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The European winter moth, Operophtera brumata L. (Lepidoptera: Geometridae), was accidentally introduced to North America on at least 4 separate occasions, where it has been hybridizing with the native Bruce spanworm, O. bruceata Hulst, at rates up to 10% per year. Both species are known to respond to the same sex pheromones and to produce viable offspring, but whether they differ in the seasonal timing of their mating flights is unknown. Therefore, we collected adult male moths weekly along 2 transects in the northeastern United States and genotyped individuals using polymorphic microsatellite markers as males of these 2 species cannot be differentiated morphologically. Along each transect, we then estimated the cumulative proportions (i.e., the number of individuals out of the total collected) of each species on each calendar day. Our results indicate that there are significant differences between the species regarding their seasonal timing of flight, and these allochronic differences likely are acting to promote reproductive isolation between these 2 species. Lastly, our results suggest that the later flight observed by winter moth compared to Bruce spanworm may be limiting its inland spread in the northeastern United States because of increased exposure to extreme winter events., (© The Author(s) 2023. Published by Oxford University Press on behalf of Entomological Society of America. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Hybridization plays an important and underappreciated role in shaping the evolutionary trajectories of species. Following the introduction of a non-native organism to a novel habitat, hybridization with a native congener may affect the probability of establishment of the introduced species. In most documented cases of hybridization between a native and a non-native species, a mosaic hybrid zone is formed, with hybridization occurring heterogeneously across the landscape. In contrast, most naturally occurring hybrid zones are clinal in structure. Here, we report on a long-term microsatellite data set that monitored hybridization between the invasive winter moth, Operophtera brumata (Lepidoptera: Geometridae), and the native Bruce spanworm, O. bruceata, over a 12-year period. Our results document one of the first examples of the real-time formation and geographic settling of a clinal hybrid zone. In addition, by comparing one transect in Massachusetts where extreme winter cold temperatures have been hypothesized to restrict the distribution of winter moth, and one in coastal Connecticut, where winter temperatures are moderated by Long Island Sound, we found that the location of the hybrid zone appeared to be independent of environmental variables and maintained under a tension model wherein the stability of the hybrid zone was constrained by population density, reduced hybrid fitness, and low dispersal rates. Documenting the formation of a contemporary clinal hybrid zone may provide important insights into the factors that shaped other well-established hybrid zones., (© 2022 John Wiley & Sons Ltd.)

Specialized natural enemies have long been used to implement the biological control of invasive insects. Although research tracking populations following biological control introductions has traditionally focused on the impact of the introduced agent, recent studies and reviews have reflected an appreciation of the complex interactions of the introduced specialist agents with native generalist natural enemies. These interactions can be neutral, antagonistic, or complementary. Here we studied the invasive defoliator winter moth (Operophtera brumata) in the Northeast USA to investigate the role of native, generalist pupal predators along with the introduced, host-specific parasitoid Cyzenis albicans. Prior research in Canada has shown that predation of winter moth pupae from native generalists increased after C. albicans was established as a biological control agent. To explain this phenomenon, the following hypotheses were suggested: (H 1 ) parasitoids suppress the winter moth population to a density that can be maintained by generalist predators, (H 2 ) unparasitized pupae are preferred by predators and therefore experience higher mortality rates, or (H 3 ) C. albicans sustains higher predator populations throughout the year more effectively than winter moth alone. We tested these hypotheses by deploying winter moth pupae over 6 years spanning 2005 to 2017 and by modeling pupal predation rates as a function of winter moth density and C. albicans establishment. We also compared predation rates of unparasitized and parasitized pupae and considered additional mortality by a native pupal parasitoid. We found support for the first hypothesis; we detected both temporal and spatial density dependence, but only in the latter years of the study when winter moth densities were low. We found no evidence for the latter two hypotheses. Our findings suggest that pupal predators have a regulatory effect on winter moth populations only after populations have been reduced, presumably by the introduction of the host-specific parasitoid C. albicans., (© 2022 The Ecological Society of America.)

Winter moth, Operophtera brumata, native to Europe, invaded the northeastern United States in the late 1990s, where it caused widespread defoliation of forests and shade trees ranging from 2,266 to 36,360 ha/yr between 2003 and 2015 in Massachusetts. In 2005, we initiated a biological control effort based on the specialist tachinid parasitoid Cyzenis albicans, which had previously been introduced along with the generalist ichneumonid parasitoid Agrypon flaveolatum to control winter moth in Nova Scotia in the 1950s and British Columbia in the 1970s. Due to concerns of possible non-target impacts by A. flaveolatum, we focused entirely on the specialist C. albicans. Each year for 14 yr, we collected several thousand individuals of C. albicans from British Columbia and released them in widely spaced sites in the northeastern United States. As of 2020, we had established C. albicans at 41 of 44 sites from coastal Maine to southeastern Connecticut. By 2016, winter moth densities (pupae/m 2 ) had declined from 100-500 to 0-10 pupae/m 2 at six release sites at least 10 km apart and this was coincident with the onset of 10-40% parasitism. At one site in Wellesley, Massachusetts, the decline occurred in 2012 and winter moth densities have remained low for seven subsequent years. Defoliation in Massachusetts has been reduced to undetectable levels by aerial survey since 2016. DNA sequencing of the barcoding region of the mitochondrial gene CO1 confirmed that all C. albicans reared from winter moth matched the C. albicans collected from Vancouver Island and were distinct from parasitic flies (presumably a native species) reared from a native congener of winter moth, Bruce spanworm (O. bruceata). Successful establishment of C. albicans on winter moth represents a rare, if not the only, example of the biological control of a major forest defoliator that attacks a wide range of tree species anywhere in the world by the establishment of a single specialist natural enemy., (© 2021 by the Ecological Society of America.)

We collected data on mortality of late-instar gypsy moth, Lymantria dispar (L.), from outbreak populations over 4 wk in June 2017 at 10 sites in the New England region of the United States, along with estimated rainfall at these sites. Deposition of airborne conidia of the fungal pathogen, Entomophaga maimaiga Humber, Shimazu & R.S. Soper, was measured at these same sites as well as at seven other locations in New England. We also quantified the geographical distribution of gypsy moth-caused defoliation in New England in 2017 and 2018 from Landsat imagery. Weekly mortality of gypsy moth larvae caused by E. maimaiga correlated with local deposition of conidia from the previous week, but not with rainfall. Mortality from this pathogen reached a peak during the last 2 wk of gypsy moth larval development and always exceeded that caused by LdNPV, the viral pathogen of gypsy moth that has long been associated with gypsy moth outbreaks, especially prior to 1989. Cotesia melanoscela (Ratzeburg) was by far the most abundant parasitoid recovered and caused an average of 12.6% cumulative parasitism, but varied widely among sites. Deposition of E. maimaiga conidia was highly correlated with percent land area defoliated by gypsy moths within distances of 1 and 2 km but was not significantly correlated with defoliation at distances greater than 2 km. This is the first study to relate deposition of airborne conidia of E. maimaiga to mortality of gypsy moths from that agent., (© The Author(s) 2019. Published by Oxford University Press on behalf of Entomological Society of America. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Acoustic signals produced by caterpillars have been documented for over 100 years, but in the majority of cases their significance is unknown. This study is the first to experimentally examine the phenomenon of audible sound production in larval Lepidoptera, focusing on a common silkmoth caterpillar, Antheraea polyphemus (Saturniidae). Larvae produce airborne sounds, resembling ;clicks', with their mandibles. Larvae typically signal multiple times in quick succession, producing trains that last over 1 min and include 50-55 clicks. Individual clicks within a train are on average 24.7 ms in duration, often consisting of multiple components. Clicks are audible in a quiet room, measuring 58.1-78.8 dB peSPL at 10 cm. They exhibit a broadband frequency that extends into the ultrasound spectrum, with most energy between 8 and 18 kHz. Our hypothesis that clicks function as acoustic aposematic signals, was supported by several lines of evidence. Experiments with forceps and domestic chicks correlated sound production with attack, and an increase in attack rate was positively correlated with the number of signals produced. In addition, sound production typically preceded or accompanied defensive regurgitation. Bioassays with invertebrates (ants) and vertebrates (mice) revealed that the regurgitant is deterrent to would-be predators. Comparative evidence revealed that other Bombycoidea species, including Actias luna (Saturniidae) and Manduca sexta (Sphingidae), also produce airborne sounds upon attack, and that these sounds precede regurgitation. The prevalence and adaptive significance of warning sounds in caterpillars is discussed.

Recent attention has focused on the harmful effects of introduced biological control agents on nontarget species. The parasitoid Compsilura concinnata is a notable example of such biological control gone wrong. Introduced in 1906 primarily for control of gypsy moth, Lymantria dispar, this tachinid fly now attacks more than 180 species of native Lepidoptera in North America. While it did not prevent outbreaks or spread of gypsy moth, we present reanalyzed historical data and experimental findings suggesting that parasitism by C. concinnata is the cause of the enigmatic near-extirpation of another of North America's most successful invaders, the browntail moth (Euproctis chrysorrhoea). From a range of approximately 160,000 km2 a century ago, browntail moth (BTM) populations currently exist only in two spatially restricted coastal enclaves, where they have persisted for decades. We experimentally established BTM populations within this area and found that they were largely free of mortality caused by C. concinnata. Experimental populations of BTM at inland sites outside of the currently occupied coastal enclaves were decimated by C. concinnata, a result consistent with our reanalysis of historical data on C. concinnata parasitism of the browntail moth. The role of C. concinnata in the disappearance of browntail moth outside these enclaves has not been reported before. Despite the beneficial role played by C. concinnata in reversing the browntail moth invasion, we do not advocate introduction of generalist biological control agents. Our findings illustrate that the impact of such organisms can be both unpredictable and far-reaching.