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Pathogen outbreaks often represent a major issue when mass rearing target insects for biological control, even when applying sterile insect techniques (SIT). Here, we aimed to improve existing screening techniques used to detect a neogregarine Farinocystis sp. that infects the West Indian sweet potato weevil Euscepes postfasciatus Fairmaire (Coleoptera: Curculionidae) mass reared under the SIT program. Our results revealed that infected late-instar larvae exhibited swelling symptoms, and that their body weight and head width significantly increased compared to uninfected larvae. The head width to body weight ratio differed between infected and uninfected larvae; yet, head width was highly correlated with body weight in both cases. Visual diagnosis based on the appearance of the late-instar larvae showed concordance of almost 90 % with the current diagnostic methods of optical microscopy and PCR. Thus, the swelling symptoms observed in late-instar larvae were directly noticeable. Therefore, visual diagnosis, based on larval appearance, represents a feasible, low-cost technique for the early detection of Farinocystis sp. infection of the West Indian sweet potato weevil.

Sterile insect technique (SIT) is widely used to eradicate target pest species. Its effectiveness depends on the ability of released sterile males to mate with wild females. In pest species with a female choice mating system, there is potential for female preference to evolve in the wild population in view of the resulting zero-fitness in females that mate with sterile males. The evolution of female preference against sterile males can, therefore, reduce the efficiency of the SIT as the likelihood decreases that sterile males will mate with wild females. We examined this experimentally by using the solanaceous fruit fly Bactrocera latifrons, a serious pest of solanaceous crops, by allowing female preference in mass-reared fruit flies to select between fertile and sterile males over 12 generations. However, this did not generate a rapid evolutionary response. Also the remating rate of females did not increase, even when the first mating partner was a sterile male. We discuss the implication of the results ...

Selection for genetic adaptation might occur whenever an animal colony is maintained in the laboratory. The laboratory adaptation of behavior such as foraging, dispersal ability, and mating competitiveness often causes difficulties in the maintenance of biological control agents and other beneficial organisms used in procedures such as the sterile insect technique (SIT). Sweet potato weevil, Cylas formicarius (Summers) (Coleoptera: Brentidae), is an important pest in sub-tropical and tropical regions. An eradication program targeting C. formicarius using SIT was initiated in Japan with weevils being mass-reared for 95 generations to obtain sufficient sterile males. The mass-reared strain of C. formicarius exhibits weaker female resistance to male mating attempts compared with the wild strain. This could affect the success of SIT programs because mating persistence of mass-reared males might be expected to decrease in response to weak female resistance. We show that high success of sperm transfer to mass-reared females was due to weak female resistance to male mating attempts. However, the mating behavior of mass-reared males did not change. In C. formicarius, the trait of male persistence to mate was not correlated with the female resistance traits. Our results suggest that mass-rearing conditions do not have negative effects on the mating ability of the sterile males of this species, and thus that the current mass-rearing procedures are suitable for production of sterile males for the weevil eradication program.

For ensuring the effectiveness of sterile insect technique (SIT) programmes, maintaining the reproductive competitiveness and dispersal ability of mass-reared sterile males is essential. Inadvertent selection is an important genetic process that frequently occurs during mass rearing to produce sterile males. We investigated the effect of mass-rearing conditions on the responsiveness to sex pheromones and spontaneous flight activity of males of the sweetpotato weevil Cylas formicarius (Coleoptera: Brentidae). There were no significant differences in the responsiveness to sex pheromones and spontaneous flight activity between wild and mass-reared strains. These results indicate that mass-reared strains of C. formicarius might not cause serious problems for implementing SIT programmes.

Behaviour during copulation can alter the fate of sperm of the mating males. This behaviour may exert selective pressure, resulting in the evolution of diverse reproductive behaviour, morphology, and physiology. This study examined the role of female copulatory behaviour on sperm fate in the sweet potato weevil, Cylas formicarius (Fabricius) (Coleoptera: Brentidae). In this species, males mount the female during copulation. The female frequently walks during copulation, carrying the male on her back. Here, we describe and quantify the copulatory behaviour of mating pairs and examine the sperm fate. Insemination success, as determined by the presence of sperm in the spermatheca, was lower when females walked for longer periods during copulation. This result emphasizes the value of studying variation in female copulatory behaviour in order to understand the factors that influence sperm fate. We discuss the implications of these results on sexual selection and utility in programs applying sterile insect techniques.

The West Indian sweet potato weevil Euscepes postfasciatus (Fairmaire) (Coleoptera: Curculionidae) is a major pest of sweet potato Ipomoea batatas (L.) in the tropical and subtropical regions. The sterile insect technique (SIT) could be used as one of the most effective methods for suppression or eradication of the weevil. The effectiveness of SIT depends on the ability of the released sterile males to mate with and inseminate wild females. However, the effect of sterile weevils on the fitness of E. postfasciatus has not been evaluated on natural density. Here, we investigated the effect of gamma-irradiated weevil density on the number of weevil progeny. When irradiated weevils were released in numbers equal to those of non-irradiated weevils, the number of progeny was reduced by half of that in the control treatment, and it remained at this state for 2 weeks. Our results show that irradiated weevils ensure adequate and efficient suppression of wild weevils. We conclude that the SIT programs will be employed as effective eradication method for E. postfasciatus.

The sweetpotato weevil Cylas formicarius (Fabricius) (Coleoptera: Brentidae) and West Indian sweetpotato weevil Euscepes postfasciatus (Fairmaire) (Coleoptera: Curculionidae) are major pests of sweet potato Ipomoea batatas (L.) Lam., in tropical and subtropical regions. Effects of intra- and inter-specific competition on the number of progeny (fecundity), body weight and developmental time of C. formicarius and E. postfasciatus were examined in single- and mixed-species treatments under laboratory conditions. Cylas formicarius tended to outcompete E. postfasciatus, whereas E. postfasciatus rather than C. formicarius tolerated higher conspecific densities. We discuss the implications of the results for pest management and resource partitioning of pestiferous weevils.

The effectiveness of the sterile insect technique (SIT) depends on the ability of released sterile males to mate with and inseminate wild females, but the use of gamma radiation to induce sterility negatively affects both somatic and reproductive cells of the sterilized insects. Recently, the effectiveness of fractionated-dose irradiation (FI), in which a sterilizing dose is delivered over time in a series of small irradiations, has been demonstrated in the West Indian sweetpotato weevil Euscepes postfasciatus (Fairmaire). FI improves male mating propensity compared with acute irradiation; however, this renewed technique takes a long time (72 h in the present circumstances) compared with the traditional technique (approximately 20 min) using single-dose irradiation (150 Gy) to fully sterilize this weevil. The extra time required by FI might negatively affect the quality of released sterile males, because weevils expend limited resources, such as metabolic energy or sperm, in mating freely in this period. We evaluated whether the temporal storage of weevils at low temperature (5°C and 15°C) improves the quality of sterile males compared with the normal condition (25°C). Temporal storage at low temperature in FI improves male mating propensity. For example, the sexually active phase of males exposed to 15°C was 18 days at least after irradiation. This period was longer than that of normal males (14 days). Meanwhile, this manner delayed male reproductive development and temporarily reduced mating competitiveness ability. If considering the long active phase of sterile males exposed to 15°C, these disadvantage would be cancelled out. We discuss the advantage of FI with temporal storage at low temperature in the eradication program using SIT for E. postfasciatus.

When there is an inbreeding depression, mating with a kin individual is generally considered maladaptive behaviour. However, in some conditions, the inclusive fitness benefits from inbreeding may outweigh the costs of inbreeding depression, and thus, inbreeding tolerance is often adaptive. Inbreeding depression and the effect of relatedness on mating behaviour in the West Indian sweet potato weevil Euscepes postfasciatus were examined. No significant inbreeding depression was detected as indicated by body weight and number of progeny emerging from sweet potato roots. Male mating performance (i.e. number of mating occurrences per night) was adversely affected by inbreeding depression, but the effect was low (fitness loss was 6.3%). Although there were no significant differences in latency to mounting, pre-copulatory guarding, copulation and post-copulatory guarding duration between full-sib and non-kin pairs, the copulation rate of full-sib pairs was significantly higher than that of non-kin pairs. These results support the theoretical prediction that when inbreeding depression is weak, copulation with close relative individuals is favoured.

Inadvertent selection is an important genetic process that frequently occurs during laboratory culture. The mass-reared strain of the sweet potato weevil Cylas formicarius exhibits stronger inbreeding depression than the wild strain does. When inbreeding depression occurs in a population, mating with a close relative is often considered maladaptive; however, in some contexts, the inclusive fitness benefits of inbreeding may outweigh the costs, favoring individuals that tolerate a low level of inbreeding depression. Theory predicts that mass-reared strain weevils will avoid inbreeding while wild strain weevils will tolerate inbreeding. To examine this prediction, we compared the effect of relatedness on the mating and insemination successes in mass-reared and wild strains of C. formicarius. While close relative pairs of the wild strain copulated less frequently than non-kin pairs, almost all mass-reared strain pairs copulated irrespective of relatedness. The results showed that the strain with weak inbreeding depression (wild strain) avoided inbreeding, whereas the strain with strong inbreeding depression (mass-reared strain) tolerated inbreeding. The contradiction between the theoretical prediction and our results is discussed from the perspective of laboratory adaptation, mating systems, and life history of C. formicarius.

Because life-history theory predicts that risky behaviours such as mating should increase as life expectancy decreases, predatory avoidance is expected to decrease with age. However, this prediction has not been examined. In the present study, the effect of age on death-feigning behaviour, a form of predatory avoidance behaviour in the sweetpotato weevil Cylas formicarius (Summers) (Coleoptera: Brentidae), is investigated by performing a longitudinal study. Because the effects of mating history and age usually cannot be distinguished, mating history is controlled. The results show that only female weevils decrease the investment in death-feigning behaviour with age, whereas male weevils do not show any age-related change. In addition, death-feigning behaviour of mated females is longer than that of virgin females, possibly because additional mating partners would be not needed by mated females.

The sterile insect technique (SIT) is widely used to suppress or eradicate target pest insect populations. Although the effectiveness of SIT depends on the ability of released sterile males to mate with and inseminate wild females, the use of gamma radiation to induce sterility negatively impacts reproductive cells as well as somatic cells. Consequently, sterilization by irradiation drastically diminishes mating performance over time. In the current study, we evaluated the effect of irradiation dose intensity on fertility, mating propensity, and mating competitiveness in sweetpotato weevil, Cylas formicarius elegantulus (Summers) (Coleoptera: Curculionidae), for 16 d after irradiation. Although the mating propensity of males irradiated with 200 Gy, the dose currently used to induce complete sterility of C. f. elegantulus in the SIT program in Okinawa Prefecture, was equal to that of nonirradiated weevils for the first 6 d, the mating propensity of males irradiated with doses between of 75 and 150 Gy was maintained for the first 12 d. The potential fertilization ability of weevils was highly depressed compared with the control weevils, even in those treated with 75 Gy. Mating performance was severely compromised in weevils that were irradiated with a dose of 100 Gy or more. These results demonstrate that partial sterilization can be highly advantageous in eradication programs for the sweetpotato weevil. We discuss the advantages of the application of partial irradiation in insect eradication programs.

In sterile insect technique (SIT) programs, the released sterile males are important for their effectiveness. The use of sex pheromones to collect males is economical; however, pre-exposure to sex pheromones may affect male mating behavior, and would thus reduce the effectiveness of the SIT programs. Males exposed to sex pheromone may become attractive to other males due to pheromone adsorption on their body impregnation, prevent mating and reduce the dispersal of sterile males. We examined the effect of pre-exposure to sex pheromone on the mating behavior of male sweetpotato weevil Cylas formicarius. Pre-exposure to high-dose sex pheromone for 24 h did not affect the male mating behavior of C. formicarius, and the exposed males did not attract other males. These results suggest that male C. formicarius that were collected from a mass-rearing facility using sex pheromone can be effectively used in SIT programs.

Inadvertent selection is an important genetic process that frequently occurs during laboratory culture and maintenance of biological control agents and other beneficial organisms used in procedures such as the sterile insect technique (SIT). We investigated effects of mass rearing and inbreeding depression on life history traits (number of progeny emerging from host plants, body weight, developmental period, and starvation tolerance) in the sweetpotato weevil, Cylas formicarius elegantulus (Summers) (Coleoptera: Brentidae). The effect of inbreeding was measured by comparing the results obtained from the full-sib crosses with those obtained from nonkin crosses in both wild and mass-reared strains. The mass-reared strain had more progeny than the wild strain. The developmental period of mass-reared strain was shorter than that of the wild strain. Other traits did not differ significantly between strains. We detected inbreeding depression effects on numbers of progeny, and the effects were more pronounced in the mass-reared strain. Hence, laboratory adaptation to mass rearing can produce changes in important biological attributes of sweetpotato weevils.

The sterile insect technique (SIT) is based on population and behavioral ecology and is widely used to suppress or eradicate target pest insect populations. The effectiveness of SIT depends on the ability of the released sterile males to mate with and inseminate wild females. The use of gamma-radiation to induce sterility is, however, associated with negative impacts not only on reproductive cells but also on somatic cells. Consequently, irradiation for sterilization diminishes mating performance over time. In this study, we evaluated the balance between the irradiation dose and both fertility and mating propensity in Euscepes postfasciatus (Fairmaire) (Coleoptera: Curculionidae) for 22 days following irradiation. The mating propensity of males irradiated with a 150-Gy dose, as currently used to induce complete sterility of E. postfasciatus in the SIT program in Okinawa Prefecture, was equal to that of non-irradiated weevils for up to 6 days, and the mating propensity of males irradiated with a dose of 125 Gy was equal to that of non-irradiated weevils for twice this period (12 days). The fertilization ability of weevils irradiated with a dose of 125 Gy was reduced by 4.6% in males and 0.6% in females, compared to the potential fertilization ability. We also discuss the possibility of the application of partially sterilized insects in eradication programs.

Persistent mating attempts by males (sexual harassment) are frequently observed among animals. For females, resisting persistent males can be costly because vigorous resistance increases both energy expenditure and the possibility of injury. Although one tactic for coping with male harassment is to cease resistance and mate with the persistent partner, the females of several species are able to prevent the fertilization of their egg(s) despite copulation. In this study, we used three different sex ratios to investigate whether a male’s mating persistence affects his mating success in the West Indian sweet potato weevil Euscepes postfasciatus, in which males mount females both before and after copulation. Consistent with our predictions, we found that female weevils resist and manipulate sperm transfer either before or during copulation according to their preferences. Female weevils were able to reject the sperm of persistent males despite having copulated with them. However, neither copulation and/or post-copulatory mounting affected insemination success. We speculate that the intensive resistance shown by females before copulation may induce mechanical sterility in E. postfasciatus.

Male body size is considered to be one of the major determinants of mating success among many insect species. Because the effectiveness of the sterile insect technique (SIT) depends on the ability of released sterile males to mate with and inseminate wild females, it is indispensable to understand the effect of male body size on the mating behavior of both sexes for the progress of the SIT program. We investigated how male body size and the presence of other rival males affect the guarding and copulatory durations of the West Indian sweetpotato weevil, Euscepes postfasciatus (Fairmaire) (Coleoptera: Curculionidae). In this species, males guard females before and after copulation. By observing the mating behavior under two sex-ratio conditions (male-to-female ratios of 1:1 and 2:1), we found that small males hastened to court females when rival males were present, but the females rejected these small males as mates. Therefore, we consider that female weevils adopt a counter-adaptation for mate preference in response to this male mating strategy. Body size did not affect the durations of copulation and post-copulatory guarding. Although we found a conditional mating strategy for body size in E. postfasciatus, it is unlikely to have a large influence on the weevil-eradication program using SIT.

The sterile insect technique (SIT) is widely used to suppress or eradicate target insect pest populations. The effectiveness of SIT depends on the ability of the released sterile males to mate with and inseminate wild females. Irradiation is effective for sterilizing mass-reared insects, and the negative impacts of this procedure are not limited to reproductive cell damage. In this study, we evaluated the death-feigning that is considered as an antipredator behavior of male Cylas formicarius (Summers) (Coleoptera: Blentidae) irradiated with 200 Gy, which is the dose used in the SIT program in Okinawa Prefecture. Irradiated male insects were observed in the laboratory for 10 d after irradiation. On d 0, fewer irradiated males feigned death than nonirradiated males. The number of irradiated and nonirradiated males who feigned death and the duration of their death-feigning were about the same from 2 d after irradiation. On the basis of our results, we discuss the optimal release time of irradiated m...

Although there have been numerous studies on the effects of mating history on mating behaviour, few studies have reported the relationship between mating history and other contextual behaviours such as foraging and predator avoidance. We examined the effect of mating history on death-feigning behaviour (an antipredator behaviour) in the sweetpotato weevil. Because mating behaviour can be divided into phases, we examined the effects of encounters with the opposite sex, copulation and insemination success on death-feigning behaviour. For females after copulation and males after multiple copulations the duration of death-feigning behaviour was reduced, whereas encounters with the opposite sex had no effect. Insemination success did not affect the duration of death feigning in males, but inseminated females reduced the duration of death feigning. We discuss the implications of these results for the effect of mating history on this antipredator behaviour.

The West Indian sweetpotato weevil Euscepes postfasciatus (Fairmaire) (Coleoptera: Curculionidae) is a major pest of sweetpotato Ipomoea batatas (L.) Lam (Solanales: Convolvulaceae) in some countries. In order to improve mass-rearing for an eradication program employing the sterile insect technique (SIT), optimal population density of E. postfasciatus in an artificial diet was examined. Six population densities (1000, 4000, 7000, 10000, 13000, and 16000 individuals per container with 200 g of artificial diet) were compared for effect on the number of eggs collected and hatchability. The total number of eggs collected after 24 d increased with an increase in population density and reached a saturation level at 13,000 individuals, whereas hatchability was not affected by population density. The results indicated that optimal population density in mass rearing was 13,000 individuals on 200 g of artificial diet. Furthermore, we examined cannibalism by adult weevils in the presence of other diets. The...

The West Indian sweetpotato weevil Euscepes postfasciatus (Fairmaire) (Coleoptera: Curculionidae) is a major pest of the sweet potato Ipomoea batatas (L.) Lam (Solanales: Convolvulaceae) in some countries. In order to improve the mass-rearing systems involved in an eradication program employing the sterile insect technique (SIT), we improved the efficiency of the method used for collecting adult weevils from artificial larval diets; this was done by using the combination of a sieve and Tullgren apparatus. Using this combination, we could collect 95.4% of weevils from artificial diets. Furthermore, the time spent in collection with this method was approximately one-fifth of that spent with the existing hand-sorting method. These results showed that this new collection method will improve the efficiency of mass rearing of West Indian sweetpotato weevils.

The sterile insect technique (SIT) is widely used for suppressing or eradicating target pest insect populations. In the eradication programs using the SIT, a large number of sterile insects are marked and released in the field. In Japan, Cylas fonnrmicarius (F.) group (Coleoptera: Brentidae) weevils are marked with a fluorescent powder dye to monitor the progress of such programs. However, this monitoring technique is not fully effective because of the disappearance or contamination of the dye. Therefore, an alternative marking method is required. Currently, a rare color morph such as piceous elytra (PE) is used for visible marking of C. formicarius group weevils. A PE-monomorphic strain has previously been established by artificial selection from a small locally distributed population; this can lead to reduced survival and genetic changes in behavioral traits due to inbreeding depression. In this study, we evaluated the survival rate and mating behavior of PE males of C. formicarius group. The characteristics of the PE males were similar to those of the wild strain (WS) males. Thus, we considered that PE males were suitable for visible marking in the eradication programs using the SIT.

The number of West Indian sweet potato weevils, Euscepes postfasciatus, being mass-reared in a facility for use in sterile insect technique (SIT) eradication programs has undergone a drastic reduction. A neogregarine protozoan pathogen Farinocystis sp. (an undescribed species) was detected in vivo in the mass-reared E. postfasciatus. We investigated the effects of this disease on the longevity and fecundity of host weevils and the incubation time of the disease in the host body under mass-rearing conditions. Our results demonstrated that infection by this Farinocystis sp. decreased both longevity and fecundity in E. postfasciatus. In particular, the pathogen severely limited the production of progeny by infected females compared to healthy females. Therefore, we consider this protozoan infection to be the major cause of the decreased E. postfasciatus production in the mass-rearing facility.

The aim of our study was to examine the effect of mass-rearing, in which there is no exposure to predators, on antipredator traits of insects for improving sterile insect technique programs. The duration of death-feigning (antipredator behaviour) in sweet potato weevil Cylas formicarius (Summers) (Coleoptera: Brentidae) after mass-rearing for 71 generations were compared with those in wild strain. There was no significant difference in the duration of death-feigning between wild and mass-reared strains. This indicated that the death-feigning behaviour of mass-reared strain was maintained for 71 generations even in the absence of predators. We discuss the reasons why death-feigning behaviour is maintained after mass-rearing.

Weevils constitute the most species-rich animal group with over 60,000 described species, many of which possess specialized symbiotic organs and harbor bacterial endosymbionts. Among the diverse microbial associates of weevils, Nardonella spp. represent the most ancient and widespread endosymbiont lineage, having co-speciated with the host weevils for over 125 million years. Thus far, however, no empirical work on the role of Nardonella for weevil biology has been reported. Here we investigated the biological role of the Nardonella endosymbiont for the West Indian sweet potato weevil, Euscepes postfasciatus. This insect is an experimentally tractable pest insect that can easily be reared on a natural diet of sweet potato root as well as on an agar-based artificial diet. By larval feeding on an antibiotic-containing artificial diet, Nardonella infection was effectively eliminated from the treated insects. The antibiotic-treated insects exhibited significantly lighter body weight and lower growth rate than the control insects. Then, the antibiotic-treated insects and the control insects were respectively allowed to mate and oviposit on fresh sweet potatoes without the antibiotic. The offspring of the antibiotic-treated insects, which were all Nardonella-negative, exhibited significantly lighter body weight, smaller body size, lower growth rate and paler body color in comparison with the offspring of the control insects, which were all Nardonella-positive. In conclusion, the Nardonella endosymbiont is involved in normal growth and development of the host weevil. The biological role of the endosymbiont probably underlies the long-lasting host-symbiont co-speciation in the evolutionary course of weevils.