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 dataset 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 find that the location of the hybrid zone appears to be independent of environmental variables and maintained under a tension model wherein the stability of the hybrid zone is 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.

Bat aggregational response to pest caterpillar emergence

Moths (Lepidoptera) are major agricultural and forest pests in many parts of the world, including Europe, with many causing great economic damage to crops, horticultural plants, stored items, and wool products. Here, we focus on two ecologically similar inchworms, Operophtera brumata and Erannis defoliaria, known for their high foliage consumption during the spring emergence of caterpillars. We hypothesise that bats could play a role in reducing pests such as caterpillars by switching to this abundant emerging prey. At two infested and one control forest sites, caterpillars were sampled during spring to determine levels of infestation. At the same time, bat flight activity was monitored during the peak in caterpillar abundance. During the spring caterpillar outbreak, we collected faecal samples of forest-dwelling bats capable of using gleaning. The majority of samples were positive for our focus species, being 51.85% for O. brumata and 29.63% for E. defoliaria faecal samples. The foraging activity of two gleaning bats, Myotis nattereri and Myotis bechsteinii, increased at both infested sites, but not at the control site, during caterpillar emergence, as did foraging of Plecotus auritus/austriacus, which used both gleaning and aerial hawking. We conclude that both specialists and occasional gleaners, which prefer different prey but are able to switch their foraging strategies, aggregate at sites during pest emergence and, as such, our results confirm the high potential of bats to reduce numbers of pest species such as caterpillars.

Trends in Outbreaks of Defoliating Insects Highlight Growing Threats for Central European Forests and Implications for Eastern Baltic Region

To identify general patterns in the effect of climate-driven changes in the outbreak frequency of forest defoliating species, we examined 60 years of records (1950–2010) of outbreaks of five defoliating species. Data on Lymantria dispar, Lymantria monacha, Bupalus piniarius, Panolis flammea, and Operophtera brumata from five Central European countries (Slovakia, Czech Republic, Austria, Hungary, and Germany), where the current climate is comparable with the projections of climate for the Eastern Baltic region by the end of the 21st century, were analyzed. Time series approach was applied to estimate the linkage between outbreaks and climate warming. Mean annual, summer, and winter deviations for the period of 1850 to 1900 were assessed as proxies of warming. To estimate the legacy effect, warming proxies were lagged by one year. Among those tested, warming proxies showed a linkage with outbreaks. Three significant outbreaks occurred in the analyzed period (at the beginning and end of the period). During the middle part of the analyzed period, the frequency and magnitude of outbreaks were low, implicating a higher insect outbreak risk with warming in Central Europe. In the latter part of the analyzed period, more frequent yet smaller outbreaks occurred, which supports the outbreak linkage with one-year lag, summer, and annual temperatures.

Seasonal and elevational variability in the induction of specialized compounds from mountain birch (Betula pubescens var. pumila) by winter moth larvae (Operophtera brumata)

The mountain birch (Betula pubescens var. pumila (L.)) forest in the Subarctic is periodically exposed to insect outbreaks, which are expected to intensify due to climate change. To mitigate abiotic and biotic stresses, plants have evolved chemical defenses, including volatile organic compounds (VOCs) and non-volatile specialized compounds (NVSCs). Constitutive and induced production of these compounds, however, are poorly studied in Subarctic populations of mountain birch. Here, we assessed the joint effects of insect herbivory, elevation, and season on foliar VOC emissions and NVSC contents of mountain birch. VOCs were sampled in situ by an enclosure technique and analyzed by gas chromatography–mass spectrometry. NVSCs were analyzed by liquid chromatography–mass spectrometry using an untargeted approach. At low elevation, experimental herbivory by winter moth larvae (Operophtera brumata) increased emissions of monoterpenes and homoterpenes over the three-week feeding period, and sesquiterpenes and green leaf volatile in the end of the feeding period. At high elevation, however, herbivory augmented only homoterpene emissions. The more pronounced herbivory effects at low elevation were likely due to higher herbivory intensity. Of the individual compounds, linalool, ocimene, 4,8-dimethylnona-1,3,7-triene, 2-methyl butanenitrile, and benzyl nitrile were among the most responsive compounds in herbivory treatments. Herbivory also altered foliar NVSC profiles at both low and high elevations, with the most responsive compounds likely belonging to fatty acyl glycosides and terpene glycosides. Additionally, VOC emissions from non-infested branches were higher at high than low elevation, particularly during the early season, which was mainly driven by phenological differences. VOC emissions varied substantially over the season, largely reflecting the seasonal variations in temperature and light levels. Our results suggest that if insect herbivory pressure continues to rise in the mountain birch forest with ongoing climate change, it will significantly increase VOC emissions with important consequences for local trophic interactions and climate.

Four times out of Europe: serial invasions of the winter moth, Operophtera brumata, to North America

Reconstructing the geographic origins of invasive species is critical for establishing effective management strategies. Frequently, molecular investigations are undertaken when the source population is not known, however; these analyses are constrained both by the amount of diversity present in the native region and by changes in the genetic background of the invading population following bottlenecks and/or hybridization events. Here we explore the geographical origins of the invasive winter moth (Operopthera brumata L.) that has caused widespread defoliation to forests, orchards, and crops in four discrete regions: Nova Scotia, British Columbia, Oregon, and the northeastern United States. It is not known whether these represent independent introductions to North America, or “stepping stone” spread among regions. Using a combination of Bayesian assignment and approximate Bayesian computation methods, we analyzed a population genetic dataset of 24 polymorphic microsatellite loci. We estimate that winter moth was introduced to North America on at least four occasions, with the Nova Scotian and British Columbian populations likely being introduced from France and Sweden, respectively; the Oregonian population likely being introduced from either the British Isles or northern Fennoscandia; and the population in the northeastern United States likely being introduced from somewhere in Central Europe. To our surprise, we found that hybridization has not played a large role in the establishment of winter moth populations even though previous reports have documented widespread hybridization between winter moth and a native congener. We discuss the impact of genetic bottlenecks on analyses meant to determine region of origin.

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.

Predicting the invasion range for a highly polyphagous and widespread forest herbivore

Here we compare the environmental niche of a highly polyphagous forest Lepidoptera species, the winter moth (Operophtera brumata), in its native and invaded range. During the last 90 years, this European tree folivore has invaded North America in at least three regions and exhibited eruptive population behavior in both its native and invaded range. Despite its importance as both a forest and agricultural pest, neither the potential extent of this species’ invaded range nor the geographic source of invading populations from its native range are known. Here we fit a climatic niche model, based on the MaxEnt algorithm, to historical records of winter moth occurrence in its native range and compare predictions of suitable distributions to records from the invaded range. We modeled this distribution using three spatial bins to overcome sampling bias for data obtained from public databases and averaged the multi-continental suitable habitat prediction. Results indicate that this species is distributed across a wide range of climates in its native range but occupies a narrower range in its invaded habitat. Furthermore, the lack of a close fit between climatic conditions in parts of its invaded range and its known native range suggests the possibility that this species has adapted to new climatic conditions during the invasion process. These models can be used to predict suitable habitats for winter moth invasions worldwide and to gain insight into possible origins of North American populations.