Characterisation of Cydia pomonella granulovirus from codling moths in a laboratory colony and in orchards of British Columbia*

Sequence analyses indicate that a granulovirus isolated from a laboratory colony of codling moth in British Columbia, Canada is identical or very similar to the Mexican isolate of Cydia pomonella granulovirus (CpGV-M). In addition to codling moth, the vims was found to cause mortality of secondary leafroller pests of North American orchards, including the fruit tree leafroller and the obliquebanded leafroller. A survey in apple growing regions of British Columbia indicates that 23% of the wild population of codling moth larvae carry CpGV.     

Potential of high‐density pheromone‐releasing microtraps for control of codling moth Cydia pomonella and obliquebanded leafroller Choristoneura rosaceana

Recent large‐cage studies with codling moth Cydia pomonella (L.) reveal that the removal of moths from an apple orchard using pheromone‐releasing traps is more effective at reducing capture in a central monitoring trap than is a mating disruption protocol without kill/capture. The present study uses open orchard 0.2‐ha plots comparing a high‐density trapping scenario with mating disruption to confirm those results. Two tortricid moth pests of tree fruit are studied: codling moth and obliquebanded leafroller Choristoneura rosaceana (Harris). Codling moth treatments include Isomate CM FLEX (ShinEtsu Ltd, Japan), nonsticky traps baited with Trécé CM lures (Trécé, Inc., Adair, Oklahoma), and sticky traps baited with Trécé CM lures, all at equal application rates of 500 dispensers ha^?1, as well as a no pheromone control. These microtraps are of a novel design, small and easy to apply, and potentially inexpensive to produce. Mating disruption using Isomate CM FLEX and nonsticky traps reduces codling moth capture in standard monitoring traps by 58% and 71%, respectively. The attract‐and‐remove treatment with sticky traps reduces capture by 92%. Obliquebanded leafroller treatments include Isomate OBLR/PLR Plus and Pherocon IIB microtraps baited with Trécé OBLR lures, both applied at 500 dispensers ha^?1, as well as a no pheromone control. Mating disruption reduces capture in monitoring traps by 69%. The attract‐and‐remove treatment reduces capture by 85%. Both studies suggest that an attract‐and‐remove approach has the potential to provide superior control of moth populations compared with that achieved by mating disruption operating by competitive attraction.     

Sprayable microencapsulated sex pheromone formulations for mating disruption of four tortricid species: effects of application height, rate, frequency, and sticker adjuvant

Several application parameters of microencapsulated (MEC) sex pheromone formulations were manipulated to determine their impact on efficacy of disruption for codling moth, Cydia pomonella (L.); oriental fruit moth, Grapholita molesta (Busck); obliquebanded leafroller, Choristoneura rosaceana (Harris); and redbanded leafroller, Argyrotaenia velutinana (Walker). Depending on the experiment, the formulations evaluated were those formerly manufactured by 3M Canada (London, ON, Canada) or those that are currently available from Suterra LLC (Bend, OR). The efficacy of MEC formulations applied by air-blast sprayer evenly throughout the entire canopy of 2-3-m-tall apple (Malus spp.) trees was equivalent to treatments in which targeted applications of MECs were made to the lower or upper 1.5 m of the canopy (at equivalent overall rates) for oriental fruit moth and both leafroller species. The realized distribution of deposited microcapsules within the tree canopy corresponded well with the intended heights of application within the canopy. The additional coapplication of the pine resin sticker Nu-Film 17 increased efficacy but not longevity of MEC formulations for oriental fruit moth; this adjuvant had no added effects for codling moth or leafroller formulations. Increasing the rate of active ingredient (AI) per hectare by 20-30-fold (range 2.5-75.0 g/ha) did not improve the disruption efficacy of MECs for codling moth or either leafroller species when both low and high rates were applied at equivalent frequencies per season. A low-rate, high-frequency (nine applications per season) application protocol was compared with a standard protocol in which two to three applications were made per season, once before each moth generation for each species. The low-rate, high-frequency protocol resulted in equivalent or better disruption efficacy for each moth species, despite using two-fold less total AI per hectare per season with the former treatment. The low-rate, frequent-application protocol should make the use of MEC formulations of synthetic pheromone more economical and perhaps more effective.     

Effect of Delayed Mating on Oviposition Pattern, Fecundity and Fertility in Codling Moth, Cydia pomonella (L.) (Lepidoptera: Tortricidae)

Female codling moth ( Cydia pomonella ) were mated either on the day of emergence or 2 or 4 d after emergence. Oviposition was checked daily and fertility (% egg hatch) of eggs laid 1–4 d and 5 + d after mating was determined from a sub-sample of females in each treatment. From these data, net reproduction rate (R₀) was calculated. the effect of delayed mating of males was also determined. Mean fecundity, fertility and net reproduction rate of females mated 4 d after emergence were significantly lower than for females mated either on the day of emergence (day 0) or 2 d after emergence. There were no significant differences between the three treatments in terms of the proportion of females that mated or female longevity, although there was a trend towards increased longevity with increasing delays in mating. Mean longevity of unmated females was significantly greater than that of mated females. Oviposition pattern was similar in females regardless of when they were mated and in all cases maximum oviposition took place 2 d after mating. There was no effect on the fecundity of 2 d old females or on the fertility of their eggs when they were paired with either 2 or 6 d old males. the significance of these findings in relation to control of codling moth by mating disruption is discussed.     

Effect of mating delay on the reproductive performance of Cnephasia jactatana (Lepidoptera: Tortricidae)

The leafroller species, Cnephasia jactatana Walker, is an important pest of kiwifruit in New Zealand. The effect of mating delay on its reproductive performance was investigated in the laboratory to provide information for the development of pheromonal pest control measures such as mating disruption. Reproductive performance of both sexes is adversely affected by mating delay. Females are more severely affected by mating delay than males in terms of reproductive potential. Delaying mating in females for 3 and 4 d reduced the reproductive potential by 29 and 88%, respectively. It is suggested that mating disruption may have potential for the control of this species.     

Potential for outbreaks of leafrollers (Lepidoptera: Tortricidae) in California apple orchards using mating disruption for codling moth suppression

Two species of leafrollers, Argyrotaenia citrana (Fernald) and Pandemis pyrusana Kearfott, represent serious obstacles to the implementation of mating disruption for control of codling moth in coastal California apple orchards. Larval and adult densities of A. citrana and P. pyrusana and subsequent fruit damage were compared under different codling moth control treatments. Leafroller larval counts and levels of fruit damage were significantly higher in most plots that were untreated or treated only with codling moth pheromone. Leafroller fruit damage levels in these plots were commonly between 10 and 15% at harvest. As summer larval counts were good predictors of fruit damage levels, larval sampling could be a useful tool for predicting leafroller outbreaks. Use of pheromone trapping for A. citrana to detect localized outbreaks within an orchard was not useful and failed to correlate with larval numbers, whereas adult monitoring for P. pyrusana appears more promising. Efforts to implement a codling moth mating-disruption program in California must include changes in strategies for monitoring and controlling leafroller species.     

Modeling the interaction of physiological time, seasonal weather patterns, and delayed mating on population dynamics of codling moth, Cydia pomonella (L.) (Lepidoptera: Tortricidae)

The effect of delaying female mating on population growth in codling moth (Cydia pomonella (L.)) was found to act on a physiological time (degree-day) basis and was predictable using a simple quadratic equation. When combined with previous work on degree-day based mortality, we were able to evaluate how the magnitude of population reduction and survival varied between sites, years, and generations at locations in California, Michigan, Pennsylvania and Washington states. In general, reductions in population growth associated with females mating 1–3 days after emergence were greater in warmer areas and during warmer times of the year. In any given year and location, the temperature profiles during peak flight were crucial in determining the population reductions, but over an 11-year period, the average seasonal temperature profile was more important. During the overwintering generation, conditions were relatively mild in all locations and only minor differences were observed in population growth rates between locations. Populations experiencing 1–3 days delay in female mating were reduced 8, 19 and 32 % compared to populations experiencing no delay, respectively. During the first summer generation, population reductions doubled compared to those seen in the overwintering generation. During the second summer flight, reductions in population growth rate at the three cooler locations decreased, while they increased in the warmer California location. Overall, the results show delayed mating can help understand how population growth is related to environmental conditions experienced naturally by insect populations and will help guide studies of the mechanisms of mating disruption, a technique used for pest suppression in agricultural and forest systems.     

Effect of anti‐hail nets on Cydia pomonella behavior in apple orchards

Over the last 10 years, the use of nets to protect pome fruit from hailstorms has increased. In this study, we investigated the effect of these nets on the behavior of the codling moth, Cydia pomonella (L.) (Lepidoptera: Tortricidae), a major pest of apple, Malus domestica Borkh. (Rosaceae). Experiments were carried out in net‐covered and uncovered apple orchards treated with conventional insecticides. The number of codling moth males caught in pheromone‐ and virgin female‐baited traps was significantly reduced in net‐covered compared to uncovered plots. In addition, inhibition of mating by the net was demonstrated by significant reductions in mating of tethered virgin females. Fruit injury was consistently lower in net‐covered plots vs. uncovered plots. Because of the reductions in male trap catch, the reduced female mating frequency, and the lower level of damage, we conclude that flat anti‐hail nets have a disruptive effect on the reproductive behavior of the codling moth.     

Comparative activity of baculoviruses against the codling moth Cydia pomonella and three other tortricid pests of tree fruit

The granulovirus of Cydia pomonella (L.) (CpGV) offers potential for selective control of codling moth. Two major limitations of CpGV are its narrow host range and lack of persistence in the orchard agroecosystem. The nucleopolyhedroviruses of the alfalfa looper Autographa californica (Speyer) (AcMNPV) and those of the celery looper Anagrapha falcifera (Kirby) (AfMNPV) have broad host ranges. Comparative assays of CpGV, AcMNPV, and AfMNPV against codling moth neonate larvae revealed a 54-93-fold greater susceptibility of codling moth to the granulovirus than to the two nucleopolyhedroviruses based on the LC(50) values for each virus. The LC(50)s for CpGV, AfMNPV, and AcMNPV were 32.7 capsules/mm(2), 1.77 x 10(3) occlusion bodies (OBs)/mm(2), and 3.05 x 10(3)OBs/mm(2), respectively. The LT(50) determined for AfMNPV using an approximate LC(95) of the virus against neonate larvae was 3.6 days. Histological examination of tissues in moribund codling moth larvae that had been treated with AfMNPV revealed the presence of nonoccluded and unenveloped virus rods in midgut tissue. Neither OBs nor signs of infection were detected in other tissues. The activity of AfMNPV was also evaluated in three other tortricid apple pests (obliquebanded leafroller, Choristoneura rosaceana (Harris); Pandemis leafroller, Pandemis pyrusana Kearfott; and the oriental fruit moth, Grapholitha molesta (Busck)). Codling and Oriental fruit moths were significantly more susceptible to AfMNPV than were the two leafroller species.     

Demographic analysis of delayed mating in mating disruption: a case study with Cryptophelbia illepida (Lepidoptera: Tortricidae)

Laboratory-derived life tables were used to determine the effect of delaying mating of adult female koa seedworm, Cryptophlebia illepida (Butler) (Lepidoptera: Tortricidae), 4 and 6 d on population growth rates. Leslie matrices were developed from the life tables and used to project the effects for approximately four generations. Delay of mating caused a decrease in population growth rate and also resulted in asynchronous population cycling between control (1-d delay) and the delayed treatments. By the fourth generation, the control population began to increase 10 and 14 d before the 4- and 6-d delay treatments, respectively. Increasing the mortality of females during the first 7 d of adult life resulted in a greater reduction of the populations where mating was delayed than in the control populations. This result suggests that even at relatively low levels of natural enemy mortality, there is a synergistic effect when mating is delayed. The implications of these effects on mating disruption management programs are discussed.     

Vertical distribution of codling moth adults in pheromone-treated and untreated plots

The vertical distribution of codling moth,Cydia pomonella (L.) within pheromone-treated and untreated apple and pear orchard canopies was determined using tethered virgin females, unbaited sticky traps, and blacklight observation of released moths. Mating of virgin females tethered at various heights in untreated orchard canopies increased with placement height from 1–4 m. Application of pheromone dispensers for mating disruption at 2 and 4 m above the ground greatly decreased mating. Greatest capture of males and females on unbaited sticky traps occurred at mid- and upper-canopy heights. Total capture of males and females in pheromone-treated plots was not statistically different than in untreated plots. The percentage of mated females captured on sticky traps did not vary with trap height or pheromone treatment. Released moths marked with flourescent powder and observed at dark with a blacklight indicated that moths are primarily distributed high in the canopy. However, males shifted to a position lower in the canopy when pheromone dispensers were placed 2.1 m above the ground. Results suggest that pheromone dispensers be placed in the upper canopy for optimal disruption of codling moth mating.     

Effect of specific and generic sex attractant blends on pheromone trap captures of four leafroller species in mid-Atlantic apple orchards

The responses of tufted apple bud moth, Platynota idaeusalis (Walker), the leafroller P. flavedana Clemens, redbanded leafroller, Argyrotaenia velutinana (Walker), and obliquebanded leafroller, Choristoneura rosaceana (Harris), to the pheromone blends of each, as well as to 3 putative generic blends (two- and three-component blends containing pheromone elements of each of the 4 species) were evaluated in small orchard plots. P. idaeusalis and P. flavedana, and A. velutinana and C. rosaceana comprise 2 pairs of species, each pair with broad overlap in pheromone blend, and quite different from one another. Each generic blend suppressed trap captures of all 4 species. The blends for P. idaeusalis and P. flavedana each reduced captures for these species. Furthermore, the blend for P. idaeusalis also suppressed captures of A. velutinana. The P. flavedana blend did not reduce captures of A. velutinana; in fact, at times captures increased. This study determines relative abilities of several sex attractant blends to reduce captures of 4 leafroller species in pheromone traps, presumably reflecting the ability of a blend to reduce orientation of males to females in a large block situation. This is a requisite 1st step in the development of a multispecies mating disruption blend.     

Overwintering of codling moth (Lepidoptera: Tortricidae) larvae in apple harvest bins and subsequent moth emergence.

Codling moths, Cydia pomonella (L.), have long been suspected of emerging from stacks of harvest bins in the spring and causing damage to nearby apple and pear orchards. With increased use of mating disruption for codling moth control, outside sources of infestation have become more of a concern for growers using pheromone based mating disruption systems. Studies were designed to provide information on bins as a source of codling moth and the pattern of codling moth emergence from stacks of bins. In these studies, codling moth larvae colonized wood harvest bins at a much higher frequency than harvest bins made of injection molded plastic (189 moths emerged from wood compared with five from plastic). There was no statistical difference in the number of moths infesting bins that had been filled with infested fruit compared with bins left empty at harvest. This suggests that codling moth enter the bins during the time that the bins are in the orchard before harvest. Emergence of laboratory reared adult codling moth from wood bins placed in stacks was found to be prolonged compared with field populations. Temperature differences within the bin stacks accounted for this attenuated emergence pattern. Covering bin stacks with clear plastic accelerated codling moth development in the upper levels of the stack. Codling moth emergence patterns from plastic-covered stacks more closely coincided with male flight in field populations. This information could be important in developing a technique for neutralizing codling moth-infested bins, and in understanding how infested bins may influence pest management in fruit orchards that are located near bin piles. Implications for control of codling moth in conventional orchards and in those using mating disruption as the principal component of an integrated pest management system include increased numbers of treatments directed at areas affected by infested bins.     

Pheromone pre-exposure and mating modulate codling moth (Lepidoptera: Tortricidae) response to host plant volatiles

Abstract 1 Two codling moth Cydia pomonella kairomonal attractants, ethyl (E,Z)‐2,4‐decadienoate (pear ester) and (E)‐β‐farnesene, were tested in an insecticide‐sprayed apple orchard and an orchard treated for mating disruption with synthetic pheromone (E,E)‐8,10‐dodecadienol (codlemone). Male captures with pear ester were higher in the pheromone‐treated than in the insecticide‐treated orchard, whereas captures with (E)‐β‐farnesene were not different. Subsequent wind tunnel experiments confirmed that pre‐exposure to sex pheromone codlemone increased the behavioural response of codling moth males to pear ester. This supports the idea that male attraction to the plant volatile pear ester and sex pheromone codlemone is mediated through the same sensory channels. 2 Pear ester is a bisexual codling moth attractant and even captures of female moths were significantly increased in the pheromone‐treated orchard. In the laboratory wind tunnel, pheromone pre‐exposure had no effect on female response to pear ester, but significantly more mated than unmated codling moth females flew upwind towards a pear ester source. Differences in mating status in insecticide‐treated vs. pheromone‐treated orchards may thus account for the differences in female trap captures with pear ester. 3 These findings are important with respect to monitoring of codling moth with pear ester in mating disruption orchards. They also emphasize the importance of host plant volatiles in pheromone‐mediated mating disruption, which has been neglected to date.     

Efficacy and mode of action of female‐equivalent dispensers of pheromone for mating disruption of codling moth

1 We evaluated the efficacy and mode of action of commercially available female‐equivalent dispensers of pheromone for mating disruption of codling moth Cydia pomonella (L.) (Lepidoptera Tortricidae). 2 Scentry fibres, but not Hercon flakes, were consistently as or more effective than Isomate C Plus dispensers when applied by hand at 50 dispensers per tree. 3 Individual Scentry fibres were slightly more attractive to male codling moth than Hercon flakes. 4 Efficacy of aerially applied Scentry fibres was equivalent to that of Isomate dispensers in disrupting male codling moth in 4‐ha commercial apple plots. 5 Initial deposition and retention of aerially applied fibres were inefficient with approximately 44% formulation loss at application, poor rainfastness and a gradual loss of dispensers from tree canopies after application. 6 Male codling moths were captured in traps baited with lures containing 0.1 mg of pheromone and mated with tethered virgin females that were surrounded by eight fibres placed 30 cm away or 16 fibres placed 45 cm away in untreated plots and plots treated with a background of 50 Isomate dispensers per hectare. 7 A plausible explanation for mating disruption of codling moth by female‐equivalent dispensers is competitive attraction without associated habituation and thus improving the effectiveness of these technologies will depend on maximizing the attractiveness of individual dispensers as well as the application density of dispensers per area of crop.     

Apple damage, pest phenology, and factors influencing the efficacy of tebufenozide for control of obliquebanded leafroller (Lepidoptera: Tortricidae).

Fruit damage by obliquebanded leafroller, Choristoneura rosaceana (Harris), was examined on four different apple cultivars during 1997-1999 in heavily infested orchards in New York State. Inital fruit damage occurred 354+/-26 degree-days base 6 degrees C (DD6) after the first moth catch from the overwintering generation and continued to increase until after the typical spray season (approximately 1,300 DD6). In addition to apple damage by late instars, fruit damage occurred by early instars of the summer and overwintering generations. The insect growth regulator tebufenozide was used as a model insecticide to determine how a slow-acting insecticide and its relative toxicity to early (neonate) and late (fourth and fifth) instars may influence the efficacy of sprays for the control of the obliquebanded leafroller. Because tebufenozide is a slow-acting compound, bioassays were conducted to determine what percentage of the total mortality to neonates occurs at each 24 h interval until 10 d. Based on a polynomial regression, half of the total mortality to larvae at the LC25, LC50, LL90 and LC99 occurred at 7.2, 5.0, 4.1, and 3.0 d, respectively. Late instars were three times more tolerant to tebufenozide than neonates.     

Evaluation of border sprays for managing the codling moth (Tortricidae: Lepidoptera) and the apple maggot (Tephritidae: Diptera) in Ontario apple orchards

The efficacy of two insecticide control programs for managing the codling moth, Cydia pomonella (L.), and the apple maggot, Rhagoletis pomonella (Walsh), were compared in the Georgian Bay, London, Niagara, and Quinte apple production areas of Ontario during 1995, 1996, and 1997. In the border spray program, an initial cover spray of organophosphorus insecticide was applied to eradicate codling moths that may have colonized a test plot during the previous growing season. Subsequent sprays were applied only to a four-tree-wide zone (approximately 20 wide) around the perimeter of the plot to control immigrating codling moths or apple maggots. In the cover spray program, all sprays of organophosphorus insecticide were applied to the entire plot. Apple maggot injury was significantly greater in border spray program plots than in cover spray program plots only during 1995 in the London production area. There was no significant difference in codling moth injury between border spray and cover spray plots in the four production areas during the three-year study. The elimination of cover sprays from border spray plots during July and August may have left the apple crop more susceptible to damage by second generation larvae of the obliquebanded leafroller, Choristoneura rosaceana (Harris), in the London production area during 1995. There was a trend of increasing codling moth injury from 1995 to 1997 in two border spray plots, and apple maggot injury was detected in these plots during the third year of the study.     

Development of 2‐phenylethanol plus acetic acid lures to monitor obliquebanded leafroller (Lepidoptera: Tortricidae) under mating disruption

We evaluated the effectiveness of 2‐phenylethanol (PET) in combination with acetic acid (AA) as a binary lure for monitoring male and female obliquebanded leafroller, Choristoneura rosaceana (Harris). Studies were conducted in apple, Malus domestica Borkhausen, orchards treated with or without sex pheromone dispensers for mating disruption (MD). Open polypropylene vials, closed membrane cups, and rubber septa loaded with AA and/or PET in varying amounts were first evaluated in a series of trapping experiments. Membrane cups loaded with 800 mg of PET were as effective as 10‐mg septa, but longer lasting, and were comparable to the open vials. A membrane cup AA lure was effective in tests, but further work is needed to increase its release rate and extend its activity. Catches of codling moth, Cydia pomonella (L.), and C. rosaceana were unaffected by combining PET with (E,E)‐8,10‐dodecadien‐1‐ol, the sex pheromone of codling moth, pear ester, (E,Z)‐2,4‐ethyl‐decadienoate and AA lures. Adding (E)‐4,8‐dimethyl‐1,3,7‐nonatriene to this blend to enhance codling moth catch significantly reduced catches of C. rosaceana. PET + AA was a more attractive binary lure than AA plus phenylacetonitrile (PAN) for C. rosaceana. The addition of PET or PAN to traps already baited with the sex pheromone of C. rosaceana significantly reduced male catches. Traps baited with PET + AA placed in blocks not treated with MD caught significantly fewer C. rosaceana than traps baited with sex pheromone. In comparison, sex pheromone‐baited traps in MD blocks caught ≤1 male moth per season which was significantly lower than total moth (>10) or female moth (≥3) catch in these blocks with PET + AA. A high proportion (>70%) of trapped females were mated in both untreated and MD‐treated orchards. Further refinement of this binary, bisexual lure using membrane cup technology may allow the establishment of action thresholds and improve management timings for C. rosaceana.     

Life history changes in Trogoderma variabile and T. inclusum due to mating delay with implications for mating disruption as a management tactic

Controlling postharvest pest species is a costly process with insecticide resistance and species‐specific control requiring multiple tactics. Mating disruption (MD) can be used to both decrease a female's access to males and delay timing of mating and decreases overall mating success in a population and population growth rate. Development of new commercially available MD products requires an understanding of life history parameters associated with mating delay. These can provide information for targeting proportions of reproducing individuals using MD. After delaying mating for females of two closely related beetle species, Trogoderma variabile and T. inclusum, we surveyed survivorship, number of eggs laid, and number of progeny emerged. With increases in mating age, total number of eggs laid and total number of progeny emerged significantly declined over time. T. inclusum typically had greater numbers of eggs laid and progeny emerged compared to T. variabile as female age at mating increased, suggesting that T. inclusum may be more resistant to long‐term delays in mating. Life span showed an increase as mating age increased but life span significantly decreased almost immediately following mating. Simulations depicting multiple distributions of mating within a population suggest that in a closed population, high levels of mating delay significantly reduced reproductive growth rates. Although reproductive growth rates were decreased with increased mating age, they are still large enough to maintain populations. This study highlights the differences in life history between two closely related species, suggesting that T. inclusum outperforms T. variabile over the course of a life span, but T. variabile has better reproductive capabilities early in life. MD may also be a viable component of a pest management system for these two species as it significantly decreased overall reproductive output and population growth.     

Reproductive performance and longevity of female European corn borer,Ostrinia nubilalis: effects of multiple mating,delay in mating,and adult feeding

In a recent study on the pheromone-mating disruption of the European corn borer, Ostrinia nubilalis (Lepidoptera: Crambidae), we recorded a significant reduction in mating frequency, as well as a marked delay in mating in feral females captured in disruptant-treated fields. In order to be able to accurately interpret the results in terms of effective population control, the current study was undertaken on the effects of multiple matings and a delay in mating on reproductive performance. Female O. nubilalis that mated at least twice had significantly higher fecundity and fertility, compared with once-mated females. In addition, multiple-mated females deposited a significantly larger portion of their egg complement, relative to single-mated or unmated females. Females that experienced a 3-day delay in mating showed a significant reduction in fecundity compared with females that mated soon after emergence. A 1-week delay in mating resulted in a further reduction in fecundity and a near zero fertility. The effect of sugar feeding on reproduction was not significant. In general, unmated females lived longer than mated females, and sugar-fed mated females had a higher longevity than water-fed mated females.