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.     

Comparison of delayed female mating on reproductive biology of codling moth and obliquebanded leafroller

Delay of mating was examined as a possible mechanism for population decreases associated with mating disruption for codling moth, Cydia pomonella L., and obliquebanded leafroller, Choristoneura rosaceana (Harris) (Lepidoptera: Tortricidae). We examined the effect of delaying female mating 0, 2, 4, or 6 d while holding male age constant on life table parameters of both species. We found that increasing delays in mating were accompanied by two responses: (1) an increase in the percentage of sterile pairs and (2) a reduction in net reproductive rate and population growth unrelated to sterility. On a percentage basis, obliquebanded leafroller population growth was more strongly affected than codling moth. However, the net fertility rate of obliquebanded leafroller was nearly eight-fold higher than that of codling moth, so that obliquebanded leafroller females that experienced a 4-d delay in mating had nearly the same reproductive rate as codling moth females that experienced no delay. Leslie matrix simulations using life tables with field-based adult longevity estimates showed that codling moth females experiencing >2-d delay in mating resulted in decreases in population density or extinction within two generations. In contrast, obliquebanded leafroller females delayed <6 d showed rapid population growth that decreased as female age at mating increased; only the 6-d delay treatment resulted in decreased population levels. Our results indicate that obliquebanded leafroller females must on average experience a much longer delay in mating to significantly reduce population growth compared with codling moth females, suggesting that delay of mating likely plays a greater role in codling moth mating disruption than for obliquebanded leafroller.     

Instar-specific phenology of Pandemis pyrusana and Choristoneura rosaceana (Lepidoptera: Tortricidae) in Washington apple orchards

Head capsule width was used to determine the instar specific phenology of the leafroller Pandemis pyrusana Kearfott and the obliquebanded leafroller, Choristoneura rosaceana (Harris) (Lepidoptera: Tortricidae), attacking apple in Washington state during 2001-2003. In total, 7012 P. pyrusana and 6122 obliquebanded leafroller larvae were measured from apple orchards from mid-March to mid-September. Degree-day accumulations from each site were paired with the head capsule data to determine the periods during which different instars were present in the field. The implications of this work for pest management and biological control of leafrollers is discussed.     

Delayed mating and reproduction in the autumn gum moth Mnesampela privata

• 1 The success of mating disruption using synthetic sex pheromones depends not only on preventing mating, but also on delaying mating in the target insect. Using the geometrid pest of Eucalyptus plantations, Mnesampela privata (Guenée), we determined the effect of delaying mating when imposed on males only, females only or on both sexes simultaneously, for 1, 3, 5 and 7 days.
• 2 Delayed mating had a significant negative impact on reproduction, with a 0.89‐fold decrease in the likelihood of mating and a 0.67‐fold decrease in the likelihood of that mating resulting in fertile eggs for every day that mating is delayed. A mating delay of 7 days reduced the mean number of viable eggs laid to 4–13% of that laid by moths paired immediately after emergence.
• 3 Male only imposed mating delays had a significantly lower effect on reducing the likelihood of pairs mating than when both sexes were delayed. A delay imposed on one sex only or on both sexes simultaneously, however, had a similar negative impact on the proportion of fertile matings as well as on the total number of fertile eggs laid.
• 4 Longevity of mated female and male M. privata was significantly different between mating delay treatments, with a significant decline in female longevity when they mated with older males.
• 5 The underlying mechanisms causing a decline in female reproductive output when a mating delay was imposed on males versus females are discussed in relation to the reproductive biology of M. privata and the potential of using mating disruption strategies to control populations in Eucalyptus plantations.

     

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.     

Discovery of a granulovirus of Pandemis pyrusana (Lepidoptera: Tortricidae), a leafroller pest of apples in Washington

Pandemis pyrusana (Kearfott) is an important leafroller pest of apples in Washington. Surveys for natural enemies discovered a pathogen infecting Pandemis leafrollers in an apple orchard in central Washington. The pathogen was propagated in the laboratory and light microscopy using an azan stain demonstrated that it infected fat body, epidermis, and tracheal matrix cells. The virus was identified morphologically as a granulovirus using electron microscopy and designated PpGV. Rates of infection were determined for each generation in an apple orchard for three years. Infection rates were variable and ranged from 2.6 to 67% of individuals collected from each generation.     

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.     

Sex ratio at mating does not modulate age fitness effects in Drosophila melanogaster

Understanding the effects of male and female age on reproductive success is vital to explain the evolution of life history traits and sex‐specific aging. A general prediction is that pre‐/postmeiotic aging processes will lead to a decline in the pre‐ and postcopulatory abilities of both males and females. However, in as much the sexes have different strategies to optimize their fitness, the decline of reproductive success late in life can be modulated by social context, such as sex ratio, in a sex‐specific manner. In this study, we used Drosophila melanogaster to investigate whether sex ratio at mating modulates age effects on male and female reproductive success. As expected, male and female age caused a decrease in reproductive success across male‐biased and female‐biased social contexts but, contrary to previous findings, social context did not modulate age‐related fitness decline in either of the two sexes. We discuss these results in the light of how sex ratio might modulate pre‐/postcopulatory abilities and the opportunity for inter‐ and intrasexual competition in D. melanogaster, and generally suggest that social context effects on these processes are likely to be species specific.     

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.     

Influence of male mating history on female reproductive success among monandrous Naryciinae (Lepidoptera: Psychidae)

1. Multiple male copulations can have detrimental effects on female fitness due to sperm limitation. 2. Monandrous Naryciinae females are immobile while the males are short‐lived and do not feed. Multiple male mating is therefore expected to lead to sperm limitation in females. Sperm limitation and male limitation are hypothesised as causes of the repeated evolution of parthenogenetic reproduction in the Psychidae. 3. In this study, the effects of multiple male mating on female reproduction are investigated in several species of Naryciinae by allowing males multiple copulations. The results for two species, Siederia listerella and Dahlica lichenella, are compared. The sex ratios of 53 natural populations are examined for indications of male limitation. 4. Previous copulations by the male increased the female's risk of remaining unfertilised. However, contrary to expectations, those unfertilised females were capable of successful re‐mating. 5. In S. listerella, the number of previous copulations of males negatively influenced female fitness. Females produced 30% fewer offspring if they mated with a previously mated male. In D. lichenella, the older the male and the lower its number of total lifetime copulations, the higher the female's reproductive success. 6. Only a fraction of the investigated populations had a female‐skewed sex ratio, but differences in development time between males and females could lead to reproductive asynchrony. 7. In conclusion, male mating history did not lead to strong sperm limitation in Naryciinae as had been suggested by their life history.     

Sex peptide causes mating costs in female Drosophila melanogaster

Conflicts between females and males over reproductive decisions are common . In Drosophila, as in many other organisms, there is often a conflict over how often to mate. The mating frequency that maximizes male reproductive success is higher than that which maximizes female reproductive success . In addition, frequent mating reduces female lifespan and reproductive success , a cost that is mediated by male ejaculate accessory gland proteins (Acps) . We demonstrate here that a single Acp, the sex peptide (SP or Acp70A), which decreases female receptivity and stimulates egg production in the first matings of virgin females , is a major contributor to Acp-mediated mating costs in females. Females continuously exposed to SP-deficient males (which produce no detectable SP ) had significantly higher fitness and higher lifetime reproductive success than control females. Hence, rather than benefiting both sexes, receipt of SP decreases female fitness, making SP the first identified gene that is likely to play a central role in sexual conflict.     

豆野螟延迟交尾和多次交尾对生殖的影响

豆野螟Maruca vitrata(Fabricius)是一种严重的泛热带豆类蔬菜害虫。本文研究了豆野螟延迟交尾和多次交尾对其生殖的影响。延迟交尾实验结果表明:豆野螟雌雄同时延迟交尾,雌雄虫的寿命、产卵量均表现为先上升后下降的趋势,但是对卵的孵化率没有显著影响;雌虫延迟交尾,随着延迟时间的增加,雌虫的寿命、产卵量、卵的孵化率表现为下降的趋势,而雄虫的寿命延迟交尾第3天达到最大值;雄虫延迟交尾,随着延迟交尾时间的增加,雌雄虫的寿命、产卵量均表现为先上升后下降的趋势,卵的孵化率随着延迟逐渐下降。多次交尾实验结果表明,随着雄虫交尾次数的增加,成功交尾率逐渐降低,用于交尾的时间延长,雌虫和相应雄虫的寿命逐渐缩短,雌虫的产卵量下降,但对卵的孵化率影响不大;豆野螟雌虫一生只交尾一次,未见到雌虫2次交尾。     

Delayed mating reduces reproductive output of female European grapevine moth,Lobesia botrana (Lepidoptera: Tortricidae)

Virgin females of Lobesia botrana Denis & Schiffermüller were mated at ages of 1, 3, 5, 8, 12 and 16 days and the effect of mating delay on reproductive output assessed. Delayed mating did not affect female mating success but fertilization was reduced in 16-day-old females. Increased mating delays substantially affected daily oviposition pattern and resulted in a significant reduction of both fecundity and fertility, so that mean number of viable eggs laid decreased from 150 to 22 in 1- and 16-day-old mated females, respectively. Heavier females were more constrained than lighter ones by mating delays and female reproductive efficiency (no. viable eggs/female body weight) was also significantly reduced. Female longevity significantly increased and oviposition period gradually declined with mating delay. The number of viable eggs was positively correlated with both female weight and oviposition period; female longevity and female weight were also significantly correlated. However, the significance of these correlations declined with increased mating delay. Results overall indicated that mating delay drastically reduces female L. botrana reproductive output. The implications of delayed mating of females are discussed from an ecological perspective in relation to L. botrana control using mating disruption.     

A survival cost to mating in a polyandrous butterfly, Colias eurytheme

Adaptations that enhance fitness in one sex may be harmful to members of the opposite sex and lead to antagonistic coevolution between the sexes. In fruit flies, for example, selection for fertilization success has rendered the male ejaculate slightly toxic to females. Here we investigated whether mating imposes a cost upon female fitness in a polyandrous pierid butterfly ( Colias eurytheme ) by comparing life history traits between once-mated females and virgins. Mated females laid relatively more eggs early in their adult life, but suffered a reduction in longevity relative to virgins held under identical experimental conditions. The effect of mating on female survivorship was statistically independent of lifetime and early life fecundity. Moreover, lifetime fecundity co-varied positively with longevity across all females, and across females within each treatment group, hence there was no phenotypic trade-off between survival and reproduction. These results suggest that the observed longevity difference between virgin and mated females represents a true cost of mating, possibly arising from a toxic side effect of the male ejaculate. However, irrespective of this cost, virgin and mated females laid an equivalent lifetime number of eggs. Female C. eurytheme are also known to use nutrients from the male ejaculate to supplement their reproductive output, hence it is presently unclear how the observed longevity cost may have influenced the evolution of lifetime mating schedules in this polyandrous species.     

Multiple mating in the traumatically inseminating Warehouse pirate bug, Xylocoris flavipes: effects on fecundity and longevity

Optimal mating frequencies differ between sexes as a consequence of the sexual differentiation of reproductive costs per mating, where mating is normally more costly to females than males. In mating systems where sexual reproduction is costly to females, sexual conflict may cause both direct (i.e. by reducing female fecundity or causing mortality) and indirect (i.e. increased risk of mortality, reduced offspring viability) reductions in lifetime reproductive success of females, which have individual and population consequences. We investigated the direct and indirect costs of multiple mating in a traumatically inseminating (TI) predatory Warehouse pirate bug, Xylocoris flavipes (Reuter) (Hemiptera: Anthocoridae), where the male penetrates the female's abdomen during copulation. This study aimed to quantify the effects of TI on female fecundity, egg viability, the lifetime fecundity schedule, longevity and prey consumption in this cosmopolitan biocontrol agent. We found no difference in the total reproductive output between mating treatments in terms of total eggs laid or offspring viability, but there were significant differences found in daily fecundity schedules and adult longevity. In terms of lifetime reproduction, female Warehouse pirate bugs appear to be adapted to compensate for the costs of TI mating to their longevity.     

Control of reproduction and a survival cost to mating in female Bicyclus anynana butterflies

Abstract.  1. Given that sexual conflict is all pervasive, investigating potential costs to mating and the control of female reproduction are important issues.
2. Here, female reproductive output and longevity are investigated in relation to mating status (virgin vs. once-mated females) and host-plant availability in the tropical butterfly Bicyclus anynana .
3. Both factors significantly affected realised fecundity in a manner demonstrating that female reproduction is not entirely under control of the female herself, but is rather subject to additive contributions of the female, her male partner, and intrinsic physiological processes. As evident from the deposition of significant egg numbers by virgin, host-deprived females, B. anynana is effectively unable to completely inhibit oogenesis.
4. Mated females suffered a reduction in adult life span, which cannot be explained as a side-effect of variation in egg size, lifetime, or early fecundity.
5. Such detrimental effects of mating per se are indicative of the cooperation–conflict balance between sexes being shifted towards conflict in B. anynana .     

Detection of female mating status using chemical signals and cues

Males of many species choose their mate according to the female's reproductive status, and there is now increasing evidence that male fitness can depend on this discrimination. However, females will also aim to regulate their mating activity so as to maximize their own fitness. As such, both sexes may attempt to dictate the frequency and timing of female mating, reflecting the potentially different costs of female signaling to both sexes. Here, I review evidence that chemical cues and signals are used widely by males to discriminate between mated and unmated females, and explore the mechanisms by which female odour changes post‐mating. There is substantial empirical evidence that mated and unmated females differ in their chemical profile, and that this variation provides males with information on a female's mating status. Although there appears to be large variation among species regarding the mechanisms by which female odour is altered post‐mating, the transfer of male substances to females during or subsequent to copulation appear to play a major role. This transfer of substances by males may be part of their strategy to suppress reproduction by competing males, particularly in species where females mate more than once.     

桑天牛成虫交配后血淋巴和生殖系统内可溶性总糖和蛋白质含量的变化

以生理状况相似的未交配桑天牛Apriona germari Hope雌、雄成虫为供试昆虫, 观察、记录交配活动;采集血淋巴,解剖生殖系统并绘图;用蒽酮比色法和福林-酚法检测交配前、后成虫血淋巴和生殖系统内可溶性总糖和蛋白质含量变化。结果表明：交配后1 h,桑天牛雄虫血淋巴内的可溶性总糖含量增加21.38%,蛋白质含量降低22.66%;雄虫生殖系统内的可溶性总糖和蛋白质含量都明显升高;雄性附腺作为某些特异性蛋白质的合成场所其内的可溶性总糖和蛋白质含量分别降低81.76%和63.76%,雌虫血淋巴和卵巢内的可溶性总糖和蛋白质含量都升高。交配后, 雄虫发生陪伴行为最短历时为4 h, 可能是其重要的生殖策略之一。     

Age-specific mating and reproductive senescence in the seven-spotted ladybird, Coccinella septempunctata

Abstract:  Adults of the seven-spotted ladybird, Coccinella septempunctata Linnaeus (Col., Coccinellidae) were paired for mating from very young to old age (1–50 days) to record the willingness to mate, attainment of sexual maturity and onset of reproductive senescence in both the sexes. Mating commenced after 4 and 2 days of emergence of male and female, respectively, and 100% mating was achieved at the young age in both cases (10 days). Willingness to mate decreased with increase in the age from 40 to 50 days of both the sexes. Ladybird exhibited protandry. Mating duration, fecundity and per cent viability of eggs of middle-aged males (20–30 days old) and females (20 days old) were the highest. Mating duration and per cent viability of eggs were male age dependent, whereas pre-oviposition and oviposition periods were mating stimulus dependent. Oviposition period and fecundity were female age-dependent responses. Fecundity was highest when 20-day-old female and 30-day-old male were paired. Onset of reproductive senescence started at the age of 30 days in males and 20 days in females. The present study confirms the effect of ageing on male and female C. septempunctata and supports the Hansen and Price [J. Evol. Biol. (1995) vol. 8, pp. 759–778] model that females mating with young and middle-aged males yield optimal quality progeny.     

The combined effects of pre‐ and post‐copulatory processes are masking sexual conflict over mating rate in Gerris buenoi

In polygynandrous animals, post‐copulatory processes likely interfere with precopulatory sexual selection. In water striders, sexual conflict over mating rate and post‐copulatory processes are well documented, but their combined effect on reproductive success has seldom been investigated. We combine genetic parentage analyses and behavioural observations conducted in a competitive reproductive environment to investigate how pre‐ and post‐copulatory processes influence reproductive success in Gerris buenoi Kirkaldy. Precopulatory struggles had antagonistic effects on male and female reproductive success: efficiently gaining copulations was beneficial for males, whereas efficiently avoiding copulations was profitable for females. Also, high mating rates and an intermediate optimal resistance level of females supported the hypothesis of convenience polyandry. Contrary to formal predictions, high mating rates (i.e. the number of copulations) did not increase reproductive success in males or decrease reproductive success in females. Instead, the reproductive success of both sexes was higher when offspring were produced with several partners and when there were few unnecessary matings. Thus, male and female G. buenoi displayed different interests in reproduction, but post‐copulatory processes were masking the effects of copulatory mating success on reproductive success. Given the high mating rates observed, sperm competition could easily counter the effect of mating rates, perhaps in interaction with cryptic female choice and/or fecundity selection. Our study presents a complex but realistic overview of sexual selection forces at work in a model organism for the study of sexual conflict, confirming that insights are gained from investigating all episodes in the reproduction cycle of polygynandrous animals.