Effects of parental age at mating on the fecundity and progeny sex ratio of <Emphasis Type="Italic">Campoletis chlorideae</Emphasis>, an endolarval parasitoid of the pod borer, <Emphasis Type="Italic">Helicoverpa armigera</Emphasis>

Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae), commonly called gram pod borer, is a serious pest of pulse crops in India and damages chickpea (Cicer arietinum L). On average a 30% crop loss is reported. One of the potential natural enemies reported for its biological control is Campoletis chlorideae Uchida (Hymenoptera: Ichneumonidae), an idiobiont parasitic wasp that attacks second instar larvae of the pod borer. Male-biased sex ratios hinder efforts to mass release parasitic Hymenoptera for biological control by making the production of females costly. Parental age at time of mating is known to affect the progeny sex ratio in some Braconidae. In this view, we examined the reproduction and survival of the parasitoid C. chlorideae in the laboratory [22 ± 4°C, 70 ± 10% R.H. and 10:14 h light:dark photoperiod]. All the nine (i.e. 3 × 3) combinations of possible mating were made for males and females, being each of different age groups (0–12, 48–60 and 96–108 h old). Second instar larvae of the pod borer were reared on chickpea and exposed to mated females of different mating age group combinations. Results revealed that (a) newly emerged females (0–12 h old) mated to aged males yielded more progeny contrasted to the older females inseminated by newly emerged males (0–12 h old). This implies that increased maternal age at mating caused greater reduction in the progeny yield than that of paternal age; (b) the progeny sex ratio was lowest (0.344 ± 0.057) (mean ± SD) when parents mated just after emergence (0–12 h old). It became more and more male biased as parents were deprived of mate and was highest (0.666 ± 0.701) for 96–108 h old parents. The analysis of variance of the data shows that mate deprived males caused greater contribution in the production of sons, resulting a male-biased progeny sex ratio. Our conclusion indicate that newly emerged parasitoids (0–12 h old) are most fecund and should be used in rearing practices in order to enhance the progeny yield, particularly with respect to daughters. Handling Editor: Torsten Meiners     

Caddisfly egg mass morphology mediates egg predation: potential costs to individuals and populations

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Female presence enhances sexual performance of sterile Anastrepha ludens males of the Tapachula-7 GSS strain

The sterile insect technique (SIT), used for the control of many tephritid fly pests, is based on the rearing and release of large numbers of sexually competitive sterile insects into a wild population. In the interest of reducing expenses and increasing SIT effectiveness, genetic sexing strains (GSS) have been developed. These strains allow the production and release of only males. The objective of our study was to assess the effects of pre-release adult exposure to methoprene and to females on the mating propensity and mating competitiveness of GSS sterile males of Anastrepha ludens (Loew) (Diptera: Tephritidae). GSS sterile males were kept on a protein-sugar (protein-fed) or a protein-sugar-methoprene diet and were exposed to different proportions of females for the normal pre-release period of 5 days. Using laboratory and field-cage bioassays, we examined the influence of methoprene and female presence on the mating success of sterile males of 3–9 days old, in competition for wild females with untreated males and with wild males. Methoprene and female exposure had no significant effects on male mating success in the laboratory, whereas age had a positive relationship with the number of copulations observed. However, in field-cage bioassays, males exposed to females obtained a higher number of copulations than unexposed control males. Possible implications of these findings for programs that use GSS and especially for the campaign against Mexican fruit flies are discussed.     

Protective function of maternal care against egg predators in the shield bug Sastragala esakii (Hemiptera: Acanthosomatidae)

Females of many acanthosomatid shield bugs attend eggs and young nymphs by covering them with their bodies. Although such form of maternal care has been suggested to have evolved independently in four genera, Elasmucha, Sastragala, Acanthosoma and Sinopla, previous studies exploring its adaptive function have solely focused on species of Elasmucha. This study pioneered an experimental examination of maternal care in the Japanese species Sastragala esakii. Field experiments demonstrated that unattended egg masses suffered intense predation, whereas egg masses attended by their mothers were rarely preyed upon. The ant Crematogaster sp. was the most common egg predator, while two other insect species were also observed to prey on eggs. The exclusion of ant workers and other walking predators from accessing egg masses through the utilization of a sticky trap resulted in a remarkably improved egg survival rate in orphaned egg masses, suggesting that, at least in our study site, maternal care of S. esakii primarily functions to protect eggs from walking predators. Laboratory observations revealed that egg protection against predators was achieved through at least two distinctive defensive behaviors shown by egg-tending females: (i) tilting their bodies when approached by predators; and (ii) fanning their wings when in contact with predators. However, females displayed a limited response to predators approaching from the posterior direction, suggesting that visual cues play a significant role in predator recognition. These results indicate a similarity in the maternal care functions between Elasmucha and Sastragala, and suggest the parallel evolution of female defensive behaviors.     

The screw-worm flyChrysomya bezziana villeneuve (Diptera: Calliphoridae) in a sterile insect release trial in Papua New Guinea

The feasibility of the sterile insect release method (SIRM) was tested against natural populations of the Old World screw-worm fly, Chrysomya bezziana in the Musa Valley, Papua New Guinea. Sterile mating frequencies were determined from egg masses laid by native females on wounded, sentinel cattle. The aerial release of sterilised puparia resulted in low frequencies of sterile matings and few trap recaptures of released material. The release of chilled adult flies resulted in higher frequencies of sterile matings and many trap recaptures. The mean density of males released was 230 males per km² per week over a target area of 361 km² (48% of the valley). Sterile masses were first detected 2 weeks after the release of chill flies commenced, reaching a weekly peak of 33% after 5 weeks of releases with 15% of egg masses found to be sterile during the final month. The levels of sterile matings achieved in this trial were similar to comparable SIRM studies made in the USA, Mexico and Guatemala to control the New World screw-worm fly, Cochliomyia hominivorax.     

Effect of mating period and time‐of‐day for bloodmeal on rearing of Asian tiger mosquito (Aedes albopictus) in laboratory conditions

Colonization and maintenance of mosquitoes in the laboratory is required to study physiology, ecology, and behavior of mosquitoes and interactions between mosquito and pathogens. Artificial blood feeding systems have been widely used to maintain the laboratory colony of Aedes albopictus. In this study, we investigated the effects of mating period (1, 3, 6, and 10 days) and time‐of‐day for bloodmeal (08:00, 13:00, and 18:00) in the use of an artificial feeding system on blood‐feeding rate, female fecundity, egg hatching rate, and developmental time of the Asian tiger mosquito, A. albopictus. Younger females mated for three or fewer days reproduced more eggs compared to those of oldest females mated for ten days. Similar to the result for eggs laid, the mean egg‐hatching rate was significantly higher from the offspring of younger females than from those of older females. However, mating period and time‐of‐day for bloodmeal had no effect on the blood feeding rate and developmental time. Taken together, we suggest that three‐day mating with bloodmeal at 18:00 is optimal for maintaining colonies of A. albopictus in laboratory conditions.     

Effects of starvation on reproduction of the predacious mite <Emphasis Type="Italic">Neoseiulus californicus</Emphasis> (Acari: Phytoseiidae)

Effects of starvation on gravid females of Neoseiulus californicus were investigated at 20°C and 85% RH. When females that had been reared with abundant prey were swapped, just after laying their first egg, to conditions without any prey and water, they laid 1.8 eggs and survived for 4.3 days. In the body of well-fed females, an egg with eggshell and/or two oocytes were observed in the ventral and dorsal regions, respectively. The larger oocyte had two roundish nuclei and abundant yolk granules, and was enveloped with a vitelline membrane. These two nuclei were not fused but were just close to each other. The smaller oocyte had a nucleus, but had not yet formed yolk granules and vitelline membrane. Females after 12 h starvation had an egg in the ventral region and an oocyte in the dorsal region of the body. After more than 24 h starvation females maintained an oocyte in the dorsal region of the body, but had no egg in the ventral region. The oocyte was filled with abundant yolk granules and contained two irregular nuclei when females were starved for 24 h, but when starved for more than 36 h it contained one irregular nucleus. These findings suggest that (1) gravid females maintained an oocyte in the dorsal region after laying two eggs during starvation, (2) the oocyte was not absorbed during starvation, (3) the oocyte advanced vitellogenesis and the fusion of two nuclei, and (4) the vitellogenic oocyte was not enveloped with an eggshell and had not started embryogenesis.     

Life cycle and behaviour of the predaceous mite <Emphasis Type="Italic">Cunaxatricha tarsospinosa</Emphasis> (Acari: Prostigmata: Cunaxidae)

Life cycle and behaviour of Cunaxatricha tarsospinosa Castro & Den Heyer from rubber trees in Brazil were studied, with Tenuipalpus heveae Baker offered as prey. The study was conducted at 25.4 ± 0.2°C, 83 ± 5% RH and 12:12 h L:D photophase. The egg stage was the longest immature stage, lasting 17.1 ± 1.3 days (mean ± SE); total juvenile development was completed in 33.2 ± 2.8 days. Lifetime fecundity was 12.0 ± 2.2 eggs. Intrinsic rate of population increase was low, suggesting that T. heveae may not be a good prey for the predator. All specimens of C. tarsospinosa collected in the field for this study were females, no males were found. Concurrently, only females were obtained in the laboratory. This seems to be the first report of thelytokous parthenogenesis for cunaxids. Similar to earlier reports for some Cunaxinae and Coleoscirinae, prey were captured when predators were actively searching for them.     

Life history of a dipteran predator (Scathophagidae: Acanthocnema) of insect egg masses in a northern California stream

1. Little is known about the predators of insect eggs in fresh waters. This study describes aspects of the life history of a scathophagid fly (Acanthocnema sp.), whose larvae are predators of aquatic insect eggs. 2. Because the Acanthocnema predator oviposits its eggs on the surface of aquatic insect egg masses, all insect egg masses were collected regularly within a 200‐m reach of Redwood Creek (California, U.S.A.) between September 2003 and June 2007. Acanthocnema predators were found predominantly within egg masses of the caddifly Neophylax rickeri (Trichoptera: Uenoidae). 3. There was a mean of 0.25 Acanthocnema individuals per N. rickeri egg mass (n = 2367 egg masses). In general, N. rickeri egg masses were more commonly found clustered in aggregations (93.7%) than singly (6.3%), and Acanthonema were found more often within the aggregations of N. rickeri (98.7%) compared to singly laid egg masses (1.3%). 4. The duration of the Acanthocnema predator life stages was: egg 2.9 ± 0.8 (mean ± SD) days, larva 15.6 ± 10.2 days, pupa 80.3 ± 24.9 days and adult 7.2 ± 4.8 days. The short duration of the Acanthocnema egg stage (1–7 days) compared to that of its prey N. rickeri (2–4 weeks) raises the probability that the undeveloped eggs of N. rickeri would be available to the young predators upon hatching. Egg consumption of N. rickeri eggs by Acanthocnema averaged 262.6 eggs per larval period. 5. Acanthocnema had a bivoltine life cycle in which the first generation fed exclusively on N. rickeri egg masses in the winter and the second generation fed on the egg masses of several species, including other Trichoptera (Brachycentridae) and Diptera (Ceratopogonidae, Chironomidae) in the spring. These findings suggest differing feeding strategies by the first and second generations of Acanthocnema in response to the seasonal availability of prey species. This type of autecological information is important for understanding mechanisms of community interactions.     

Attractiveness and competitiveness of irradiated light brown apple moths

The sterile insect technique (SIT) potentially provides a socially acceptable approach for insect eradication of new pest incursions. The light brown apple moth, Epiphyas postvittana (Walker) (Lepidoptera: Tortricidae), was discovered in Berkeley (CA, USA) in 2006, leading to an incursion response that included this technology. In this study, we assessed factors affecting mating success from a bisex release of irradiated moths: effects of radiation dose on male multiple mating, male flight competition, female sex pheromone titre and attractiveness of irradiated females to males, and identification of successful mating in vineyards of either irradiated or wild males (identified by isotope analysis of spermatophores from sentinel females). There was a significant negative relationship between male radiation dose and mating frequency. In head‐to‐head flights of irradiated males against non‐irradiated males to a pheromone lure in a wind tunnel, irradiated males reached the lure first only 31% of the time. With increasing radiation dose, the production of the major sex pheromone component in females, (E)‐11‐tetradecenyl acetate, dropped, from 0.7 ± 0.1 ng per female in non‐irradiated females to 0.2 ± 0.07 ng per female when irradiated at 300 Gy. Male catch was reduced to 11% of control females in traps containing females irradiated at 300 Gy. Isotope analysis of spermatophores found in the bursa copulatrix of females indicated that mating success of irradiated males inside the live (entry‐only) traps containing virgin females was lower (13.1 ± 3.3%) than suggested by male catch (21.2 ± 3.8%) in pheromone traps, the current standard for assessing field competitiveness. Impacts of irradiation on male and female moth fitness should be taken into account to improve estimates of irradiated to wild male E. postvittana overflooding ratios needed for population suppression.     

Sex peptide receptor is required for the release of stored sperm by mated Drosophila melanogaster females

The storage of sperm in mated females is important for efficient reproduction. After sperm are transferred to females during mating, they need to reach and enter into the site(s) of storage, be maintained viably within storage, and ultimately be released from storage to fertilize eggs. Perturbation of these events can have drastic consequences on fertility. In Drosophila melanogaster, females store sperm for up to 2 weeks after a single mating. For sperm to be released normally from storage, Drosophila females need to receive the seminal fluid protein (SFP) sex peptide (SP) during mating. SP, which binds to sperm in storage, signals through the sex peptide receptor (SPR) to elicit two other effects on mated females: the persistence of egg laying and a reduction in sexual receptivity. However, it is not known whether SPR is also needed to mediate SP’s effect on sperm release. By phenotypic analysis of flies deleted for SPR, and of flies knocked down for SPR, ubiquitously or in specific tissues, we show that SPR is required to mediate SP’s effects on sperm release from storage. We show that SPR expression in ppk⁺ neurons is needed for proper sperm release; these neurons include those that mediate SP’s effect on receptivity and egg laying. However, we find that SPR is also needed in the spermathecal secretory cells of the female reproductive tract for efficient sperm release. Thus, SPR expression is necessary in both the nervous system and in female reproductive tract cells to mediate the release of stored sperm.     

Female response to predation risk alters conspecific male behaviour during pre‐copulatory mate guarding

Mating behaviour often increases predation risk, but the vulnerability within mating pairs differs between the sexes. Such a sex difference is expected to lead to differences in responses to predation risk between the sexes. In the two‐spotted spider mite Tetranychus urticae, males engage in pre‐copulatory mate guarding because only the first mating results in fertilisation. We investigated (i) whether pre‐copulatory pairs are more conspicuous to the predatory mite Phytoseiulus persimilis than solitary females, (ii) whether the vulnerability to the predator differs between sexes within the pre‐copulatory pair, (iii) whether each sex of T. urticae responds to predation risk during pre‐copulatory mate guarding and (iv) whether T. urticae's response to predation risk affects predator behaviour. Because T. urticae females are immobile during pre‐copulatory mate guarding, we observed male behaviour to evaluate effects of predation risk. We found that the predators detect more pre‐copulatory pairs than solitary females and that more females than males of the pre‐copulatory pairs are preyed upon by the predators. The preference of spider mite males for pre‐copulatory pairs versus solitary females was affected by whether or not the female had been exposed to predators during development. Male T. urticae exposed to predation risk did not alter their behaviour. These results suggest that only the most vulnerable sex, that is the female, responds to predation risk, which modifies male behaviour. Regardless of T. urticae females’ experience, however, P. persimilis detected more T. urticae pre‐copulatory pairs than solitary females, suggesting that pre‐copulatory mate guarding itself is dangerous for T. urticae females when these predators are present. We discuss our results in the context of sex‐dependent differences in predation risk.     

Interrelation of mating,flight, and fecundity in navel orangeworm females

The navel orangeworm, Amyelois transitella (Walker) (Lepidoptera: Pyralidae, Phycitini), is an economically important pest of nut crops in California, USA. Improved management will require better understanding of insect dispersal, particularly relative to when mating occurs. A previous study demonstrated a more robust laboratory flight capacity compared to other orchard moth pests, but it was unclear how mating affects dispersal, and how dispersal affects fecundity. In this study, 1‐ and 2‐day‐old females were allowed to fly overnight on a flight mill either before or after mating, respectively, and were then allowed to oviposit. Data on fecundity were compared between treatments to minimally handled or tethered‐only control females. Females that mated before flight flew longer and covered a greater distance than those flying prior to mating. However, timing of flight relative to mating did not affect fecundity, nor did any measure of flight performance. There was no effect on fecundity when females were forced to fly for designated durations from 3 min to 2 h. Together, our data revealed no obvious trade‐off between flight activity and reproductive output. Distances measured on the flight mills (mean ca. 15 km for mated females) may overestimate net displacement in the field where flight tracks are often meandering. The results suggest that most females mate and oviposit in or near their natal habitat, but that some may disperse potentially long distances to oviposit elsewhere.     

Sexual attractiveness shared by both sexes mediates same‐sex sexual behaviour in the parasitoid wasp Telenomus triptus

The mating behaviour of a quasi‐gregarious egg parasitoid Telenomus triptus Nixon (Hymenoptera: Platygastridae), which exploits egg masses of a stink bug Piezodorus hybneri (Heteroptera: Pentatomidae), is examined in the laboratory. In this parasitoid wasp, male adults that emerge earlier stay at the natal egg mass and mate with subsequently emerging females. In the present study, a male adult that encounters the emergence of another male always waits for it to egress, and then mounts the newly emerging male. To examine why males of T. triptus show same‐sex sexual behaviour, male adults are presented with a parasitized host egg mass or a freshly killed wasp. Male adults are observed to remain at host egg masses from which only male wasp(s) had emerged. In addition, male adults attempt to copulate with freshly killed young male wasps. It is suggested that newly emerging male wasps are targets of same‐sex sexual behaviour because they possess cues for male sexual behaviour similar to the cues of females. Both the sex and age of freshly killed wasps affect the frequency of the sexual behaviour of male adults: females are more attractive than males, although their attractiveness declines with age. When the mating opportunity is restricted to the natal egg mass, the costs of failing to notice newly emerging female adults should be extremely high. Therefore, males are forced not to discriminate the sex, resulting in same‐sex sexual behaviour.     

Swarming and mating activity of Anopheles gambiae mosquitoes in semi‐field enclosures

Anopheles gambiae Giles sensu stricto (Diptera: Culicidae) is the major Afro‐tropical vector of malaria. Novel strategies proposed for the elimination and eradication of this mosquito vector are based on the use of genetic approaches, such as the sterile insect technique (SIT). These approaches rely on the ability of released males to mate with wild females, and depend on the application of effective protocols to assess the swarming and mating behaviours of laboratory‐reared insects prior to their release. The present study evaluated whether large semi‐field enclosures can be utilized to study the ability of males from a laboratory colony to respond to natural environmental stimuli and initiate normal mating behaviour. Laboratory‐reared males exhibited spatiotemporally consistent swarming behaviour within the study enclosures. Swarm initiation, peak and termination time closely tracked sunset. Comparable insemination rates were observed in females captured in copula in the semi‐field cages relative to females in small laboratory cages. Oviposition rates after blood feeding were also similar to those observed in laboratory settings. The data suggest that outdoor enclosures are suitable for studying swarming and mating in laboratory‐bred males in field‐like settings, providing an important reference for future studies aimed at assessing the comparative mating ability of strains for SIT and other vector control strategies.     

Estimate of mating pattern of a paternal nest brooder goby of <Emphasis Type="Italic">Rhinogobius</Emphasis>, using egg density in the nest

The mating pattern of the paternal nest brooder Rhinogobius sp. CB (cross band type) exhibits both monogamy and polygyny. However, it is difficult to infer the actual conditions of their mating from their egg stages in the clusters, because multiple females spawn simultaneously or sequentially within short intervals. I examined whether the egg density of the egg clusters could be used as an index of the number of females, based on laboratory and field experiments. Egg densities increased with the number of spawning females. Egg densities of egg clusters collected in the field were categorized into three normal distributions which corresponded to those for the number of mating females in the spawning experiment. Egg density in the nest, therefore, seems to be a useful index for clarifying the mating pattern of this species in the field. By adopting the present method, it is suggested that this species frequently mates polygynously in the wild.     

Egg mass destroying behaviour of the female giant water bugLethocerus deyrollei Vuillefroy (Heteroptera: Belostomatidae)

After being laid on emergent aquatic vegetation, the egg masses ofLethocerus deyrollei are brooded by the male. In laboratory studies, females were observed to destroy egg masses and ingest the fluid of eggs. Brooding males fought with these attackers at first, but then gave up the defense and mated with them. After destroying the egg masses, females laid new ones at the same sites on the same night or the following one, and males brooded the new offspring. The population density of this species is very low. Most males engage in brooding and cease to become acceptable mates after most of the females lay the first egg masses. It is costly for females to search out free males in the aquatic vegetation. By destroying egg masses, however, individual females can gain new mates with little exertion and can make them brood their own offspring. Furthermore, the survival of their own offspring increases with the elimination of their competitors.     

Remating in Ceratitis capitata sterile males: Implications in sterile insect technique programmes

Sterile insect technique (SIT) is used, among other biological control tools, as a sustainable measure for the management of Ceratitis capitata Wiedemann (Diptera: Tephritidae) in many agricultural regions where this pest can trigger severe economic impacts. The tendency of wild females to remate multiple times has been deeply studied; it has been a common point of controversy when evaluating SIT programmes. Nevertheless, the remating potential of the released sterile males remains unknown. Here, under laboratory conditions, the remating capability of mass-reared sterile males was determined. Wild-type virgin females were offered to sterile males (Vienna-8 strain), which had the opportunity to mate up to four consecutive times. The remating assays were carried out at 24 hr, 48 hr, 4 days and 7 days after the first mating. At the end of each tested time period, males were divided according to their mating response, mated or unmated, and subsequently reused for the next round of mating assays. The frequency of successful remating in each tested time period was obtained. Insemination was confirmed by determining the sperm transfer in mated female spermathecae by quantitative real-time PCR. Our results demonstrate that 73% of the mass-reared sterile males were able to remate 24 hr after the first mating, 55% of which remated again the day after. Close to 25% of the V8 sterile males tended to copulate in all of the four mating opportunities. The qPCR analysis of the spermathecae contents verified an effective transfer of V8 sperm to wild females with every mating; 99% of copulations resulted in sperm transfer. These findings shed light on the remating potential of V8 sterile males, an aspect until now underestimated in many SIT programmes.     

Mating behaviour and mating success in woodfrogs Rana sylvatica

Mating behaviour of woodfrogs was investigated in a woodland pond in southeastern Michigan. Males became sexually mature one year before females and outnumbered females by 5.6 to 1 at the breeding site. Males employed searching behaviour to obtain mates. Yearly male mating success varied from 0 to 2 matings, and larger males had a greater probability of mating than smaller males. The mating season lasted less than 10 days. Most egg masses were deposited in 1 m² of a 256-m² pond. Experimental introduction of egg masses to the breeding site indicated that presence of eggs was sufficient to stimulate egg deposition. The influence of selection for synchronous sexual receptivity in females, communal egg deposition, and delayed female sexual maturation on male mating behaviour and variation in male mating success is discussed.