Endogenous and exogenous factors affecting parasitism of gypsy moth egg masses by shape Ooencyrtus kuvanae

Factors affecting the orientation, reproduction, and sex ratio of the egg parasitoid Ooencyrtus kuvanae Howard were examined. Adult females were attracted to airborne volatiles from the egg mass and accessory gland of the primary host, the gypsy moth Lymantria dispar L. Visual cues also affected host selection. Background colors against which egg masses were placed affected oviposition preference. In the absence of egg masses, color variation did not affect wasp behavior. Light is required for parasitism by O. kuvanae. The age and density of both the host and parasitoid affected wasp reproduction and sex ratios. Older egg masses issued relatively fewer wasps and higher proportions of males than did young egg masses. Likewise, wasp reproduction and the proportion of females declined with wasp age. Larger egg masses produced more wasps and lower proportions of males than did smaller egg masses. The number of offspring per female, and the proportion of female offspring, were inversely related to wasp density. Implications to biological control of the gypsy moth and parasitoid ecology are discussed.     

中国舞毒蛾六个地理种群的RAPD分析及SCAR标记构建

舞毒蛾Lymantria dispar L.是世界性农林害虫, 包含不同的亚种, 其中亚洲舞毒蛾的雌蛾具有较强的飞行能力, 已成为国际性的重要检疫性有害生物。然而, 不同舞毒蛾亚种及种群间形态难辨, 因此采用传统的手段鉴别舞毒蛾亚种种群是很困难的。本研究首先采用RAPD标记分析了中国舞毒蛾6个地理种群的遗传多态性。结果表明, 所检测的舞毒蛾种群的遗传分化系数G_st为0.7571, 由此推算出的平均有效迁移数（基因流参数）N_em为0.1604, 说明不同舞毒蛾种群间的遗传分化程度较高, 缺乏广泛的基因流动。本研究在RAPD遗传分析基础之上, 筛选出了4个舞毒蛾种群的特异性遗传位点, 然后对这些特异性位点进行了克隆测序、 序列分析和位点特异性引物设计。结果表明, 其中2个舞毒蛾种群的位点特异性引物可产生序列特征性扩增区域（SCAR）标记。经验证, 这些标记可被用来鉴别特定的舞毒蛾地理种群, 因此有助于对这些舞毒蛾地理种群的分布与扩散进行监测。     

The influence of geographic population,age, and mating status on the flight activity of the Asian gypsy moth <Emphasis Type="Italic">Lymantria dispar</Emphasis> (Lepidoptera: Erebidae) in China

The gypsy moth Lymantria dispar (Linnaeus) is a highly polyphagous herbivore that defoliates a wide variety of tree species. The influence of age and mating status on the flight activity of females was assessed in seven geographically isolated populations from China. We examined flight activity of tethered females using a computer-linked flight mill in the laboratory and found that age significantly influenced flight parameters, including total flight distance, time, and speed. Flight distance peaked in the youngest females (at 1 day of age) and declined with age. No significant differences in flight parameters were found between unmated and mated females. Females from Guizhou showed the strongest flight activity among all seven populations. The total flight distances, flight times, and maximum flight speeds at 1 day of age were 7.50 ± 2.28 km, 2.17 ± 0.51 h, and 6.18 ± 0.51 km/h, respectively. These results provide valuable information regarding the scale and speed of dispersal in hotspots and can improve the management and control of forest damage.     

Predators ofLymantria dispar (Lep. lymantriidae) egg masses: Spatio-temporal variation of their impact during the 1988–89 pest generation in the mamora cork oak forest (Morocco)

In the Mamora cork oak forest (Morocco), oophagous predators of the gypsy mothLymantria dispar (L.) (Lepidoptera: Lymantriidae) are one of the main mortality factors influencing pest population dynamics. The predators destroy egg masses more by disrupting their cohesiveness than by predation. From 1987 to 1990, the impact of oophagous enemies significantly contributed to the collapse of a localized gypsy moth outbreak. Variation of egg mass dislocation intensity was greater between trees than among the different strata of a tree. The number of gypsy moth egg masses and egg predator attacks increased when oaks were large and unhealthy. Forest degradation probably explains why egg mass destruction rates were so high (60 to 90% of the eggs) in the infested forest. Oophagous predators find food and shelter under the dehiscent bark of unhealthy cork oaks.     

Population quality,dispersal and numerical change in the gypsy moth,Lymantria dispar (L.)

Summary First instars from small and large gypsy moth eggs differ significantly in their head capsule width, weight, hatching time and the length of thoracic setae. Pupal weight and the developmental period of immature stages of the gypsy moth originating from small or large eggs do not differ significantly. The mean number of eggs per mass produced by females originating from small eggs is greater than that of females from large eggs although not statistically significant. Highly significant differences in mean egg size of egg masses of each type of female were also observed. The relationship between egg size and dispersal strategies are discussed.Paper No. 2229 Massachusetts Agricultural Experiment Station. University of Massachusetts at Amherst. This research supported (in part) from Experiment Station Project No. 355     

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.     

Spatial analysis of gypsy moth populations in Sardinia using geostatistical and climate models

• 1 Spatial fluctuations of the Sardinian population of the gypsy moth Lymantria dispar (L.) (Lepidoptera: Lymantriidae) were characterized using geostatistical and climate models. Data on gypsy moth egg mass abundance recorded at 282 permanent monitoring sites from 1980 to 2004 were incorporated in a geographic information system with the vegetational, geomorphological and pedological features of the sites.
• 2 Statistical analyses revealed that the relative outbreak frequency was related to the predominant host tree, slope and elevation of the monitoring sites, whereas there was no correlation between outbreak frequency and exposure and soil type.
• 3 By using bioclimatic modelling, probability maps of gypsy moth outbreaks were generated. The model identified a probability surface with climatic conditions favourable to gypsy moth outbreaks and thus potentially subject to defoliation. The maps included 92 sites where outbreaks never occurred, suggesting that the Sardinian climate may not be a determinant factor for gypsy moth outbreaks.
• 4 The geostatistical method cokriging with outbreak frequency as a covariate was found to be the most suitable technique to estimate gypsy moth egg mass abundance. Semivariograms showed spatial correlation of egg mass abundance within the range 18.5–53 km. The results obtained were used to create regional gypsy moth distribution maps by cokriging, which demonstrated the outbreak foci and different infestation levels at each monitoring area. These results can help to delimit the treatment areas and develop rational gypsy moth management programmes.

     

World distribution of female flight and genetic variation in Lymantria dispar (Lepidoptera: Lymantriidae)

Female gypsy moths, Lymantria dispar L., from 46 geographic strains were evaluated for flight capability and related traits. Males from 31 of the same strains were evaluated for genetic diversity using two polymorphic cytochrome oxidase I mitochondrial DNA restriction sites, the nuclear FS1 marker, and four microsatellite loci. Females capable of strong directed flight were found in strains that originated from Asia, Siberia, and the northeastern parts of Europe, but flight capability was not fixed in most strains. No flight-capable females were found in strains from the United States or southern and western Europe. Wing size and musculature were shown to correlate with flight capability and potentially could be used in predicting female flight capability. The mtDNA haplotypes broadly separated the gypsy moth strains into three groups: North American, European/Siberian, and Asian. Specific microsatellite or FS1 alleles were only fixed in a few strains, and there was a gradual increase in the frequency of alleles dominant in Asia at both the nuclear and microsatellite loci moving geographically from west to east. When all the genetic marker information was used, 94% of the individuals were accurately assigned to their broad geographic group of origin (North American, European, Siberian, and Asian), but female flight capability could not be predicted accurately. This suggests that gene flow or barriers to it are important in determining the current distribution of flight-capable females and shows the need for added markers when trying to predict female flight capability in introduced populations, especially when a European origin is suspected.     

Spatial distribution of the gypsy moth [Lepidoptera: Lymantriidae] and some of its parasitoids within a forest environment

Aspects of the microhabitat distributions of the gypsy moth,Porthetria dispar (L.) (Lepidoptera: Lymantriidae), and some of its parasitoids were investigated in the field by means of sticky panels and gypsy moth egg masses exposed at different heights in trees, by egg masses exposed within forested and cleared areas, and by gypsy moth pupal collections from different heights in trees.Ooencyrtus kuwanai (Howard)(Hymenoptera: Encyrtidae), Apanteles melanoscelus Ratzeburg (Hymenoptera: Braconidae), and gypsy moths were caught most frequently on stickly panels placed in upper portions of trees. In contrast,Apanteles laeviceps Ashmead, a parasitoid of cutworms, was most often caught near the forest floor.O. kuwanai attacked equally egg masses exposed at different heights in trees, but parasitized those in a clearing less often than those within the forest prosper.Brachymeria intermedia (Nees) (Hymenoptera: Chalcididae) emerged mostly from pupae collected near the tops of trees and not at all from those collected below 5 m. The results are discussed as they relate to field sampling procedures, behavioral activities of gypsy moth and parasitoid adults, and integrated control possibilities for the gypsy moth.     

Evaluation of a nonconventional insecticide and appropriate application timing for destruction of gypsy moth (Lepidoptera: Lymantriidae) egg masses

Two field studies were conducted in 2001-2002 and 2003 to evaluate the effectiveness and appropriate application timing of Golden Pest Spray Oil (GPSO) for destruction of gypsy moth, Lymantria dispar (L.) (Lepidoptera: Lymantriidae), egg masses in Wisconsin. GPSO is a commercially available, registered pesticide that is predominantly comprised of a soybean-oil base (93%); its primary mode of action is by means of suffocation. Because gypsy moth spends the majority (>75%) of its life cycle in the egg stage (August-April), the potential utility of this product by arborists, city foresters, landscapers, and homeowners is high, especially because GPSO is a United States Environmental Protection Agency registered, nonconventional pesticide that is considered relatively nontoxic. When GPSO was applied at a 1:1 ratio with water, >96% control of gypsy moth egg masses was achieved, regardless of application timing (October, 3 d before egg hatch).     

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.     

Rapid identification of the Asian gypsy moth and its related species based on mitochondrial DNA

The gypsy moth—Lymantria dispar (Linnaeus)—is a worldwide forest defoliator and is of two types: the European gypsy moth and the Asian gypsy moth. Because of multiple invasions of the Asian gypsy moth, the North American Plant Protection Organization officially approved Regional Standards for Phytosanitary Measures No. 33. Accordingly, special quarantine measures have been implemented for 30 special focused ports in the epidemic areas of the Asian gypsy moth, including China, which has imposed great inconvenience on export trade. The Asian gypsy moth and its related species (i.e., Lymantria monocha and Lymantria xylina) intercepted at ports are usually at different life stages, making their identification difficult. Furthermore, Port quarantine requires speedy clearance. As such, it is difficult to identify the Asian gypsy moth and its related species only by their morphological characteristics in a speedy measure. Therefore, this study aimed to use molecular biology technology to rapidly identify the Asian gypsy moth and its related species based on the consistency of mitochondrial DNA in different life stages. We designed 10 pairs of specific primers from different fragments of the Asian gypsy moth and its related species, and their detection sensitivity met the need for rapid identification. In addition, we determined the optimal polymerase chain reaction amplification temperature of the 10 pairs of specific primers, including three pairs of specific primers for the Asian gypsy moth (L. dispar asiatic), four pairs of specific primers for the nun moth (L. monocha), and three pairs of specific primers for the casuarina moth (L. xylina). In conclusion, using our designed primers, direct rapid identification of the Asian gypsy moth and its related species is possible, and this advancement can help improve export trade in China.     

Underestimated mitochondrial diversity in gypsy moth Lymantria dispar from Asia

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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.     

Interpopulation quality in gypsy moths with implications for success of two pupal parasitoids: Brachymeria intermedia (Nees) and Coccygomimus turionellae (L.)*

Abstract.

• 1 Gypsy moth egg masses were collected from innocuous, release and outbreak populations and reared in the laboratory on synthetic diet under identical conditions.
• 2 Outbreak population gypsy moths hatched sooner, were smaller and less fecund than innocuous or release gypsy moths, but had a higher concentration of total carbohydrates in their haemolymph.
• 3 Pupae from each population source were submitted to parasitization by two pupal parasitoids. Emerging B.intermedia, an established parasitoid of the gypsy moth associated with outbreak populations, were largest on outbreak source gypsy moths. C. turionellae, not a usual parasitoid of the gypsy moth, were largest when emerging from innocuous or release population gypsy moths. Implications for population dynamics of the gypsy moth are discussed.

     

Pheromone‐trapping the nun moth,Lymantria monacha (Lepidoptera: Lymantriidae) in Inner Mongolia,China

The nun moth, Lymantria monacha L., is one of the most important defoliators of Eurasian coniferous forests. Outbreaks during 2011–2015 in the natural/planted larch, and larch‐birch mixed forests of the Greater Khingan Range in Inner Mongolia, China, caused tremendous timber losses from severe defoliation and tree mortality. A series of trapping experiments were conducted in these outbreak areas to evaluate the efficacy of a synthetic species‐specific pheromone lure based on the female pheromone blend of European nun moth populations. Our results clearly show that the nun moth in Inner Mongolia is highly and specifically attracted to this synthetic pheromone, with few gypsy moths (Lymantria dispar) captured. Flight activity monitoring of L. monacha male moths using pheromone‐baited Unitraps at 2 locations during the summer of 2015 indicated that the flight period started in mid‐July, peaking in early August at both locations. Based on male moth captures, there was a strong diurnal rhythm of flight activity throughout the entire scotophase, peaking between 22:00 and 24:00. Unitraps and wing traps had significantly and surprisingly higher catches than the gypsy moth traps. Unitraps fastened to tree trunks 2 m above ground caught significantly more male moths than those at the ground level or at 5 m height. Male L. monacha moths can be attracted to pheromone‐baited traps in open areas 150–200 m distant from the infested forest edge. Our data should allow improvement on the performance of pheromone‐baited traps for monitoring or mass‐trapping to combat outbreaks of this pest in northeastern China.     

Inheritance of female flight in Lymantria dispar (Lepidoptera: Lymantriidae)

A clinal female flight polymorphism exists in the gypsy moth, Lymantria dispar L., where female flight diminishes from east to west across Eurasia. A Russian population where females are capable of sustained ascending flight and a North American population with females incapable of flight were crossed: parentals, reciprocal F(1) hybrids, double reciprocal F(2) hybrids, and all possible backcrosses to both the parental lines were compared. Heritabilities were estimated using a threshold model, female offspring on female parent regressions, and joint-scaling analyses. Heritability of female flight capability measured using a free flight test was at least 0.60, and variation in wing size, muscle strength, and flight behaviors contributed to the flight polymorphism. Relative wing size varied continuously and had a heritability of 0.70. Environmental variation accounted for >90% of the variation in female preflight weight and relative flight muscle strength, as estimated by an inverted female's ability to right herself. Preflight walking behavior and early deposition of eggs were each inherited through a single gene with two co-dominant alleles. There was no evidence for sex-linkage or maternal effects in female flight capability or associated traits. Continued vigilance to exclude and eradicate introductions of strains capable of female flight in North America is warranted even in areas where no females fly, because some of the alleles needed for full flight capability may not be present in the North American populations, and some flight capability is maintained in the hybrids that could increase the rate of spread of L. dispar.     

The effect of flight on reproduction in an outbreaking forest lepidopteran

Post‐flight reproductive investment by female insects may be limited as a result of a trade‐off in resource allocation between flight and reproduction. Outbreaking forest pests reduce their habitat quality as a result of severe defoliation when population densities are high. Female relocation to better‐quality habitats can increase offspring survival but reduce their reproductive fitness through flight. In the present study, the effect of flight on the capacity of female Choristoneura conflictana (Walker) (Lepidoptera: Tortricidae) to mate and produce eggs is examined. Females are flown on flight mills, and the subsequent reproductive capacity of each moth is assessed through measures of mating success and egg production. There is no effect of flight on commencement or the duration of mating. Although flight does not affect egg production directly, energy expenditure as a result of flight (as measured by weight loss) shows a negative correlation to potential fecundity, possibly indicating the resorption of eggs in some females. The effect of female size on fecundity is dependent on mating status, suggesting that energy allocated to reproduction is not dependent on flight treatment. Female moth longevity also has a significant effect on egg production but is dependent on flight and mating treatments. There is a relationship between energy expenditure to flight and reproduction in C. conflictana. Females that fly away from dense populations may produce fewer offspring, although this cost may be mitigated by improved offspring survivorship in less defoliated habitats.     

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.     

Geographical variation in the periodicity of gypsy moth outbreaks

The existence of periodic oscillations in populations of forest Lepidoptera is well known. While information exists on how the periods of oscillations vary among different species, there is little prior evidence of variation in periodicity within the range of a single Lepidopteran species. The exotic gypsy moth is an introduced foliage-feeding insect in North America. Its populations are characterized by high amplitude oscillations between innocuously low densities and outbreak levels during which large regions of forests are defoliated. These outbreaks are recognized to arise periodically with considerable synchrony across much of the gypsy moth's North American range. Our analysis indicates that gypsy moth outbreaks in North America are periodic but they exhibit two dominant periodicities: a primary period of 8–10 yr (as previously reported) and a secondary period of 4–5 yr (a new finding in this study). The outbreak periodicity varied geographically and this variation was associated with forest type. We found that in the most susceptible forest types, those on xeric sites where oak is often mixed with pines, outbreak periodicity had a more dominant 5-yr period while in forest types characteristic of more mesic sites where oak was mixed with maples and other species, cycles were clearly operating on a 10-yr period.