Seasonal differences in the timing of flight between the invasive winter moth and native Bruce spanworm promotes reproductive isolation.

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.

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.

Successful establishment, spread, and impact of the introduced parasitoid Spathius galinae (Hymenoptera: Braconidae) on emerald ash borer (Coleoptera: Buprestidae) populations in postinvasion forests in Michigan.

Spathius galinae is a larval parasitoid native to the Russian Far East that was approved for release in the United States in 2015 for biological control of the emerald ash borer (EAB), Agrilus planipennis, an invasive beetle from Asia responsible for widespread mortality of ash trees (Fraxinus spp.) in North America. From 2015 to 2017, 1,340-1,445 females of S. galinae along with males were released into each release plot, paired with a nonrelease control plot (1-12.5 km apart), at 6 postinvasion forested sites containing abundant pole-sized ash trees in Michigan. By 2018, S. galinae had spread to all but one control plot. Based on the first year that S. galinae was found in trees in each control plot and the distances of those trees to the parasitoid release point within each site, we estimated that S. galinae spread at 3.7 (±1.9) km per year after its initial releases in 2015. The proportion of sampled trees with S. galinae broods, brood densities within sampled trees, and parasitism of EAB larvae increased sharply in both control and release plots after the last field releases in 2017, with the highest parasitism rates (42.8-60.3%) in 2020. Life table analysis showed that S. galinae alone reduced EAB's net population growth rate by 35-55% across sites from 2018 to 2020. These results demonstrate that S. galinae has established an increasing population in Michigan and now plays a significant role in reducing EAB populations in the area.

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.

Tracking the push towards extinction: combining dispersal and management data to monitor Asian longhorned beetle eradication in the U.S.

Introduction: Based on the threat posed by the Asian longhorned beetle (Anoplophora glabripennis Motschulsky), many countries including the United States have adopted policies of eradication. The eradication of infestations that cover hundreds of square kilometers can require multiple visual surveys of millions of individual trees. At these scales, eradication may take several decades and span multiple beetle generations. During this period the infestation of new trees adds spatially-explicit risk to the landscape while surveys and the removal of infested trees reduce it. Methods: To track dynamic risk on the landscape we have developed the Asian Longhorned Beetle Hazard Management and Monitoring Tool. The geospatial tool combines data documenting; the locations, levels of infestation, and dates of detection of infested trees; the locations, methods, and timing of survey and host removal activities; and a reconstruction of beetle movement within the infested landscape to generate annual spatial estimates of infestation risk based on the combination of beetle dispersal and survey and host removal activities. Results: The analyses of three eradication programs highlight similar patterns in risk through time with risk peaking at the time infestations are detected and declining as management activities slow beetle spread and reduce risk through surveys. However, the results also highlight differences in risk reduction among the eradication programs associated with differences in beetle dispersal among infestations and the size of the infested landscape, highlighting the importance of applying local information to structure eradication programs. Discussion: The Asian Longhorned Beetle Hazard Management and Monitoring Tool provides a quantitative repeatable approach to tracking changes in infestation risk using local beetle behavior and management efforts. In addition to this, the tool may provide a structure to optimize eradication efforts by allowing managers to estimate expected risk reduction based on proposed survey and host removal strategies.

Real-time geographic settling of a hybrid zone between the invasive winter moth (Operophtera brumata L.) and the native Bruce spanworm (O. bruceata Hulst).

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.

High Rainfall May Induce Fungal Attack of Hemlock Woolly Adelgid (Hemiptera: Adelgidae) Leading to Regional Decline.

Hemlock woolly adelgid (HWA; Adelges tsugae Annand (Hemiptera: Adelgidae)) is the cause of widespread mortality of Carolina and eastern hemlock (Tsuga caroliniana Engelmann and T. canadensis (L.) Carrière) throughout the eastern United States (U.S.). Since its arrival in the northeastern U.S., HWA has steadily invaded and established throughout eastern hemlock stands. However, in 2018, anecdotal evidence suggested a sharp, widespread HWA decline in the northeastern U.S. following above-average summer and autumn rainfall. To quantify this decline in HWA density and investigate its cause, we surveyed HWA density in hemlock stands from northern Massachusetts to southern Connecticut and analyzed HWA density and summer mortality in Pennsylvania. As native fungal entomopathogens are known to infect HWA in the northeastern U.S. and rainfall facilitates propagation and spread of fungi, we hypothesized high rainfall facilitates fungal infection of aestivating nymphs, leading to a decline in HWA density. We tested this hypothesis by applying a rain-simulation treatment to hemlock branches with existing HWA infestations in western MA. Our results indicate a regional-scale decline and subsequent rebound in HWA density that correlates with 2018 rainfall at each site. Experimental rain treatments resulted in higher proportions of aestivating nymphs with signs of mortality compared to controls. In conjunction with no evidence of increased mortality from extreme winter or summer temperatures, our results demonstrate an indirect relationship between high rainfall and regional HWA decline. This knowledge may lead to better prediction of HWA population dynamics.

Sin Nombre virus prevalence from 2014-2017 in wild deer mice, Peromyscus maniculatus, on five of the California Channel Islands.

Sin Nombre virus (SNV) is a zoonotic virus that is highly pathogenic to humans. The deer mouse, Peromyscus maniculatus, is the primary host of SNV, and SNV prevalence in P. maniculatus is an important indicator of human disease risk. Because the California Channel Islands contain permanent human settlements, receive hundreds of thousands of visitors each year, and can have extremely high densities of P. maniculatus, surveillance for SNV in island P. maniculatus is important for understanding the human risk of zoonotic disease. Despite the importance of surveillance on these heavily utilized islands, SNV prevalence (i.e. the proportion of P. maniculatus that test positive to antibodies to SNV) has not been examined in the last 13-27 years. We present data on 1,610 mice sampled for four consecutive years (2014-2017) on five of the California Channel Islands: East Anacapa, Santa Barbara, Santa Catalina, San Nicolas, and San Clemente. Despite historical data indicating SNV-positive mice on San Clemente and Santa Catalina, we detected no SNV-positive mice on these islands, suggesting very low prevalence or possible loss of SNV. Islands historically free of SNV (East Anacapa, Santa Barbara, and San Nicolas) remained free of SNV, suggesting that rates of pathogen introduction from other islands and/or the mainland are low. Although continued surveillance is warranted to determine whether SNV establishes on these islands, our work helps inform current human disease risk in these locations and suggests that SNV prevalence on these islands is currently very low.

Defoliation severity is positively related to soil solution nitrogen availability and negatively related to soil nitrogen concentrations following a multi-year invasive insect irruption.

Understanding connections between ecosystem nitrogen (N) cycling and invasive insect defoliation could facilitate the prediction of disturbance impacts across a range of spatial scales. In this study we investigated relationships between ecosystem N cycling and tree defoliation during a recent 2015-18 irruption of invasive gypsy moth caterpillars (Lymantria dispar), which can cause tree stress and sometimes mortality following multiple years of defoliation. Nitrogen is a critical nutrient that limits the growth of caterpillars and plants in temperate forests. In this study, we assessed the associations among N concentrations, soil solution N availability and defoliation intensity by L. dispar at the scale of individual trees and forest plots. We measured leaf and soil N concentrations and soil solution inorganic N availability among individual red oak trees (Quercus rubra) in Amherst, MA and across a network of forest plots in Central Massachusetts. We combined these field data with estimated defoliation severity derived from Landsat imagery to assess relationships between plot-scale defoliation and ecosystem N cycling. We found that trees in soil with lower N concentrations experienced more herbivory than trees in soil with higher N concentrations. Additionally, forest plots with lower N soil were correlated with more severe L. dispar defoliation, which matched the tree-level relationship. The amount of inorganic N in soil solution was strongly positively correlated with defoliation intensity and the number of sequential years of defoliation. These results suggested that higher ecosystem N pools might promote the resistance of oak trees to L. dispar defoliation and that defoliation severity across multiple years is associated with a linear increase in soil solution inorganic N.

The Reliability of Genitalia Morphology to Monitor the Spread of the Invasive Winter Moth (Lepidoptera: Geometridae) in Eastern North America.

Winter moth, Operophtera brumata L. (Lepidoptera: Geometridae), causes widespread defoliation in both its native and introduced distributions. Invasive populations of winter moth are currently established in the United States and Canada, and pheromone-baited traps have been widely used to track its spread. Unfortunately, a native species, the Bruce spanworm, O. bruceata (Hulst), and O. bruceata × brumata hybrids respond to the same pheromone, complicating efforts to detect novel winter moth populations. Previously, differences in measurements of a part of the male genitalia called the uncus have been utilized to differentiate the species; however, the accuracy of these measurements has not been quantified using independent data. To establish morphological cutoffs and estimate the accuracy of uncus-based identifications, we compared morphological measurements and molecular identifications based on microsatellite genotyping. We find that there are significant differences in some uncus measurements, and that in general, uncus measurements have low type I error rates (i.e., the probability of having false positives for the presence of winter moth). However, uncus measurements had high type II error rates (i.e., the probability of having false negatives for the presence of winter moth). Our results show that uncus measurements can be useful for performing preliminary identifications to monitor the spread of winter moth, though for accurate monitoring, molecular methods are still required. As such, efforts to study the spread of winter moth into interior portions of North America should utilize a combination of pheromone trapping and uncus measurements, while maintaining vouchers for molecular identification.

Large-scale patterns of seed removal by small mammals differ between areas of low- versus high-wolf occupancy.

Because most tree species recruit from seeds, seed predation by small-mammal granivores may be important for determining plant distribution and regeneration in forests. Despite the importance of seed predation, large-scale patterns of small-mammal granivory are often highly variable and thus difficult to predict. We hypothesize distributions of apex predators can create large-scale variation in the distribution and abundance of mesopredators that consume small mammals, creating predictable areas of high and low granivory. For example, because gray wolf (Canis lupus) territories are characterized by relatively less use by coyotes (C. latrans) and greater use by foxes (Vulpes vulpes, Urocyon cinereoargentus) that consume a greater proportion of small mammals, wolf territories may be areas of reduced small-mammal granivory. Using large-scale, multiyear field trials at 22 sites with high- and low-wolf occupancy in northern Wisconsin, we evaluated whether removal of seeds of four tree species was lower in wolf territories. Consistent with the hypothesized consequences of wolf occupancy, seed removal of three species was more than 25% lower in high-wolf-occupancy areas across 2 years and small-mammal abundance was more than 40% lower in high-wolf areas during one of two study years. These significant results, in conjunction with evidence of seed consumption in situ and the absence of significant habitat differences between high- and low-wolf areas, suggest that top-down effects of wolves on small-mammal granivory and seed survival may occur. Understanding how interactions among carnivores create spatial patterns in interactions among lower trophic levels may allow for more accurate predictions of large-scale patterns in seed survival and forest composition.

Effects of Extreme Low Winter Temperatures on the Overwintering Survival of the Introduced Larval Parasitoids Spathius galinae and Tetrastichus planipennisi: Implications for Biological Control of Emerald Ash Borer in North America.

Climate change has been linked to shifts in the distribution and phenology of species although little is known about the potential effects that extreme low winter temperatures may have on insect host-parasitoid interactions. In late January 2019, northern regions of the United States experienced a severe cold wave caused by a weakened jet stream, destabilizing the Arctic polar vortex. Approximately 3 mo later at six study sites in southern Michigan and three in southern Connecticut, we sampled the overwintering larvae of the emerald ash borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), and two larval parasitoids, Spathius galinae (Hymenoptera: Braconidae) and Tetrastichus planipennisi (Hymenoptera: Eulophidae), that are being introduced as emerald ash borer biocontrol agents in North America. At these nine study sites, emerald ash borer-infested ash trees and/or saplings were debarked and each overwintering emerald ash borer and parasitoid larva was then examined for cold-induced mortality, as indicated by a brown coloration, flaccid, and watery consistency. In early spring in Michigan, we found 4.5-26% of emerald ash borer larvae, 18-50% of S. galinae larvae, and 8-35% of T. planipennisi larvae were killed by cold. In Connecticut where temperatures were more moderate than in Michigan during the 2019 cold wave, <2% of the larval hosts and parasitoids died from cold injury. Our findings revealed that cold-induced mortality of overwintering larvae of emerald ash borer and its larval parasitoids varied by location and species, with higher mortality of parasitoid larvae in most Michigan sites compared to host larvae. The potential impacts of our findings on the management of emerald ash borer using biocontrol are discussed.

A slow opportunist: physiological and growth responses of an obligate understory plant to patch cut harvesting.

Understory light environments change rapidly following timber harvest, and while many understory species utilize and benefit from the additional light, this response is not ubiquitous in shade-obligate species. I examined the effects of patch cut timber harvest on the physiology and growth of an obligate forest understory species to determine if disturbances via timber harvest are physiological stressors or whether such disturbances provide physiological benefits and growth increases in understory species. Forest canopy structure, along with photosynthesis, respiration, water use efficiency, stomatal conductance, and growth rates of American ginseng were quantified one summer before and two summers after patch cut timber harvest. Survival following timber harvest was lower than that observed at undisturbed populations; however, growth of survivors increased post-harvesting, with growth increasing as a function of canopy openness. Light response curves as well as photosynthesis and respiration rates indicated that plants were not well acclimated to higher light levels in the growing season after timber harvest, but rather to two growing seasons after harvest. Relative growth rate formed a positive linear relationship with maximum photosynthesis following timber harvest. My study suggests that ginseng is a "slow opportunist", because while it benefits from sudden light increases, acclimation lags at least one growing season behind canopy changes. American ginseng is surprisingly resilient in the face of a discrete environmental shift and may benefit from forest management strategies that mimic the natural disturbance regimes common in mature forests throughout its range.

"I've tried the switch but he laughs through the tears:" the use and conceptualization of corporal punishment during the Machine Age, 1924-1939.

OBJECTIVE: To examine attitudes, conflicts, images, circumstances, and time-period effects associated with corporal punishment and other forms of adult-to-child violence during the early 20th century in the United States. METHOD: A sample of 147 letters, referencing corporal punishment and dating from 1924 to 1939, were analyzed using both qualitative and quantitative techniques. The letters were addressed to Angelo Patri (1876-1965), a popular child-rearing expert during the interwar years (also known as the Machine Age), and written primarily by middle class parents with everyday worries about child rearing and proper discipline. RESULTS: People who sought advice emphasized the practical significance of corporal punishment over and above the idea that it violated children's rights to be protected against harm. One in four letters cited conflicts with significant others about corporal punishment. Generally, children were perceived as frail, defiant, or feral. Rarely, were they seen as devilish or, conversely, innocent. Children's disobedience and disrespect were cited more than other misbehaviors as reasons for corporal punishment. Age and gender of the focal child varied by time period when letters from the 1920s and 1930s were compared. CONCLUSIONS: A full understanding of parent-to-child violence cannot be achieved without a firm grasp of its genealogy. The growing popularity of child psychology during the Machine Age had a measurable impact on how children were viewed. A utilitarian frame of interpretation was an important part of the everyday "work" associated with child rearing during this time, foreshadowing the tendency today to emphasize efficacy more than rights when evaluating the legitimacy of corporal punishment.