Sperm Numbers and Female Fertility in the Moth Plodia interpunctella (Hübner) (Lepidoptera; Pyralidae)

I investigated two possible reasons for remating in female Plodia interpunctella: i) females remate to obtain sufficient sperm to maintain fertility; and ii) male investment in non-sperm components increases female fecundity and longevity. The number of sperm and the mass of the spermatophore transferred by males decreases on successive matings. Sperm numbers and potential male investment were varied by allowing females to mate either once or twice with males either on their first or second mating. Females receiving a single small spermatophore containing few sperm (from a male on his second mating) had sufficient sperm to fertilize all their eggs. Females did not show increased fecundity or longevity as a result of obtaining more spermatophore material. I discuss why females remate when they already have sufficient sperm to fertilize all their eggs.     

Effects of different male remating intervals on the reproductive success of Choristoneura rosaceana males and females

The mass of the spermatophore transferred by a previously mated Choristoneura rosaceana male increases with time elapsed since the last mating but, even after 4 days, it never reaches the mass of the spermatophore of a virgin male. However, spermatophore mass is clearly not a good indicator of the male reproductive investment as the quantity of sperm in the second ejaculate of a previously mated male is the same as that of his first, if he is allowed a 2 (eupyrene sperm) to 3 day (apyrene sperm) recovery period. The interval between the first two matings had no influence on female fecundity or longevity but significantly affected fertility if the male had only 1 day to recover. The length of the post-copulatory refractory period was also shorter in females mated with previously mated males than in those mated with virgins, regardless of the male's remating interval. Furthermore, a significant variation in the eupyrene sperm content of the spermatophore transferred by virgin males had no influence on the length of the female refractory period. Globally, these results support the hypothesis that a factor, other than sperm numbers in the spermatheca, is responsible for maintaining the inhibition of pheromone production in this species.     

Delayed mating and its cost to female reproduction in the butterfly,Eurema hecabe

The timing of mating of females under semi-natural condition, male ejaculate production and their effects on female fecundity were examined inEurema hecabe. Age of the first mating of females varied, and the number of matings increased with age. Male spermatophore production depended on age and body mass. The spermatophore mass at the second mating depended only on the interval between the first and second matings. The timing of the first mating and the spermatophore mass did not affect female fecundity. The timing of mating of females relative to the role of male spermatophores in female fecundity and male mating strategy are discussed.     

Multiple mating of male and female codling moth (Lepidoptera: Tortricidae) in apple orchards treated with sex pheromone

Studies were conducted with codling moth, Cydia pomonella L., to evaluate the mating status of male and female moths in apple, Malus domestica (Borkhausen), orchards treated with and without sex pheromone dispensers. Laboratory studies first examined the effect of multiple mating of male and female moths on female fecundity and egg fertility. Females that had mated three times had a significantly higher fecundity than singly mated moths. Sequential mating by male moths had no effect on the fecundity of female moths or egg fertility. However, male moth age did impact female fecundity, with significantly fewer eggs laid after mating with virgin 1- versus 3-d-old males. The mean size of the first spermatophore transferred by males was significantly larger than all subsequent spermatophores. Classifying spermatophores based on size was used in field sampling to categorize the mating status of the female's partner. The proportion of mated females with small spermatophores (partner had previously mated) was significantly higher in treated versus untreated orchards. The proportion of female moths caught in traps baited with pear ester that were virgin was low (相似文献   

Selection on female remating interval is influenced by male sperm competition strategies and ejaculate characteristics

Female remating rate dictates the level of sperm competition in a population, and extensive research has focused on how sperm competition generates selection on male ejaculate allocation. Yet the way ejaculate allocation strategies in turn generate selection on female remating rates, which ultimately influence levels of sperm competition, has received much less consideration despite increasing evidence that both mating itself and ejaculate traits affect multiple components of female fitness. Here, we develop theory to examine how the effects of mating on female fertility, fecundity and mortality interact to generate selection on female remating rate. When males produce more fertile ejaculates, females are selected to mate less frequently, thus decreasing levels of sperm competition. This could in turn favour decreased male ejaculate allocation, which could subsequently lead to higher female remating. When remating simultaneously increases female fecundity and mortality, females are selected to mate more frequently, thus exacerbating sperm competition and favouring male traits that convey a competitive advantage even when harmful to female survival. While intuitive when considered separately, these predictions demonstrate the potential for complex coevolutionary dynamics between male ejaculate expenditure and female remating rate, and the correlated evolution of multiple male and female reproductive traits affecting mating, fertility and fecundity.     

Male Remating Behavior and Its Effect on Female Reproductive Fitness in Cnephasia jactatana Walker (Lepidoptera: Tortricidae)

Male remating behavior and its effect on the female reproductive fitness of a New Zealand leafroller, Cnephasia jactatana, were investigated in the laboratory. With a recovery period of at least 24 h between matings, most males were able to mate four times and only about 25% could mate six times during their lifespan. Only 5% of males managed to mate twice within 1 day. Mated males transferred 31–51% smaller spermatophores than virgin males. When mating with a mated male, the female was subject to a 20–51% and 23–51% reduction in fecundity and fertility, respectively. With the increasing number of matings her partner had achieved before mating with her, the female's fertility declined significantly faster than fecundity. Courtship period and mating duration remained similar regardless of the male's mating history but males required increasingly longer latency to start courtship display with the increasing number of matings achieved.     

Male larval nutrition in Choristoneura rosaceana (Lepidoptera: Tortricidae): an important factor in reproductive success

This study examines how Choristoneura rosaceana male quality, as determined by larval diet, age and mating history, affects the reproductive success of both sexes. While the size of the spermatophore produced at first mating increased linearly with male age, the frequency of mating was significantly higher for middle-aged males (2–4 days old) than younger (0–2 days old) or older (6–8 days old) individuals, when both sexes were fed on artificial diet. However, the duration of copulation was longer in couples with older than younger males. The observed age-related changes in spermatophore size had no significant effect on female longevity, fecundity or fertility, suggesting no direct relationship between male investment and spermatophore size under these experimental conditions. Different larval food sources (artificial diet, maple and hazelnut) did not affect the proportion of 2-day-old virgin males that mated; however, the proportion that remated was significantly higher for males reared on high-quality food (maple and artificial diet) than those on hazelnut, a poorer food source. There was a 5-fold decline in spermatophore size between the first and second matings on all diets, but female reproductive output was reduced by only 25%. In contrast, while the first spermatophore produced by males on hazelnut was 1.5 times smaller than those produced on maple and artificial diet, the fecundity of their mates was 40% less than those mated with high-quality virgin males. These results provide additional support to the idea that spermatophore size is not a valuable indicator of male quality. Most tethered females placed in the field during the first flight period mated with virgin males (based on the size of the spermatophore), suggesting that female choice exists in this species. These results are discussed in relation to the incidence of polyandry in naturally occurring populations of Choristoneura and the potential use of size and/or chemical cues by females to assess male quality.     

Pattern of sperm storage and migration in the reproductive tract of the swallowtail butterfly Papilio xuthus: cryptic female choice after second mating

Females of the swallowtail butterfly Papilio xuthus L. (Lepidoptera: Papilionidae) mate multiply during their life span and use the spermatophores transferred to increase their longevity as well as fecundity. Sperm from different males may be stored in the sperm storage organs (bursa copulatrix and spermatheca). To clarify the pattern of sperm storage and migration in the reproductive tract, mated females are dissected after various intervals subsequent to the first mating, and the type and activity of sperm in the spermatheca are observed. When virgin females are mated with virgin males, the females store sperm in the spermatheca for more than 10 days. Sperm displacement is found in females that are remated 7 days after the first mating. Immediately after remating, these females flush out the sperm of the first male from the spermatheca before sperm migration of the second male has started. However, females receiving a small spermatophore at the second mating show little sperm displacement, and the sperm derived from the small spermatophore might not be able to enter the spermatheca. Females appear to use spermatophore size to monitor male quality.     

Ejaculate size, second male size, and moderate polyandry increase female fecundity in a seed beetle

Several hypotheses have been proposed to explain the evolutionof polyandry in species that provide nuptial gifts. When nuptialgifts are in the form of nutritional elements in the ejaculateand ejaculate size is correlated with male body size, femalescan accrue both direct (nutritional) and indirect (genetic)benefits from multiple mating. We examined remating decisionsin females of the seed beetle Stator limbatus and, using pathanalysis, examined the effects of male body size on the sizeof his ejaculate, the amount of ejaculate that was successfullytransferred to females, and the overall effect of these variableson female fecundity. Larger males produced larger ejaculatesand consequently transferred a larger ejaculate to females,but the effects on female fecundity differed between the females'first and second mates. Both larger first and second males wereable to transfer more of their ejaculate to females than weresmaller males. Both the total amount of ejaculate transferredby these males and polyandry (number of matings) were positivelycorrelated to female fecundity independently of each other.However, larger second males were more successful at stimulatingfemale fecundity independently of how much ejaculate they transferred.We also provide evidence that females are choosy during theirsecond mating opportunity. Both female choosiness and higherfemale investment after mating with larger second males suggestthat females may benefit from both direct and indirect effectsfrom multiple mating. We also conclude that male body size isunder both directional fecundity selection and directional sexualselection.     

Spermatophore and Sperm Allocation in Males of the Monandrous Butterfly Pararge aegeria: the Female’s Perspective

In insects, spermatophore production represents a non‐trivial cost to a male. Non‐virgin males have been shown to produce small spermatophores at subsequent matings. Particularly in monandrous species, it may be an issue to receive a sufficiently large spermatophore at the first and typically only mating. Females of the monandrous Speckled wood butterfly Pararge aegeria (L.) produce fewer offspring after mating with a non‐virgin male. After mating, females spend all their active time selecting oviposition sites and typically ignore other males. Here, we show that females did not discriminate between a virgin male and a recently mated male in our laboratory experiments. We demonstrate that the number of eupyrene sperm bundles relative to spermatophore mass differed with subsequent male matings. Males transferred a significantly smaller spermatophore after the first copulation, but the spermatophore mass did not decrease further with subsequent matings. However, the number of eupyrene sperm bundles decreased linearly. Therefore, there was proportionally more eupyrene sperm in the male’s second spermatophore compared with the first and the later spermatophores. Such a pattern has been shown in polyandrous species. Hence, it suggests that differences in sperm allocation strategy between polyandrous and monandrous butterflies may be quantitative rather than qualitative. There was also a tendency for females that had mated with a recently mated male to have higher propensity to remate than did females that had mated with a virgin male. We discuss the results relative to the mating system in P. aegeria, including female remating opportunities in the field and male mate‐locating behaviour.     

Ejaculate investment and attractiveness in the stalk‐eyed fly,Diasemopsis meigenii

The phenotype‐linked fertility hypothesis proposes that male fertility is advertised via phenotypic signals, explaining female preference for highly sexually ornamented males. An alternative view is that highly attractive males constrain their ejaculate allocation per mating so as to participate in a greater number of matings. Males are also expected to bias their ejaculate allocation to the most fecund females. We test these hypotheses in the African stalk‐eyed fly, Diasemopsis meigenii. We ask how male ejaculate allocation strategy is influenced by male eyespan and female size. Despite large eyespan males having larger internal reproductive organs, we found no association between male eyespan and spermatophore size or sperm number, lending no support to the phenotype‐linked fertility hypothesis. However, males mated for longer and transferred more sperm to large females. As female size was positively correlated with fecundity, this suggests that males gain a selective advantage by investing more in large females. Given these findings, we consider how female mate preference for large male eyespan can be adaptive despite the lack of obvious direct benefits.     

Effects of male mating interval on spermatophore formation,transfer, and subsequent female receptivity and fecundity in the sorghum plant bug,Stenotus rubrovittatus

Males of the sorghum plant bug, Stenotus rubrovittatus (Matsumura) (Heteroptera: Miridae), transfer a spermatophore to females during copulation. After a 1‐day interval between the first and second copulation, males transferred both sperm and a spermatophore to females during the second copulation. However, when male mating interval was <1 h, they transferred sperm but no spermatophores to females during the second copulation. Therefore, the male mating interval probably produces two types of mated females, those with and those without a spermatophore. Mated females of S. rubrovittatus do not remate for at least 3 days after mating, even when courted, and lay more eggs than virgin females at the beginning of the oviposition period. The effects of spermatophores on female sexual receptivity and fecundity were examined using mated females with or without a spermatophore. Only one of the 40 (2.5%) mated females with a spermatophore remated, whereas 10 of the 26 (38.5%) without a spermatophore remated. Furthermore, mated females with a spermatophore laid more eggs than those without a spermatophore. These results suggest that spermatophores participate in reducing female sexual receptivity and enhancing female fecundity in S. rubrovittatus.     

Reduction of mating receptivity and changes in longevity of rice leaf bug, <Emphasis Type="Italic">Trigonotylus caelestialium</Emphasis> (Heteroptera: Miridae), females after mating

To develop new and improved pest management strategies against Trigonotylus caelestialium (Kirkaldy) (Heteroptera: Miridae), a major pest of rice in Japan, it is important to understand the mating behavior of this species. In this study, we examined the effect of mating on subsequent mating receptivity and longevity in female T. caelestialium. After mating, females temporarily exhibited decreased mating receptivity. In addition, the cumulative remating frequency of females that mated with a male that had just mated with another female was higher than that of females who mated with virgin males. As a result, we hypothesized that the male ejaculate reduces female mating receptivity. Furthermore, mated females survived longer without food and water than virgin females; on the other hand, the life span of mated females with access to food and water was less than that for virgin females.     

Effect of Age, Body Weight and Multiple Mating on Copitarsia decolora (Lepidoptera: Noctuidae) Reproductive Potential and Longevity

Successful behavioral-based control methods rely on the accurate knowledge of the mating dynamics of the target insect. Age at first mating affects reproductive potential and the chance of multiple mating. The cabbage moth, Copitarsia decolora (Guenée) is an important pest of a number of commodities. We investigated combinations of age, body size and mating history to determine how these variables affect insect reproductive capacity. Fecundity and fertility decreased as the age of mating pairs increased, heavy and average sized females laid more eggs than light females. Female multiple mating did not enhance fecundity nor fertility potential. Furthermore, spermatophore size did not determine female re-mating behavior. However, female fecundity and fertility was related to the male mating history. Our results show that SIT is a valuable tool for controlling this pest.     

Evolutionary Consequences of Desiccation Resistance in the Male Ejaculate

Avoiding water loss for insects is critical for survival. Selection for reduced water loss will depend on trade-offs between resources allocated for reproduction and those allocated for resisting desiccation. However, we lack knowledge on how selection for desiccation resistance can affect the male ejaculate. Furthermore, as male ejaculate composition is complex, desiccation resistant females could evolve traits that enable them to derive longevity benefits from mating. Here, we assessed how selection for desiccation resistance impacts male testes and accessory gland size, protein content of these organs, female sperm storage and male ability to inhibit female remating behavior, in the Mexican fruit fly Anastrepha ludens. Additionally, we tested if mating increased longevity and fecundity in desiccation resistant females. Males selected for resistance to desiccation stress had smaller accessory glands and seminal vesicles and females mating with these males stored less sperm compared to control males. Females mating with resistant males had lower fecundity compared to females mating with control males. Desiccation resistant females lived longer than control females, yet this was irrespective of mating. Rapid evolutionary responses to hydric stress can have correlated effects in reproductive capabilities, which are not restricted to pre-copulatory traits. Trade-offs between resistance to desiccation stress are reflected in decreased allocation of resources to reproductive organs. Thus, production of the ejaculate may be costly for A. ludens males. Knowledge on the evolution of ejaculate traits and reproductive organ size in response to directional selection for desiccation resistance, will aid our understanding of differential sex-specific responses to environmental stress.     

Prolonged copulation and spermatophore size ejaculated in the sulfur butterfly,Colias erate (lepidoptera: pieridae) under selective harassments of mated pairs by conspecific lone males

Uninterrupted copula durations between virgin male and virgin female in the sulfur butterflyColias erate successfully lasted about 45 min in the greenhouse. Without harassment, the copula duration of virgin females with wild young males was similar to that in the greenhouse. The male structured a spermatophore about 40 min after copula initiation. Young males ejaculated bigger spermatophores than old males, while age did not influence copula duration. The mating behaviors of the wild males and the tethered females were observed and the copula duration was recorded. All copulating pairs were harassed by single males in the field, and the copula duration was increased, though there was no difference in spermatophore size. Single males harassed pairs with old males significantly longer duration than pairs with young males. Severe harassment elicited no particular response by the pairs. There was a successful mating takeover by a single male. Females tended to be more likely to re-mate when they had fewer spermatophores in the bursa copulatrix. The females that accepted re-mating retained smaller ejaculates in the bursa copulatrix than those that refused re-mating. Lone males seemed to be able to discriminate old and young males in copula. The harassment of pair with old males might be advantageous for single males to increase mating success.     

The function of multiple mating in oviposition and egg maturation in the seed beetle Callosobruchus maculatus

Abstract.  To comprehend the significance of multiple mating in female Callosobruchus maculatus , the material-benefits hypothesis is tested, as well as the potential function and mechanism of additional mating for females. The results show that longevity and lifetime fecundity are significantly higher in doubly-mated females, whereas females subject to an interrupted second copulation (which transfers no ejaculate) show resemblance with singly-mated females in lifetime fecundity and daily fecundity, supporting the material-benefits hypothesis. Female lifetime fecundity increases with ejaculate size in doubly-, but not singly-mated females. Doubly-mated females exhibit an immediate increase in fecundity on the day of remating. Moreover, dissection of ovaries after remating shows that the number of unlaid eggs is lower in doubly-mated females in comparison with singly-mated females, whereas the total number of unlaid and laid eggs shows no significant difference. This suggests that the second mating increases the oviposition rate, but not the egg maturation rate in female C. maculatus . Thus, ejaculate serves as a stimulus for oviposition, at least on the remating day. However, because doubly-mated females lay more eggs in total than singly-mated females, doubly-mated females appear to mature more eggs than the singly-mated counterparts. This is attributed to the 'indirect-driven' hypothesis, which states that the reduction in the number of mature eggs in the oviducts brought about by oviposition stimulates the maturation of oocytes in ovarioles to replenish the number of mature eggs.     

The effect of male mating history on paternal investment, fecundity and female remating in the seed beetle Callosobruchus maculatus

1. In many organisms, males provide nutrients to females via ejaculates that can influence female fecundity, longevity and mating behaviour. The effect of male mating history on male ejaculate size, female fecundity, female longevity and female remating behaviour in the seed beetle Callosobruchus maculatus was determined.
2. The quantity of ejaculate passed to females declined dramatically with successive matings. Despite the decline, a male's ability to fertilize a female fully did not appear to decline substantially until his fourth mating.
3. When females multiply mated with males of a particular mated status, the pattern of egg production was cyclic, with egg production increasing after mating. Females multiply mated to virgins had higher fecundity than females mated to non-virgins, and females mated to twice-mated males had disproportionately increased egg production late in their life.
4. Females that mated to multiple virgins, and consequently laid more eggs, experienced greater mortality than females mated only once or mated to non-virgins, suggesting that egg production is costly, and rather than ameliorating these costs, male ejaculates may increase them by allowing or stimulating females to lay more eggs.
5. Females mating with non-virgin males remated more readily than did females mated to virgins. Females given food supplements were less likely to remate than females that were nutritionally stressed, suggesting that females remate in part to obtain additional nutrients.     

Why Do Female Callosobruchus maculatus Kick Their Mates?

Sexual conflict is now recognised as an important driver of sexual trait evolution. However, due to their variable outcomes and effects on other fitness components, the detection of sexual conflicts on individual traits can be complicated. This difficulty is exemplified in the beetle Callosobruchus maculatus, where longer matings increase the size of nutritious ejaculates but simultaneously reduce female future receptivity. While previous studies show that females gain direct benefits from extended mating duration, females show conspicuous copulatory kicking behaviour, apparently to dislodge mating males prematurely. We explore the potential for sexual conflict by comparing several fitness components and remating propensity in pairs of full sibling females where each female mated with a male from an unrelated pair of full sibling males. For one female, matings were terminated at the onset of kicking, whereas the other’s matings remained uninterrupted. While fecundity (number of eggs) was similar between treatments, uninterrupted matings enhanced adult offspring numbers and fractionally also longevity. However, females whose matings were interrupted at the onset of kicking exhibited an increased propensity to remate. Since polyandry can benefit female fitness in this species, we argue that kicking, rather than being maladaptive, may indicate that females prefer remating over increased ejaculate size. It may thus be difficult to assess the presence of sexual conflict over contested traits such as mating duration when females face a trade off between direct benefits gained from one mating and indirect benefits from additional matings.     

Water Availability Affects Female Remating in the Seed Beetle,Callosobruchus chinensis

Benefits of multiple mating to females may come from the acquisition of water in male ejaculates. This hypothesis seems plausible in species in which males provide females with large ejaculates and has been tested with the prediction that females mate more frequently when an external source of water is unavailable. My study observed that females deprived of water were more likely to remate than females given water in the adzuki bean beetle, Callosobruchus chinensis. This result suggests that females may absorb the water in male ejaculates and thus change their remating receptivity according to the need for additional water. However, compared with related species, the ejaculate size is smaller, so ejaculatory hydration benefits are expected to be small in this species. There were no significant differences in lifetime fecundity and longevity between females that were allowed to receive one ejaculate from remating and females that were not allowed to do so when water was unavailable. This provides no evidence that receiving an additional ejaculate enhances female fitness. Thus, obtaining water from male ejaculates may partly compensate the costs of remating to females, although it alone would be insufficient to explain polyandry in C. chinensis. Increased mating frequency in water‐deprived females would not necessarily support the hypothesis that females remate for ejaculatory hydration benefits.