Butterflies tailor their ejaculate in response to sperm competition risk and intensity

Males of many insects eclose with their entire lifetime sperm supply and have to allocate their ejaculates at mating prudently. In polyandrous species, ejaculates of rival males overlap, creating sperm competition. Recent models suggest that males should increase their ejaculate expenditure when experiencing a high risk of sperm competition. Ejaculate expenditure is also predicted to vary in relation to sperm competition intensity. During high intensity, where several ejaculates compete for fertilization of the female''s eggs, ejaculate expenditure is expected to be reduced. This is because there are diminishing returns of providing more sperm. Additionally, sperm numbers will depend on males'' ability to assess female mating status. We investigate ejaculate allocation in the polyandrous small white butterfly Pieris rapae (Lepidoptera). Males have previously been found to ejaculate more sperm on their second mating when experiencing increased risk of sperm competition. Here we show that males also adjust the number of sperm ejaculated in relation to direct sperm competition. Mated males provide more sperm to females previously mated with mated males (i.e. when competing with many sperm) than to females previously mated to virgin males (competing with few sperm). Virgin males, on the other hand, do not adjust their ejaculate in relation to female mating history, but provide heavier females with more sperm. Although virgin males induce longer non-receptive periods in females than mated males, heavier females remate sooner. Virgin males may be responding to the higher risk of sperm competition by providing more sperm to heavier females. It is clear from this study that males are sensitive to factors affecting sperm competition risk, tailoring their ejaculates as predicted by recent theoretical models.     

Vibratory courtship in a web-building spider: signalling quality or stimulating the female?

Courtship behaviour in spiders in the form of premating vibrations by males may function (1) as a male identity signal used for species recognition, (2) in suppression of female aggressiveness, (3) to stimulate female mating behaviour, or (4) as a quality signal used in female choice. We investigated the function of web vibration by male Stegodyphus lineatus in a series of experiments. Regardless of vibratory performance, all males mated successfully with virgin females but only 56.4% of males mated with nonvirgin females. Vibratory performance did not influence male mating success, but heavier males had a higher probability of mating with mated females. Males vibrated less often and produced fewer vibrations when introduced on the web of a mated female. Males that vibrated webs of virgin females mated faster than nonvibrating males, but there was no effect of vibration rate or body mass. There was no effect of male vibratory effort or vibration rate on female reproductive success measured as time to egg laying, clutch size, number of hatched young, number of dispersed young and offspring body mass after a single mating. Males vibrated on abandoned virgin female webs but the response decreased with increasing duration of female absence, suggesting that females produce a web-borne pheromone, which elicits male vibrating behaviour. Mated females were less receptive and not stimulated by male vibrating behaviour. We conclude that male premating vibrations in S. lineatus do not function as a male quality signal selected via female choice. Rather, the primary function of this behaviour may be to stimulate a receptive female to mate. Copyright 2003 Published by Elsevier Ltd on behalf of The Association for the Study of Animal Behaviour.      

Host‐searching responses to herbivory‐associated chemical information and patch use depend on mating status of female solitary parasitoid wasps

1. In a tritrophic interaction system consisting of plants, herbivores, and their parasitoids, chemicals released from plants after herbivory are known to play important roles for many female parasitoids to find their hosts efficiently. On the plant side, chemical information associated with herbivory can act as an indirect defence by attracting the natural enemies of the host herbivores. 2. However, mated and virgin females of haplodiploid parasitoids might not necessarily respond to such chemical cues in the same way. Since virgin females can produce only sons, they might refrain from searching for hosts to invest eggs until copulation, in order to produce both sexes. 3. Here, we investigated differential host‐searching behaviours shown by mated and virgin females in the solitary parasitoid wasp, Cotesia vestalis, in response to herbivory‐associated chemical information from cruciferous plants infested by their host larvae, Plutella xylostella. 4. Mated females showed a significantly higher flight preference for host‐infested plants over intact plants, while no preference was observed with virgin females. Mated females also showed more intensive antennal searching and ovipositor probing behaviours to leaf squares with wounds caused by hosts than did virgin females. Furthermore, mated females stayed longer in host patches with higher parasitism rates than virgin females. 5. These results indicate that mating status of C. vestalis females clearly influences their host‐searching behaviour in response to herbivory‐associated chemical information and patch exploitation. Female parasitoids seem to forage for hosts depending on their own physiological condition in a tritrophic system.     

Is Male Cohabitation Costly for Females of the Spider Stegodyphus lineatus (Eresidae)?

Reproductive interests of females and males can vary over a number of issues, including the number of matings and the occurrence and duration of mate guarding and cohabitation. The mating system of the spider Stegodyphus lineatus (Eresidae) is characterized by an exceptional sexual conflict where males induce females to remate by committing infanticide. Females experience high costs because of this male strategy, while males benefit. During the mating season, males tend to stay with females for several days. We examined whether this male strategy of cohabitation is also disadvantageous for females of S. lineatus. A field experiment revealed that male presence in a female's nest negatively affected her body condition, whereas cohabiting males gained weight because they fed on prey caught in webs of females. As a response, females did not renew their webs when males were present. Thus, females with a cohabiting male experienced the combined cost of prey loss and loss of time available for foraging. These costs are expected to increase with every additional cohabiting male. Mated females behaved more aggressively toward males than did virgin females. This behavior may be interpreted as an adaptive response to reduce mating costs.     

Sperm numbers,their storage and usage in the fly Dryomyza anilis

In the fly Dryomyza anilis females have two kinds of sperm storage organs: one bursa copulatrix and three spermathecae (two spermathecae with a common duct form the doublet, and the third is a singlet spermathecal unit). At the beginning of a mating the male deposits his sperm in the bursa copulatrix. After sperm transfer the male taps the female''s abdomen with his claspers. This behaviour has been shown to increase the male''s fertilization success. After mating, the female discharges large quantities of sperm before oviposition. To find out where the sperm remaining in the female are stored, I counted the number of sperm in the droplet and in the female''s sperm storage organs after different types of mating. I carried out three mating experiments. In experiment 1, virgin females were mated with one male and the matings were interrupted either immediately after sperm transfer or after several tapping sequences. The results show that during male tapping more sperm moved into the singlet spermatheca. In addition, the total number of sperm correlated with sperm numbers in all sperm storage organs, and male size was positively related to the number of sperm remaining in the bursa. In experiment 2, females mated with several males. The number of sperm increased with increasing number of matings only in the doublet spermatheca. No increase in the number of sperm in the singlet spermatheca during consecutive matings suggests that sperm were replaced or did not reach this sperm storage organ. In experiment 3, virgin females were mated with a single male and half of them were allowed to lay eggs. The experiment showed that during egglaying, females primarily used sperm from their singlet spermatheca. The results from the three experiments suggest that sperm stored in the singlet spermatheca is central for male fertilization success and male tapping is related to sperm storage in the singlet spermatheca. The different female''s sperm storage organs in D. anilis may have separate functions during sperm storage as well as during sperm usage.     

Sexual Inhibition in <Emphasis Type="Italic">Spalangia endius</Emphasis> Males After Mating and Time for Ejaculate Replenishment

Although males are often stereotyped as always sexually responsive, a previous study with the parasitoid wasp Spalangia endius showed that males exhibit a post-mating decrease in sexual responsiveness. The present study examined the duration of the decrease and whether waiting to remate increased the amount of ejaculate that a male provided to his second mate. Mated males were back to being as sexually responsive as virgin males by about 5 min after mating. The amount of ejaculate that females received from already mated males was not significantly related to the duration between the male’s first and second mating. The short-term decrease in sexual responsiveness of a mated male may prevent him from immediately trying to remate with the same female because it gives her time to get away.     

Impact of mating on Anagyrus kamali Moursi (Hym., Encyrtidae) lifetime fecundity, reproductive longevity, progeny emergence and sex ratio

Abstract: The solitary endoparasitoid Anagyrus kamali Moursi (Hym., Encyrtidae) and the Hibiscus mealybug Maconellicoccus hirsutus Green (Hom., Pseudococcidae), were used as a parasite/host model to test the effect of mating on several fitness parameters, i.e. longevity, lifetime fecundity, progeny emergence and sex ratio. At 27 ± 2°C, 8 h light : 16 h dark, mating significantly affected the survival of male parasitoids. Virgin males lived longer (32.2 ± 9.51 days) than mated males (23.9 ± 7.52 days). Female longevity (40.7 ± 16.3 days for virgins and 36.2 ± 10.7 days for mated females) was not affected by mating. The lifetime fecundity of female parasitoids and their oviposition period was not significantly affected by mating. However, the number of hosts parasitized was greater for mated wasps (7.54 ± 4.85 hosts parasitized/day) compared with virgin ones (5.12 ± 2.19 hosts parasitized/day). This resulted in greater progeny production from mated A. kamali females. The progeny of virgin females consisted only of males, whereas the mated ones had a more female‐biased sex ratio. The lowest sex ratio (0.41 M/F ± 0.123) was attained when females had free access to males and were multi‐mated.     

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.     

Mating Status Effects on Sexual Response of Males and Females in the Parasitoid Wasp <Emphasis Type="Italic">Urolepis rufipes</Emphasis>

Although mate preferences are most commonly examined in females, they are often found in both sexes. In the parasitoid wasp Urolepis rufipes, both female and male mating status affected certain aspects of sexual interactions. Female mating status mattered only in the later stages of mating. Males did not discriminate between virgin and mated females in terms of which they contacted or mounted first. However, once mounted, most virgin females were receptive to copulation, whereas very few mated females were. Whether a male’s mating status affected his own sexual response depended on the female’s ability to respond and the stage of mating. Examining male behavior toward dead females allowed elimination of the role of female behavior in how males responded. Virgin and mated males are both attracted to dead females as evidenced by their fanning their wings at such females. However, mated males were quicker than virgin males to contact and to mount in an experiment with dead females, whereas there was no such differential response in an experiment with live females. This difference is consistent with greater female sexual responsiveness to virgin males. Male mating status also affected female receptivity to copulate. Once mounted, live virgin females were less likely to become receptive to copulation by mated males than to virgin males, but only in a choice experiment, not in a no-choice experiment.     

Female Remating Behavior in a Lekking Moth

In polyandrous species, male reproductive success will at least partly be determined by males' success in sperm competition. To understand the potential for post‐mating sexual selection, it is therefore important to assess the extent of female remating. In the lekking moth Achroia grisella, male mating success is strongly determined by female choice based on the attractiveness of male ultrasonic songs. Although observations have indicated that some females will remate, only little is known about the level of sperm competition. In many species, females are more likely to remate if their first mating involved an already mated male than if the first male was virgin. Potentially, this is because mated males are less well able to provide an adequate sperm supply, nutrients, or substances inhibiting female remating. This phenomenon will effectively reduce the strength of pre‐copulatory sexual selection because attractive males with high mating success will be more susceptible to sperm competition. We therefore performed an experiment designed both to provide a more precise estimate of female remating probability and simultaneously to test the hypothesis that female remating is influenced by male mating history. Overall, approximately one of five females remated with a second male. Yet, although females mated to non‐virgin males were somewhat more prone to remate, the effect of male mating history was not significant. The results revealed, however, that heavier females were more likely to remate. Furthermore, we found that females' second copulations were longer, suggesting that, in accordance with theory, males may invest more sperm in situations with an elevated risk of sperm competition.     

Mating Effect on Sex Pheromone Production of the Oriental tobacco budworm,Helicoverpa assulta

This study was undertaken to clarify the suppression phenomenon of sex pheromone production after mating and its relationship to the physiological mechanism in adult females of Helicoverpa assulta, and determine the mating factor from males causing depletion of sex pheromonc production. Sex pheromone production of H. assulta females was mostly terminated in 3 hours after mating. Mated females maintained with a low titer of sex pheromone until 3 days when it started to increase again, which showed a characteristic of species mating more than once. The mated female again produced pheromone upon injection of pheromone biosynthesis activating neuropeptide (PBAN) or extracts of brain-suboesophageal ganglion complexes (Br-Sg) of mated female, which were shown similar pheromonotropic activities as compared with virgin females. These results indicated that the mating did not inhibit the receptivity of pheromone gland itself and PBAN biosynthesis in suboesophageal ganglion of the mated females. And it seems to support that the depletion of sex pheromone production is responsible for blocking of PBAN release from head. To investigate the mating factor from adult males, when extracts of reproductive organs of male were injected into hemocoel of virgin females evoking depletion of sex pheromone production as shown in mated female. The results suggest that a chemical substance(s) from the male reproductive organs could be responsible for the loss of sex pheromone biosynthesis in H. assulta.     

Virgin and recently mated males are equally able to achieve new matings in Neoleucinodes elegantalis

Lepidopteran male mating success is recognized to be directly related to physical and behavioural traits such as ability, vigour, activity and persistence in courtship. In the tomato fruit borer Neoleucinodes elegantalis Guenée, the mating system is known to be monogamic and therefore males' sexual investment is apparently low. The hypothesis that recently mated males have a remating probability equal to that of virgin males is tested. The impact of body size in remating success and the cost of remating are also analyzed. Mated males show as much propensity to remate as naïve ones. Copula duration and the time taken to copulate are similar in mated and virgin males. However, spermatophore size is not related to male size. The results suggest the ability of N. elegantalis males to remate within 24 h between mating events and their propensity to remate are not affected by copula investment. © 2013 The Royal Entomological Society     

Virgins in the wild: mating status affects the behavior of a parasitoid foraging in the field

In haplodiploid organisms, virgin females can produce offspring, albeit only sons. They may therefore face a trade-off between either: (1) searching for hosts and producing sons immediately; or (2) searching for mates and perhaps producing both sons and daughters later in life. Although this trade-off raises a theoretical interest, it has not been approached experimentally. The objective of this article is thus to document the effect of mating status on the foraging behavior of a haplodiploid parasitoid. For this, we recorded the behavior of virgin and mated female Lysiphlebus testaceipes (Hymenoptera: Braconidae) after being released, in the field, on a colony of their aphid hosts. Half of the virgin females were mated by a wild male after less than 10 min of foraging. Evidently, virgin females attract males while foraging on host patches, so that the two activities are not mutually exclusive. Nonetheless, virgin females stayed motionless more often and for longer periods than mated females. Consequently, they attacked aphids at a lower rate, and in turn, attacked fewer aphids on each patch. Moreover, contrary to mated females, virgins did not aggregate their progeny on large patches. We conclude that in L. testaceipes, the trade-off may not be as hypothesized. By dispersing across patches more than mated females, virgins could promote future mating opportunities for their sons and increase their inclusive fitness. However, by moving too frequently, females may lose immediate mating opportunities for themselves and the immediate advantage of producing offspring of both sexes. The observed behavior of virgin L. testaceipes females on host patches could reflect an optimal solution to such a trade-off.     

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.     

Body size influences mating success of the Eucalyptus longhorned borer (Coleoptera: Cerambycidae)

Both sexes of adultPhoracantha semipunctata F. (Coleoptera: Cerambycidae) congregate on stressedEucalyptus that are the larval hosts. In a field study, 721 adultP. semipunctata captured on host trees varied considerably in body size with the largest individuals being about twice the length of the smallest. Females that were paired with a mate were similar in size to solitary females, suggesting that the probability of a female being mated was not affected by her size. However, large males had greater success than smaller males in obtaining mates. MaleP. semipunctata rely on antennal contact to locate and identify females on the larval host. Therefore, the rate at which males search for mates is a function of the area swept by their antennae per unit time. Because of their greater antennal spread, large males were able to search for females at double the rate of the smallest males. Large males also dominated in aggressive contests for females. The superior abilities of large maleP. semipunctata in both locating and defending mates account for the influence of body size on mating success.     

Host selection by the autoparasitoid Encarsia pergandiella on primary (Bemisia tabaci) and secondary (Eretmocerus mundus) hosts

In autoparasitoids, females are generally primary endoparasitoids of Hemiptera, while males are hyperparasitoids developing in or on conspecific females or other primary parasitoids. Female‐host acceptance can be influenced by extrinsic and/or intrinsic factors. In this paper, we are concerned with intrinsic factors such as nutritional status, mating status, etc. We observed the behavior of Encarsia pergandiella Howard (Hymenoptera: Aphelinidae) females when parasitizing primary (3rd instar larvae of Bemisia tabaci Gennadius [Homoptera: Aleyrodidae]) and secondary hosts (3rd instar larvae and pupae of Eretmocerus mundus Mercet [Hymenoptera: Aphelinidae]) for a period of 1 h. Females had different reproductive (virgin or mated younger) and physiological (fed elder or mated elder) status. Virgin females killed a large number of secondary hosts while investing a long time per host. However, they did not feed upon them. Mated females killed a lower number of secondary hosts and host feeding was observed in both consuming primary and secondary hosts. It was common to observe host examining females of all physiological statues tested repeatedly stinging the same hosts when parasitizing, killing or rejecting them. Fed elder females parasitized more B. tabaci larvae than E. mundus larvae or pupae, while investing less time on the primary host than on the secondary host. They also parasitized more B. tabaci larvae than mated elder females, while investing less time per host. The access of females to honey allowed them to lay more eggs.     

斜纹夜蛾的性选择行为

【目的】为了解斜纹夜蛾Spodoptera litura Fabricius体重、日龄、交配经历及形态特征对其性选择行为的影响。【方法】本实验通过标记成虫后,采用观察记录的方法对其进行探究。【结果】斜纹夜蛾雌虫的体重对雄虫的性选择影响比较明显,体重较大雄虫优先选择体重较小的雌虫(71.43%),而体重较小雄虫喜欢选择体重较大的雌虫(72.00%)。体重较大和体重较小的雄虫都能获得体重较大雌虫的交配选择,但是体重较大者被选择的机会更大(70.00%),体重较小的雄虫不能获得体重较小雌虫的选择。雄虫仅选择1日龄的雌虫,而雌虫偏向选择3日龄和5日龄雄虫。交配经历影响斜纹夜蛾的性选择,未交配的雄虫优先选择未交配的雌虫(86.67%),但未交配的雌虫则优先选择已交配的雄虫(66.67%)。雄虫的形态特征(体长、翅展、腹长、复眼间距和触角长)对雌虫性选择有较明显的影响,但雌虫的形态特征除翅展的大小外,其体长、腹长、复眼间距和触角长等形态特征在雄虫选择进行交配中的作用不大。【结论】体重、日龄、交配经历及形态特征都能不同程度影响斜纹夜蛾的性选择行为。     

Mutual mate choice by mountain pine beetles: size‐dependence but not size‐assortative mating

1. Mutual mate choice may be rare, occurring when both sexes invest heavily in reproduction, mating opportunities are abundant, and individuals differ in quality. 2. Mountain pine beetles, Dendroctonus ponderosae (Curculionidae: Scolytinae) appear to meet the conditions for mutual mate choice. We introduced males to females in breeding sites and observed the occurrence and speed of a male entering a female's gallery. We tested for consequences of mutual mate choice, namely condition‐dependent choosiness and assortative mating. 3. Males were more likely to enter a female's gallery when the gallery was in a smaller tree with less resin production and when the gallery was larger. Female body size and condition did not influence the probability of entry. Larger males were less likely to enter a gallery than were smaller males, probably because of size‐dependent choosiness rather than physical limitations. 4. Small males took longer to enter galleries of large females than of small females, whereas large males entered as quickly into galleries of large females as small females. This suggests size‐dependent choosiness by females. 5. No assortative mating by body size was detected, probably because males appeared to choose on the basis of female‐associated resources rather than on female traits.     

Mating and host density affect host feeding and parasitism in two species of whitefly parasitoids

Abstract The parasitoids in the genera of Encarsia and Eretmocerus (Hymenoptera: Aphelinidae) are important biological control agents of whiteflies, and some of them not only parasitize hosts but also kill them with strong host‐feeding capacity. Two whitefly parasitoid species, Encarsia sophia and Eretmocerus melanoscutus were examined to determine if mating and host density affected their host feeding and parasitism. The whitefly host, Bemisia tabaci, was presented to these two wasp species in densities of 10, 20, 30, 40, 50 and 60 third‐instar nymphs per clip cage. Mated whitefly parasitoid females fed on more hosts than unmated females under a range of host densities (under all six host densities for En. sophia; under the densities of 40 nymphs or more for Er. melanoscutus). Meanwhile, mated females parasitized more whitefly nymphs than unmated females under all host densities for both species. With increase of host density, mated or unmated Er. melanoscutus females killed more hosts by host feeding and parasitism. Mated En. sophia females killed more hosts by host feeding with increase of host density, whereas unmated females did not parasitze whitefly nymphs at all. Our results suggest that only mated female parasitoids with host‐feeding behavior should be released in crop systems to increase their bio‐control efficiency.     

FEMALES ARE CHOOSIER IN THE DARK: ENVIRONMENT‐DEPENDENT RELIANCE ON COURTSHIP COMPONENTS AND ITS IMPACT ON FITNESS

A broad understanding of multimodal courtship function necessitates knowledge of the potential information content of signal components, the efficacy of signal components in eliciting the appropriate receiver response, and the fitness consequences of mating decisions based upon various signal components. We present data addressing each of these requirements for the multimodal‐signaling wolf spider, Schizocosa floridana Bryant. Using diet manipulations, we first demonstrate that both visual and seismic courtship signals are condition‐dependent. Next, using high‐ and low‐quantity diet individuals in mate choice trials across manipulated signaling environments, we demonstrate that the seismic signal is crucial for mating success and further show that female choosiness is environment‐dependent. Females mated more with high diet males only in the absence of visual signals, showing no discrimination in the presence of visual signals. Finally, by quantifying the number of offspring produced by our mated females, we reveal that a female's mating environment, in conjunction with her potential resource availability, influences her fitness—in environments in which females exerted choice, heavier females produced more offspring. Together, this comprehensive set of experiments demonstrates that female choosiness varies across environments, leading to direct fitness consequences.