Long-distance dispersal of the gypsy moth (Lepidoptera: Lymantriidae) facilitated its initial invasion of Wisconsin

Gypsy moth (Lymantria dispar L.) spread is dominated by stratified dispersal, and, although spread rates are variable in space and time, the gypsy moth has invaded Wisconsin at a consistently higher rate than in other regions. Allee effects, which act on low-density populations ahead of the moving population that contribute to gypsy moth spread, have also been observed to be consistently weaker in Wisconsin. Because a major cause of an Allee effect in the gypsy moth is mate-finding failure at low densities, supplementing low-density populations with immigrants that arrive through dispersal may facilitate establishment and consequent spread. We used local indicator of spatial autocorrelation methods to examine space-time gypsy moth monitoring data from 1996 to 2006 and identify isolated, low-density colonies that arrived through dispersal. We measured the distance of these colonies from the moving population front to show that long-distance dispersal was markedly present in earlier years when Wisconsin was still mainly uninfested. Recently, however, immigrants arriving through long-distance dispersal may no longer be detected because instead of invading uninfested areas, they are now supplementing high-density colonies. In contrast, we observed no temporal pattern in the distance between low-density colonies and the population front in West Virginia and Virginia. We submit that long-distance dispersal, perhaps facilitated through meteorological mechanisms, played an important role in the spread dynamics of the initial Wisconsin gypsy moth invasion, but it currently plays a lesser role because the portion of Wisconsin most susceptible to long-distance immigrants from alternate sources is now heavily infested.     

Forest edges enhance mate‐finding in the invasive European gypsy moth,Lymantria dispar

Habitat type, fragmentation, and edge effects can play important roles in the mate‐finding abilities of many species. These effects can be particularly pronounced in low‐density populations, which are often found at the margins of species' ranges or at the leading edge of an invasion. The European gypsy moth, Lymantria dispar (L.) (Lepidoptera: Erebidae), is a non‐native insect defoliator in the USA and Canada, where flightless females attract male moths through pheromone production and local extirpation of low‐density populations can be due to mate‐finding failure. To assess the effects of habitat edges on the ability of gypsy moths to find mates, we conducted a release experiment with male gypsy moths using female‐baited trap arrays in fields, at forest edges, and in the forest interior. Reduced mate‐finding was expected in fields and near forest edges based on geographic variation in invasion rates, male flight behavior, and pheromone plume dynamics. However, we found that mate‐finding was highest at forest edges, reduced in fields, and lowest within the forest interior. Within an array, traps closest to the forest edge also had the highest mate‐finding, suggesting that habitat characteristics can influence male flight direction in addition to pheromone cues. These results suggest that a moderate level of forest fragmentation enhances mate‐finding ability in the gypsy moth. Understanding the relationship between habitat heterogeneity and mate‐finding success in invasive species can inform predictions of future spread and assist with management plans that target mating disruption.     

Optimization of pheromone dosage for gypsy moth mating disruption

The effect of aerial applications of the pheromone disparlure at varying dosages on mating disruption in low‐density gypsy moth, Lymantria dispar (L.) (Lepidoptera: Lymantriidae), populations was determined in field plots in Virginia, USA during 2000 and 2002. Six dosages [0.15, 0.75, 3, 15, 37.5, and 75 g active ingredient (AI)/ha] of disparlure were tested during the 2‐year study. A strongly positive dose–response relationship was observed between pheromone dosages and mating disruption, as measured by the reduction in male moth capture in pheromone‐baited traps and mating successes of females. Dosages of pheromone 15 g AI/ha (15, 37.5, and 75 g AI/ha) reduced the mating success of females by >99% and significantly reduced male moth catches in pheromone‐baited traps compared to untreated plots. Pheromone dosages <15 g AI/ha also reduced trap catch, but to a lesser extent than dosages 15 g AI/ha. Furthermore, the effectiveness of the lower dosage treatments (0.15, 0.75, and 3 g AI/ha) declined over time, so that by the end of the study, male moth catches in traps were significantly lower in plots treated with pheromone dosages 15 g AI/ha. The dosage of 75 g AI/ha was initially replaced by a dosage of 37.5 g AI/ha in the USDA Forest Service Slow‐the‐Spread (STS) of the Gypsy Moth management program, but the program is currently making the transition to a dosage of 15 g AI/ha. These changes in applied dosages have resulted in a reduction in the cost of gypsy moth mating disruption treatments.     

Variation in growth and developmental responses to supraoptimal temperatures near latitudinal range limits of gypsy moth Lymantria dispar (L.), an expanding invasive species

Variation in thermal performance within and between populations provides the potential for adaptive responses to increasing temperatures associated with climate change. Organisms experiencing temperatures above their optimum on a thermal performance curve exhibit rapid declines in function and these supraoptimal temperatures can be a critical physiological component of range limits. The gypsy moth, Lymantria dispar (L.) (Lepidoptera: Erebidae), is one of the best‐documented biological invasions and factors driving its spatial spread are of significant ecological and economic interest. The present study examines gypsy moth sourced from different latitudes across its North American range for sensitivity to high temperature in constant temperature growth chamber experiments. Supraoptimal temperatures result in higher mortality in northern populations compared with populations from the southern range extent (West Virginia and coastal plain of Virginia, U.S.A.). Sublethal effects of high temperature on traits associated with fitness, such as smaller pupal mass, are apparent in northern and West Virginia populations. Overall, the results indicate that populations near the southern limits of the range are less sensitive to high temperatures than northern populations from the established range. However, southern populations are lower performing overall, based on pupal mass and development time, relative to northern populations. This suggests that there may be a trade‐off associated with decreased heat sensitivity in gypsy moth. Understanding how species adapt to thermal limits and possible fitness trade‐offs of heat tolerance represents an important step toward predicting climatically driven changes in species ranges, which is a particularly critical consideration in conservation and invasion ecology.     

Effects of SPLAT® GM sprayable pheromone formulation on gypsy moth mating success

Several integrated pest management programs rely on the use of mating disruption tactics to control insect pests. Some programs specifically target non‐native species, such as the gypsy moth, Lymantria dispar (L.) (Lepidoptera: Lymantriidae). We evaluated SPLAT^® GM, a new sprayable formulation of the gypsy moth sex pheromone disparlure, for its ability to disrupt gypsy moth mating. The study was conducted in 2006, 2007, and 2008 in forested areas in Virginia, USA. Mating success of gypsy moth females was reduced by >99% and male moth catches in pheromone‐baited traps by >90%, in plots treated with SPLAT^® GM at dosages ranging from 15 to 75 g of active ingredient (a.i.) ha^?1. Dosage‐response tests conducted in 2008 indicated that SPLAT^® GM applied at a dosage of 7.5 g a.i. ha^?1 was as effective as a 15 g a.i. ha^?1 dosage.     

Ground application of mating disruption against the gypsy moth (Lepidoptera: Erebidae)

The gypsy moth, Lymantria dispar (L.) (Lepidoptera: Erebidae), is a non‐native defoliating insect that continues to expand its range in North America and undergo periodic outbreaks. In management efforts to suppress outbreaks, slow its spread and eradicate populations that arrive outside of the invaded range, aerial deployments of mating disruption tactics and pesticides are generally used. However, in some cases, such as in heavily urbanized areas or other landscapes where aerial deployments are not feasible or permitted, ground applications are required. Ground applications tend to be labour‐intensive to ensure adequate coverage. To better inform optimal deployment of ground applications of mating disruption, we measured the effectiveness of a pheromone formulation designed for ground application, SPLAT® GM, in forested areas of Virginia from 2011 to 2014 using different dosages and number of point applications. We observed that SPLAT® GM applied to the tree trunks at the dosages of 49.4 and 123.6 g AI/ha in 11 × 11 systematic grids (i.e., every 11 m) reduced male trap catch by >90% relative to untreated control plots, which based on previous studies corresponds to >95% reduction in gypsy moth mating success. Our observations suggest that ground applications of gypsy moth mating disruption can be a successful management tool when circumstances require it.     

Forest type affects predation on gypsy moth pupae

Abstract 1 Predation by small mammals has previously been shown to be the largest source of mortality in low‐density gypsy moth, Lymantria dispar (L.), populations in established populations in north‐eastern North America. Fluctuations in predation levels are critical in determining changes in population densities. 2 We compared small mammal communities and levels of predation on gypsy moth pupae among five different oak‐dominated forest types along this insect's western expanding population front in Wisconsin. Comparisons of predator impact can provide critical information for predicting variation in susceptibility among forest types. 3 The results indicated that small mammals caused more mortality than did invertebrates. 4 Both abundance of Peromyscus sp. predators and predation levels were lower in urban and xeric forest types than in mesic sites. 5 These results suggest that, because predation pressures will probably be greater in the mesic sites, gypsy moths may be less likely to develop outbreaks in these habitats, and that defoliation will probably be more frequent in urban and xeric oak‐dominated sites.     

Climate suitability and management of the gypsy moth invasion into Canada

The gypsy moth has become established throughout southern Canada east of Lake Superior where the climate is suitable for the completion of its univoltine life cycle. The spread of the gypsy moth to the north and west in Canada has so far been prevented by climatic barriers and host plant availability as well as by aggressive eradication of incipient populations. Climate change is expected to increase the area of climatic suitability and result in greater overlap with susceptible forest types throughout Canada, especially in the west. At the same time, the gypsy moth is spreading west in the USA into states bordering western Canadian provinces. These circumstances all lead to a greatly increased risk of further invasion into Canadian forests by the gypsy moth. Management actions need to be intensified in different ways in different parts of the country to reduce the impacts of spread in eastern Canada and to prevent the gypsy moth from invading western regions.     

Comparison of methods for deploying female gypsy moths to evaluate mating disruption treatments

1 Mating disruption is the primary tactic used to reduce rates of gypsy moth population spread in the United States Department of Agriculture’s Slow‐the‐Spread of the gypsy moth programme (STS). Because STS targets very low‐density gypsy moth populations within which it is extremely difficult to collect females or egg masses, mating success in native populations cannot be determined. Therefore, the evaluation of mating disruption treatments in field experiments such as those designed to test new formulations and application methods requires deploying and recovering laboratory‐reared female moths to determine mating success. 2 Five methods of deploying females were evaluated for cost, rates of female and egg mass recovery, and female mating success. The deployment methods tested were: modified delta trap, square barrier, single and double trunk bands, and tethered females. 3 Deployment of tethered females had the highest cost and mating success rate, but it did not yield the highest rates of female and egg mass recovery. Deployment of females in delta traps produced the lowest cost and mating success rate, but yielded the highest recovery rate. Neither of these deployment methods is recommended because of unacceptably high cost (tethered female) or low mating success (delta trap). 4 There were no significant differences in cost or mating success among the other three deployment methods. 5 The differences among the square barrier, single trunk band, and double trunk band methods in cost, female and egg mass recovery, and mating success are too small to recommend any one over the others.     

Effects of the 3M™ MEC Sprayable Pheromone® formulation on gypsy moth mating success

The study was conducted during 2000, 2001, 2003 and 2004 in forested areas in Virginia, USA to evaluate the 3M™ MEC-GM Sprayable Pheromone® formulation of the gypsy moth sex pheromone, disparlure, for its ability to disrupt mating in gypsy moth, Lymantria dispar (Lep.: Lymantriidae). Both mating success of gypsy moth females and male moth catches in pheromone-baited traps were significantly reduced in plots treated with the 3M™ MEC-GM formulation at dosages ranging from 15 to 75 g of active ingredient/ha. However, the 3M™ MEC-GM formulation reduced trap catch to a lesser extent than did the currently registered Hercon Disrupt® II plastic flakes used as a positive control and applied at similar or lower dosages. Furthermore, the effectiveness of the 3M™ sprayable formulation declined through time, so that by the end of the male flight season, male moth catches in traps were significantly higher than in plots treated with Hercon plastic flakes. Based on the reported results, 3M™ MEC-GM Sprayable Pheromone® formulation was never integrated into the operational treatment projects of USDA Forest Service Cooperative Slow-the-Spread of the Gypsy Moth management programme.     

Factors influencing the effectiveness of an attracticide formulation against the Oriental fruit moth, Grapholita molesta

An attracticide formulation, LastCall?OFM, was tested against the Oriental fruit moth Grapholita molesta (Busck) (Lepidoptera: Tortricidae) in replicated small plot field trials in apple, Malus domestica (Borkhausen), orchards in South‐eastern Pennsylvania, USA. Attracticide treatments were applied using a calibrated hand pump, and treated plots were compared to similar untreated plots. Male moth activity was monitored using virgin female‐baited traps, and the potential for reduction in mating activity was assessed using sentinel virgin females. A comparison of application rates showed that 1500 droplets per ha of the attracticide formulation was as effective as 3000 droplets per ha, and both application rates reduced captures in synthetic pheromone‐baited traps for prolonged periods. Droplets placed either at high or low positions within the canopy significantly reduced trap capture and mating with sentinel females. In addition, the only sentinel females that mated in the treated plots were located in the untreated portion of the tree canopy. Mate finding behaviour was equally disrupted by formulations with and without insecticide. Therefore, under the test conditions, the mechanism by which the attracticide formulation worked was by disruption of male orientation, and not by the removal of males due to insecticide poisoning. Two field cage experiments tested the impact of population density on the competitiveness of the attracticide formulation compared to virgin females. A significant proportion of males were captured in female‐baited traps at the highest female‐to‐droplet ratio tested. Equal proportions of males were captured in attracticide‐baited traps at male moth densities of 10, 20, 40, and 80 males per cage. These results clarify some of the factors influencing the effectiveness and possible mechanisms of an attracticide management tactic against the Oriental fruit moth.     

Dispersion in time and space affect mating success and Allee effects in invading gypsy moth populations

1. Understanding why invading populations sometimes fail to establish is of considerable relevance to the development of strategies for managing biological invasions. 2. Newly arriving populations tend to be sparse and are often influenced by Allee effects. Mating failure is a typical cause of Allee effects in low-density insect populations, and dispersion of individuals in space and time can exacerbate mate-location failure in invading populations. 3. Here we evaluate the relative importance of dispersal and sexual asynchrony as contributors to Allee effects in invading populations by adopting as a case study the gypsy moth (Lymantria dispar L.), an important insect defoliator for which considerable demographic information is available. 4. We used release-recapture experiments to parameterize a model that describes probabilities that males locate females along various spatial and temporal offsets between male and female adult emergence. 5. Based on these experimental results, we developed a generalized model of mating success that demonstrates the existence of an Allee threshold, below which introduced gypsy moth populations are likely to go extinct without any management intervention.     

Benefits and harm caused by the introduced generalist tachinid, <Emphasis Type="Italic">Compsilura concinnata</Emphasis>, in North America

We review North American research on Compsilura concinnata (Meigen), a highly generalist tachinid parasitoid that was introduced in 1906 to control two invasive forest insects: gypsy moth and browntail moth. The impact on gypsy moth in natural populations in North America has long thought to be minor, a view confirmed by recent work in Canada and Wisconsin, USA. However, research with experimentally created populations of gypsy moth suggests that it may be more important than previously realized. Studies on browntail moth in both naturally occurring and experimental populations suggest that C. concinnata is probably the main reason browntail moth disappeared from most of its former range in North America. Research on giant silk moths suggests that C. concinnata has become the major source of mortality among several species and may be responsible for the notable decline in their densities that has occurred over the last century. C. concinnata has been touted as a premier example of the generalist natural enemy that should be avoided in classical biological control introductions, yet in the case of browntail moth its effect has been extremely beneficial.     

Effects of intentional gaps in spray coverage on the efficacy of gypsy moth mating disruption

Abstract:  The study was conducted during 2001 and 2002 in forested areas in Virginia, US to examine the effects of gaps in coverage of pheromone on gypsy moth, Lymantria dispar (L.) (Lep., Lymantriidae), mating disruption. Gypsy moth male moth catches in pheromone-baited traps were significantly reduced in plots treated with the gypsy moth sex pheromone, disparlure, at an overall application rate of 37.5 g of active ingredient (AI)/ha but with untreated gaps of 30 or 90 m between 30-m wide treated swaths. In one of the two plots with 90 m gaps, significantly more males were captured in traps in the untreated areas compared with the treated areas within the plot. However, in another plot, significant differences in trap catches between treated and untreated areas were not observed. No difference in male moth catches in the pheromone-baited traps was observed between treated and untreated areas within the plots treated with 30 m gaps. Female mating success did not differ significantly between treated and untreated areas within the one plot in which it was measured. These results suggest that it may be possible to lower costs associated with gypsy moth mating disruption applications by alternating treated and untreated swaths, which would reduce flight time and fuel costs, without a reduction in efficacy.     

Effects of alternative prey on predation by small mammals on gypsy moth pupae

Previous work shows that predation by small mammals is a dominant cause of mortality of low-density gypsy moths in North America and that declines in small mammal density result in increases in gypsy moth density. Here we examined whether predation by small mammals is density dependent by way of a type III functional response, and how predation is influenced by alternative prey. First we showed that the preference of predators for gypsy moth pupae was low compared to other experimental prey items, such as mealworm pupae and sunflower seeds. Predation on gypsy moth pupae was characterized by a type II functional response with percent predation highest at the lowest prey densities, whereas the functional response to sunflower seeds was characterized by a type III functional response in which predation increased with increasing prey density. These results suggest that predation by small mammals is unlikely to stabilize low-density gypsy moth populations.     

Anthropogenic drivers of gypsy moth spread

The gypsy moth, Lymantria dispar (L.), is a polyphagous defoliator introduced to Medford, Massachusetts in 1869. It has spread to over 860,000 km² in North America, but this still only represents ¼ of its susceptible host range in the United States. To delay defoliation in the remaining susceptible host range, the government maintains a barrier zone and a quarantine, reflecting a presumption that anthropogenic factors are important in the spread of gypsy moth. We develop a model framework that relates these factors along with biophysical characteristics to a county’s susceptibility to gypsy moth invasion. We then compile a dataset for counties within 200 km of the infested area and use trap catch data from 1999 to 2007 to estimate the probability of gypsy moth presence. As expected, gypsy moth is more likely to be found close to the population front and to traps that recorded moths in the previous year. However, when controlling for these factors, our most robust finding is that the use of wood for home heating and energy is consistently positively correlated with the presence of gypsy moth. In contrast, the movement of wood products by industry, which is actively regulated by state and federal governments, is rarely correlated with the presence of gypsy moth. This is consistent with effective regulation of the movement of goods by industry, but not by the public. Our findings provide empirical support for the importance and challenge of firewood as a vector for non-native forest insects.     

Prey‐processing by avian predators enhances virus transmission in the gypsy moth

The Lymantria dispar nucleopolyhedrovirus (LdNPV) is one of the most important regulators of gypsy moth populations, but some aspects of its transmission remain poorly understood, particularly its high rate of spatial spread and ability to persist in low‐density populations. We tested the role of predatory birds in the transmission of this virus using experimental gypsy moth populations in an aviary. Predatory birds captured virus‐infected caterpillars and facilitated viral dispersal via two processes: 1) by ingesting infected caterpillars and passing viral occlusion bodies (OBs) through their guts, and 2) by scattering OBs during predator‐specific processing behaviors, a mechanism documented here for the first time. The relative importance of both pathways differed by predator species. After eating virus‐infected gypsy moth larvae, red‐eyed vireos and black‐capped chickadees passed more gypsy moth nucleopolyhedrovirus in feces than did gray catbirds. During prey‐processing, the repetitive beating of caterpillars by red‐eyed vireos, a behavior that was rarely utilized by chickadees and catbirds, resulted in the scattering of infectious virus. Due to the combination of efficient gut passage and virus spread from prey beating, higher rates of transmission occurred in experimental gypsy moth populations exposed to red‐eyed vireos than those exposed to catbirds or chickadees. Our results show that effective virus transmission was achieved when virus was vectored by predatory birds through a combination of both behavioral and physiological traits.     

Elevational gradient in the cyclicity of a forest-defoliating insect

Observed changes in the cyclicity of herbivore populations along latitudinal gradients and the hypothesis that shifts in the importance of generalist versus specialist predators explain such gradients has long been a matter of intense interest. In contrast, elevational gradients in population cyclicity are largely unexplored. We quantified the cyclicity of gypsy moth populations along an elevational gradient by applying wavelet analysis to spatially referenced 31-year records (1975–2005) of defoliation. Based on geographically weighted regression and nonlinear regression, we found either a hump-shaped or plateauing relationship between elevation and the cyclicity of gypsy moth populations and a positive relationship between cyclicity and the density of the gypsy moth’s preferred host-tree species. The potential effects of elevational gradients in the density of generalist predators and preferred host-tree species on the cyclicity of gypsy moth populations were evaluated with mechanistic simulation models. The models suggested that an elevational gradient in the densities of preferred host tree species could partially explain elevational patterns of gypsy moth cyclicity. Results from a model assuming a type-III functional response of generalist predators to changes in gypsy moth density were inconsistent with the observed elevational gradient in gypsy moth cyclicity. However, a model with a more realistic type-II functional response gave results roughly consistent with the empirical findings. In contrast to classical studies on the effects of generalist predators on prey population cycles, our model with a type-II functional response predicts a unimodal relationship between generalist-predator density and the cyclicity of gypsy moth populations.     

Field evaluation of effect of temperature on release of disparlure from a pheromone-baited trapping system used to monitor gypsy moth (Lepidoptera: Lymantriidae)

Traps baited with disparlure, the synthetic form of the gypsy moth, Lymantria dispar (L.) (Lepidoptera: Lymantriidae), sex pheromone are used to detect newly founded populations and estimate population density across the United States. The lures used in trapping devices are exposed to field conditions with varying climates, which can affect the rate of disparlure release. We evaluated the release rate of disparlure from delta traps baited with disparlure string dispenser from 1 to 3 yr across a broad geographic gradient, from northern Minnesota to southern North Carolina. Traps were deployed over approximately 12 wk that coincided with the period of male moth flight and the deployment schedule of traps under gypsy moth management programs. We measured a uniform rate of release across all locations when considered over the accumulation of degree-days; however, due to differences in degree-day accumulation across locations, there were significant differences in release rates over time among locations. The initial lure load seemed to be sufficient regardless of climate, although rapid release of the pheromone in warmer climates could affect trap efficacy in late season. Daily rates of release in colder climates, such as Minnesota and northern Wisconsin, may not be optimal in detection efforts. This work highlights the importance of local temperatures when deploying pheromone-baited traps for monitoring a species across a large and climatically diverse landscape.     

Genetic split between coastal and continental populations of gypsy moth separated by Dinaric Alps

The gypsy moth, a polyphagous herbivore species, infests mainly deciduous trees in the northern hemisphere, being invasive in North America. In Croatia, gypsy moth is infesting both continental and coastal forests, with the Dinaric Alps posing a physical migratory barrier between two regions. During outbreaks, caterpillars cause severe damages in both regions, though with different outbreak dynamics, which suggests genetic differences between populations. Representative populations from these two regions were screened by sequencing a region of the mitochondrial COI gene. Ninety‐nine sequences resulted in seventeen haplotypes, and analyses revealed a significant genetic differentiation between coastal and continental populations, quite likely attributed to geographic isolation and post‐glacial history. This differentiation arises from significantly higher genetic variability in Mediterranean population, indicating their higher adaptability, an intriguing fact in case of possible northward range shift of gypsy moth.