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.     

What causes outbreaks of the gypsy moth in North America?

The gypsy moth has been present in North America for more than 100 years, and in many of the areas where it has become established outbreaks occur with varying degrees of periodicity. There also exists extensive spatial synchrony in the onset of outbreaks over large geographic regions. Density-dependent mortality clearly limits high-density populations, but there is little evidence for strong regulation of low-density populations. Predation by small mammals appears to be the major source of mortality affecting low-density populations, but because these are generalist predators and gypsy moths are a less preferred food item, mammals do not appear to regulate populations in a density-dependent fashion. Instead, predation levels appear to be primarily determined by small mammal abundance, which is in turn closely linked to the production of acorns that are a major source of food for overwintering predator populations. Mast production by host oak trees is typically variable among years, but considerable spatial synchrony in masting exists over large geographic areas. Thus, it appears that the temporal and spatial patterns of mast production may be responsible for the episodic and spatially synchronous behavior of gypsy moth outbreaks in North America. This multitrophic relationship among mast, predators, and gypsy moths represents a very different explanation of forest insect outbreak dynamics than the more widely applied theories based upon predator–prey cycles or feedbacks with host foliage quality. Received: September 8, 1999 / Accepted: September 20, 2000     

Inference of adult female dispersal from the distribution of gypsy moth egg masses in a Japanese city

1 The native range of the gypsy moth Lymantria dispar (L.) spans the temperate forests of Eurasia. Across this region, a clinal female flight polymorphism exists; gypsy moth females in eastern Asia are mostly capable of directed flight, those in western and southern Europe are largely incapable of flight and populations distributed across the centre of the distribution exhibit a range of intermediate flight behaviours. 2 Although information exists about the timing and duration of female flight from laboratory and wind tunnel studies, little or no quantitative data are available on average distances flown by Asian gypsy moth females prior to oviposition in the field. This information is critical for estimating risk of contamination at specific ports and transit terminals, as well as for predicting the spread of populations that might someday invade currently uninfested regions of the world. 3 In the present study, an extensive visual survey of gypsy moth egg masses was conducted during a walk through streets and paths in a 3.92 × 5.76 km area in Kanazawa, Japan. This area consisted of a matrix of urban, agricultural and forest land uses. The distribution of egg masses relative to distances from host forests was used to infer the magnitude of pre‐ovipositional female flight. 4 A total of 3172 egg masses was recorded from surveys conducted during the search of a path totalling 384 km. Within urban areas, egg masses were most abundant in the area <1 km from the edge of forest land. 5 These results suggest that most female gypsy moth flight is limited to the area within 1 km of host forests. They also suggest that shipping containers and other parcels located >1 km from forests are at a much lower risk of contamination with Asian gypsy moth egg masses.     

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.     

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.     

Temporal events of gypsy moth vitellogenesis and ovarian development

Abstract The vitellogenic period of gypsy moth ovarian development starts on day 3 of the pupal stage and continues through adulthood. During this period, rapid increases occur in follicle size, protein content, and wet weight of the ovary. Patency is observed on day 3 of the pupal stage.
Pre-vitellogenic follicles are formed in the last larval stadium. Newly formed follicles detach from the germarium on day 4, and increase rapidly to 140 per ovariole at the end of the last larval stadium. The pre-vitellogenic follicles are uniformly around 50 um in diameter. No vitellogenin is incorporated into the oocytes until the pupal stage.
Polyacrylamide gel electrophosesis (PAGE) in the presence of sodium dodecylsulphate (SDS) analysis of male and female haemolymph samples and vitellogenic ovaries demonstrates the presence of two female-specific subunits of vitellogenin of 180 kD and 160 kD. These proteins are detected only in haemolymph and ovarian extracts of vitellogenic females. The molecular weight of the native protein determined by size exclusion chromatography is approximately 400–420 kD.
A highly sensitive double antibody sandwich enzyme-linked immunosorbent assay (ELISA) was developed to monitor the temporal changes in vitellogenin titre in haemolymph. Vitellogenin production starts on day 2 of the last larval stadium, reaching a maximum level by day 6 of the last larval stadium, and decreasing in the late pupal stage as vitellogenin was internalized into the oocytes. This is the first report of vitellogenin production occurring in the larval stage of a holometabolous insect. The fact that vitellogenin production and uptake occur during different stages of development in the gypsy moth, opens up some interesting questions concerning the underlying regulatory mechanisms controlling each process.     

Underestimated mitochondrial diversity in gypsy moth Lymantria dispar from Asia

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Two-step regulation of sex-pheromone decline in mated gypsy moth females

The regulation of post-mating decline of sex-pheromone in the gypsy moth, Lymantria dispar, was studied. An initial, transient suppression of pheromone production was found to be caused by the introduction of male genitalia into the bursa copulatrix, which results in mechanical pressure being transmitted via innervation of the bursa. However, if sperm was not transferred during mating, pheromone production resumed and females returned to calling behavior. Permanent suppression of pheromone production resulted from an adequate supply of sperm in the spermatheca and could be prevented in females from which spermatheca was removed. During the initial period of suppression of pheromone production females were sexually receptive and could remate. They became nonreceptive only when pheromone production was terminated and oviposition begun.     

Characterisation of three Alphabaculovirus isolates from the gypsy moth,Lymantria dispar dispar (Lepidoptera: Erebidae), in Turkey

In this study, Lymantria dispar dispar larvae, collected from three different localities in Turkey, were examined for the presence of inclusion bodies under phase contrast and electron microscopes. Inclusion bodies from infected larvae were subjected to polymerase chain reaction using the conserved primers for polyhedrin (polh), late expression factor 8 (lef-8) and late expression factor 9 (lef-9) genes. Sequence analysis confirmed that larvae collected from the three different localities contained multiple nucleopolyhedrosis viruses (MNPVs). These isolates were designated LdMNPV-T1, LdMNPV-T2 and LdMNPV-T3. Phylogenetic analyses of these isolates were performed using target genes polh, lef-8 and lef-9. Restriction endonuclease analysis of the three geographic isolates with EcoRI and PstI enzymes demonstrated some differences existed among the isolates. According to the EcoRI profile, the mean estimated size for the complete genome of each isolate (LdMNPV-T1, LdMNPV-T2 and LdMNPV-T3) was calculated to be approximately 170, 153 and 170?kb, respectively. Insecticidal activities of each isolate were tested on L. d. dispar larvae using four different viral concentrations between 10³ and 10⁶?OBs/ml. Results showed that the mortalities for LdMNPV-T1, -T2 and -T3 ranged between 13–53%, 47–100% and 46–93%, respectively. The LC₅₀ and LC₉₅ values of LdMNPV-T2 were not significantly different from the respective corresponding values of the other two isolates. However, isolate LdMNPV-T2 killed larvae with a LC₅₀ value that was lower than the other two isolates. Our results suggested there are promising LdMNPV isolates in Turkey that can be used for microbial control of L. d. dispar larvae.     

Influence of mineral fertilization on food quality of oak leaves and utilization efficiency of food components by the gypsy moth

Host plant quality is a key determinant of the performance of larvae of herbivorous insects. The effects of nitrogen and dolomite fertilization on the quality of pedunculate oak, Quercus robur L. (Fagaceae) foliage, as a food for gypsy moth, Lymantria dispar L. (Lepidoptera: Lymantriidae) larvae were evaluated. The seedlings were divided into five fertilization treatments (nonfertilized control, commercial nutrient solution, commercial nutrient solution + (NH₄)₂SO₄, commercial nutrient solution + KNO₃, and commercial nutrient solution + dolomite). The experiment was performed in Petri dishes, in each of which a fresh leaf from one treatment and one larva were placed. Insect performance assays, survival, development, growth, and food utilization were evaluated for each fertilization treatment. Leaf samples were assayed for nitrogen and other main nutrients, soluble carbohydrates, and phenolic compounds. The fertilizer treatment with added ammonium improved gypsy moth performance, and the amount of food eaten was the lowest in this treatment. Utilization of elements from the food depended on the element and on the fertilization treatment. The insect bodies retained 50–64% of the nitrogen and 55–79% of the phosphorus. The results show that the efficiency of conversion of ingested food (ECI) and the efficiency of conversion of digested food (ECD) differ among the fertilization treatments, but it is not possible to define a general trend. Our results suggest that fertilization (especially ammonium) of host plants can increase herbivore performance, decrease the amount of food needed, and increase its utilization efficiency.     

Behaviour of late-instar gypsy moth larvae in high and low density populations

ABSTRACT. 1. Using scaffolding and night-vision equipment, we observed fifth and sixth instars of the gypsy moth, Lymantria dispar (L.), on Quercus velutina Lam. in the field.
2. In low-density populations, larvae fed at night and spent the day resting in sheltered sites away from the canopy. In high-density populations, larvae remained in the canopy throughout the day and night, and the amount of feeding during daylight hours increased with population density.
3. Larvae at all population densities used a similar sequence of behaviours and sampled a number of leaves when selecting feeding sites, but larvae in high-density populations switched feeding sites more frequently and fed continuously for shorter periods.
4. Larvae seldom interfered with each other's feeding in any of the populations.
5. When fifth instars were collected from the field and held for 24 h in an electronic feeding monitor, they maintained feeding rhythms that were characteristic of their source populations. Larvae spent more time crawling and less time feeding when offered foliage from high-density rather than low-density populations.     

Parasitoid complex of the Asian gypsy moth (Lymantria dispar) (Lepidoptera: Lymantriidae) in Primorye Territory,Russian Far East

A study of the moth parasitoid complex attacking gypsy moth, Lymantria dispar (L.) was carried out in Primorye territory, Russia Far East. Season-long collections at three sites in the Vladivostok area and collections at seven sites of central region of Primorye revealed the 18 primary parasites: one egg parasitoid, 11 larval parasitoids, one larval-pupal parasitoid, one parasitic nematode, one ectoparasitoid, and three diseases including NPV (nuclear polyhedrosis virus) and Entomophaga maimaiga. Phobocampe species (Ichneumonidae) dominated the parasitoid complex, parasitizing 5.5% of the larvae in the Vladivostok area and 9.3% in central Primorye, rates which are much higher than those detected from other Asiatic regions of Russia and Northeastern Asia. The insect parasitoid complex was found to be somewhat depauperate. The 11.8% average total parasitism in eastern Russian is similar to the 12% recorded in the US. Both regions have large gypsy moth outbreaks, but other factors including diseases have compensated for the rather low mortality exerted by the parasitoid complex in the Russian Far East.     

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.     

Oxaspiropentane derivatives as effective sex pheromone analogues in the gypsy moth: electrophysiological and behavioral evidence

A number of oxaspiropentane derivatives (OXPs) were tested as potential (+)-disparlure analogues, with the aim of identifying any possible interaction of these compounds, be it additive, synergetic, or inhibitory, with the pheromone response in the male gypsy moth Lymantria dispar. As assessed by male electroantennograms, 2 OXPs, 2-decyl-1-oxaspiro[2.2]pentane (OXP-01) and 4-(1-oxaspiro[2.2]pent-2-yl)butan-1-ol (OXP-04), were found to be effective. OXP-01 had no stimulatory effect but strongly decreased the response to (+)-disparlure in a blend in a 1:1 ratio. By contrast, OXP-04 proved to be more stimulating than (+)-disparlure and also had an additive effect in the blend. Single-cell recordings from the sensilla trichoidea showed the activity of 2 cells, one of which responded to (+)-disparlure. OXP-01 reduced the stimulating effectiveness of pheromone by silencing the pheromone-responding unit when the 2 compounds were presented in blend, whereas OXP-04 mimicked the pheromone response, evidenced by exciting the pheromone-responding neuron when tested alone. Behavioral observations are in agreement with electrophysiological results.     

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.     

Intraspecific acoustic communication and mechanical sensitivity of the tympanal ear of the gypsy moth Lymantria dispar

Tympanal ears of female gypsy moths Lymantria dispar dispar (L.) (Lepidoptera: Erebidae: Lymantriinae) are reportedly more sensitive than ears of conspecific males to sounds below 20 kHz. The hypothesis is tested that this differential sensitivity is a result of sex‐specific functional roles of sound during sexual communication, with males sending and females receiving acoustic signals. Analyses of sounds produced by flying males reveal a 33‐Hz wing beat frequency and 14‐kHz associated clicks, which remain unchanged in the presence of female sex pheromone. Females exposed to playback sounds of flying conspecific males respond with wing raising, fluttering and walking, generating distinctive visual signals that may be utilized by mate‐seeking males at close range. By contrast, females exposed to playback sounds of flying heterospecific males (Lymantria fumida Butler) do not exhibit the above behavioural responses. Laser Doppler vibrometry reveals that female tympana are particularly sensitive to frequencies in the range produced by flying conspecific males, including the 33‐Hz wing beat frequency, as well as the 7‐kHz fundamental frequency and 14‐kHz dominant frequency of associated clicks. These results support the hypothesis that the female L. dispar ear is tuned to sounds of flying conspecific males. Based on previous findings and the data of the present study, sexual communication in L. dispar appears to proceed as: (i) females emitting sex pheromone that attracts males; (ii) males flying toward calling females; and (iii) sound signals from flying males at close range inducing movement in females, which, in turn, provides visual signals that could orient males toward females.     

Larval densities and a life-table for the gypsy moth, Lymantria dispar, estimated using the head-capsule collection method

ABSTRACT. 1. In order to estimate the absolute larval density in each stage of a larval population of the gypsy moth, Lymantria dispar L., in a deciduous forest in northern Japan, the head-capsule collection method was used. An estimate by this method was compared with estimates based on two kinds of frass collection methods.
2. Twenty-one traps made of cloth were put in a study plot. Each trap was of 24.5 cm diameter. Larval head-capsules falling into the traps were collected and sorted by hand. On the first sampling occasion, the population was also estimated using the frass-collection method.
3. Larval numbers estimated by the head-capsule collection method were almost identical to estimates by the two frass methods. Larval numbers entering the four larval instar were successfully estimated by the head-capsule collection method, and an age-specific life table was established using the resultant estimates.     

Caterpillar guts and ammonia volatilization: retention of nitrogen by gypsy moth larvae consuming oak foliage

The gypsy moth (Lymantria dispar L.), a major defoliator of hardwood forests in the eastern U.S., has a highly alkaline midgut pH. We hypothesized that the high pH would cause high rates of ammonia (NH₃) volatilization as larvae consumed foliage, leading to potentially large losses of N from the ecosystem to the atmosphere during gypsy moth outbreaks. We measured NH₃ emission during the consumption of oak foliage by larvae in the laboratory. Surprisingly, we found very low amounts of NH₃ release of about 0.1% of the N consumed in foliage. We speculate that digestive mechanisms may limit NH₃ production in the midgut, and that the acidic environment of the hindgut traps most of the small amount of NH₃ that is produced, effectively preventing a potentially very large N loss from both larvae and ecosystem. The estimated rate of NH₃ emission from a defoliated forest is small compared to other inputs and outputs of N from the ecosystem, but could potentially enhance the neutralization of atmospheric acidity during the defoliation period. Received: 12 May 1998 / Accepted: 28 July 1998     

Changes in protein patterns in sperm and vas deferens during the daily rhythm of sperm release in the gypsy moth

In the gypsy moth, Lymantria dispar, the release of sperm bundles from the testis into the upper vas deferens (UVD) is precisely timed within each 24 h period by a circadian mechanism located in the reproductive system. In males kept under light:dark cycles of 16:8, release of sperm bundles is limited to the 3 h period that starts before lights off. Sperm released from the testis remains in the UVD for about 12 h and then moves into the seminal vesicles, so that the UVD stays empty until the next cycle of sperm release begins. The rhythm of release appears to play a role in the terminal stages of sperm maturation and is essential for the fertility of males. Sperm bundles undergo substantial morphological changes during the release from the testis and while they are retained in the UVD. In this study, using gel electrophoresis, we compared protein patterns in sperm and in the UVD during the daily cycle of sperm release and maturation. Several protein bands evident in the sperm bundles contained in the testis were missing from the sperm bundles that had passed from the testis into the UVD. Furthermore, a number of new proteins appeared in the sperm bundles as they remained in the UVD. Some of these proteins appeared to be secreted from the UVD epithelium into the UVD lumen before being incorporated into sperm bundles. Correlations between changes in protein patterns and ultrastructural changes in sperm during the cycle of sperm release and maturation are discussed. © 1994 Wiley-Liss, Inc.