern United States, documentation in interior areas of the northeastern United States is limited. Establishment of L. osakensis in the northeastern United States has not yet been documented. Release locations in the northeastern United States were surveyed for L. nigrinus and L. osakensis establishment to examine the relationship between establishment success and winter temperatures, as winter minimum temperatures likely limit the northern range of introduced Laricobius species. Our results suggest that L. nigrinus establishment is limited by winter minimum temperatures and that the probability of establishment declines as absolute minimum temperature declines. We found L. nigrinus established at sites in Maine, New York, and Pennsylvania, but did not recover any L. nigrinus in Massachusetts, New Hampshire, or Vermont. Similarly, we found L. osakensis established at sites in New York and Pennsylvania and recovered individuals in Maine, though further sampling is necessary to confirm presence of the F3 generation. We also report the first field observation of reproduction of silver flies, Leucotaraxis argenticollis (Diptera: Chamaemyiidae), released predator of HWA, in the eastern United States.

Effects of emamectin benzoate trunk injections on protection of neighboring ash trees against emerald ash borer (Coleoptera: Buprestidae) and on established biological control agents.

The emerald ash borer (EAB), Agrilus planipennis Fairmaire, is the most destructive invasive pest on ash (Fraxinus spp.) in the United States. We determined whether ash trees injected with emamectin benzoate (EB) could protect untreated neighboring ash trees. We also determined whether the selective treatment of ash trees with EB injections had adverse effects on the establishment of introduced larval parasitoids Tetrastichus planipennis Yang and Spathius galinae Belokobylskij & Strazenac. In experiment one, trees were treated with EB and then retreated 3 years later. Five years post initial treatment, we found that 90% of treated ash trees retained healthy crowns, significantly higher than those of untreated control ash trees (16%). For experiment two, trees only received one treatment of EB and after 2 years 100% of treated ash trees retained healthy crowns, significantly higher than those of untreated ash trees (50%). In both experiments, we found that distance from the central EB-treated tree was not a significant predictor for tree health or presence of EAB exit holes. Although distance from the EB-treated trees appeared to have a significant positive relationship with woodpecker feeding signs on neighboring trees, such relationships did not result in significant differences in the proportion of neighboring ash trees retaining healthy crowns between EB treatment and control plots. The introduced EAB parasitoids appeared to have established equally well between treatment and control plots. Findings are discussed in the context of integration of EB trunk injection with biological control for protection of North American ash against EAB.

Establishment and Early Impact of Spathius galinae (Hymenoptera: Braconidae) on Emerald Ash Borer (Coleoptera: Buprestidae) in the Northeastern United States.

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.