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.     

Methoprene application and diet protein supplementation to male melon fly,Bactrocera cucurbitae,modifies female remating behavior

Methoprene (an analogue of juvenile hormone) application and feeding on a protein diet is known to enhance male melon fly, Bactrocera cucurbitae Coquillett (Diptera: Tephritidae), mating success. In this study, we investigated the effect of these treatments on male B. cucurbitae's ability to inhibit female remating. While 14‐d‐old females were fed on protein diet, 6‐d‐old males were exposed to one of the following treatments: (i) topical application of methoprene and fed on a protein diet; (ii) no methoprene but fed on a protein diet; (iii) methoprene and sugar‐fed only; and (iv) sugar‐fed, 14‐d‐old males acted as controls. Treatments had no effect on a male's ability to depress the female remating receptivity in comparison to the control. Females mated with protein‐deprived males showed higher remating receptivity than females first mated with protein‐fed males. Methoprene and protein diet interaction had a positive effect on male mating success during the first and second mating of females. Significantly more females first mated with sugar‐fed males remated with protein‐fed males and females first mated with methoprene treated and protein‐fed males were more likely to remate with similarly treated males. Females mating latency (time to start mating) was significantly shorter with protein‐fed males, and mating duration was significantly longer with protein‐fed males compared with protein‐deprived males. These results are discussed in the context of methoprene and/or dietary protein as prerelease treatment of sterile males in area‐wide control of melon fly integrating the sterile insect technique (SIT).     

Remating Behavior of Cnephasia jactatana Walker Females (Lepidoptera: Tortricidae)

Cnephasia jactatana Walker is an important pest of kiwifruit in New Zealand. We investigated, under laboratory conditions, the effects of multiple mating on the reproductive performance of C. jactatana females and how such effects varied with male virginity and larval nutrition. We found that in permanent pairs, remating increased female fecundity and fertility but suboptimally fed females benefited more from remating. Regardless of this benefit, mass-reared pairs had a lower remating frequency. Females remating with a virgin male or a male that had delivered a spermatophore presented similar fecundity and fertility; however, females receiving a second ejaculate from a virgin male had increased daily fecundity. Female weight clearly affected remating behavior since those that received a second ejaculate were significantly heavier. Neither mating length nor size of the first spermatophore influenced female remating. Further, mass-reared and individually reared males produced spermatophores of similar size.     

Sperm competition and the significance of female multiple mating in the predatory miteParasitus fimetorum

Females of the predatory miteParasitus fimetorum (Gamasida; Parasitina) inhabiting animal manure indiscriminately copulate with many mates. The sperm competition between the males was estimated by electrophoresis of allozymes and the effects of multiple mating on female reproduction were investigated. When females were forced to mate only once, their fecundity decreased drastically compared to the case of multiple mating (but longevity was unaffected). When one female mated with two males, the outcome of sperm competition depended greatly upon the mating interval. When the second mating occurred immediately after the first, the female fecundity decreased as in the case of single mating and the second male fertilized only a few eggs. However, when there was an interval of 1 day between the two matings, the females achieved normal fecundity and the second male fertilized approximately half the eggs. This suggests that the spermatophore deposited by the first male may act as a short-term copulatory ‘plug’ in the female's genital opening. When one female mated with several males with 1 day intervals, three or more males shared fertilization of the eggs. This study suggests that the multiple mating of females is a necessary stimulus to continue oogenesis and some physiological factors for this stimulation may exist in spermatophores.     

Determinants of paternity in a butterfly

Success in sperm competition is of fundamental importance to males, yet little is known about what factors determine paternity. Theory predicts that males producing high sperm numbers have an advantage in sperm competition. Large spermatophore size (the sperm containing package) also correlates with paternity in some species, but the relative importance of spermatophore size and sperm numbers has remained unexplored. Males of the small white butterfly, Pieris rapae (Lepidoptera: Pieridae), produce large nutritious spermatophores on their first mating. On their second mating, spermatophores are only about half the size of the first, but with almost twice the sperm number. We manipulated male mating history to examine the effect of spermatophore size and sperm numbers on male fertilization success. Overall, paternity shows either first male or, more frequently, second male sperm precedence. Previously mated males have significantly higher fertilization success in competition with males mating for the first time, strongly suggesting that high sperm number is advantageous in sperm competition. Male size also affects paternity with relatively larger males having higher fertilization success. This may indicate that spermatophore size influences paternity, because in virgin males spermatophore size correlates with male size. The paternity of an individual male is also inversely correlated with the mass of his spermatophore remains dissected out of the female. This suggests that females may influence paternity by affecting the rate of spermatophore drainage. Although the possibility of female postcopulatory choice remains to be explored, these results clearly show that males maximize their fertilization success by increasing the number of sperm in their second mating.     

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   

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.     

Transfer and fate of male secretions deposited in the spermatophore of females of Acanthoscelides obtectus Say (Coleoptera Bruchidae)

During mating, males of Acanthoscelides obtectus deposit a spermatophore in the female genital tract. Spermatophore structure subsequently undergoes considerable modification, especially the central portion, which becomes vacuolated. Two methods were used to show that certain male secretions could thus pass into female haemolymph. When young males were injected with [¹⁴C]-arginine or [¹⁴C]-histidine, the accessory glands actively incorporated the isotope and the resulting spermatophores were radioactive. After mating, spermatophore radioactivity declined and then appeared in the haemolymph of females and in the oöcytes after a delay. Immunoelectrophoresis also showed that antigens appeared in the haemolymph of females after mating which reacted against male-gland antiserum. This technique, however, did not enable us to detect the presence of male antigens in the oöcytes formed after mating. The fate of some male secretions in the female and their physiological importance in the control of the female reproductive function were analysed in the present work.     

Extreme Costs of Mating for Male Two-Spot Ladybird Beetles

Male costs of mating are now thought to be widespread. The two-spot ladybird beetle (Adalia bipunctata) has been the focus of many studies of mating and sexual selection, yet the costs of mating for males are unknown. The mating system of A. bipunctata involves a spermatophore nuptial gift ingested by females after copulation. In this study, we investigate the cost to males of mating and of transferring spermatophores in terms of lifespan, ejaculate production and depletion of nutritional reserves. We found that males faced a strong trade-off between mating and survival, with males that were randomly assigned to mate a single time experiencing a 53% reduction in post-mating lifespan compared to non-mating males. This is among the most severe survival costs of a single mating yet reported. However, spermatophore transfer did not impact male survival. Instead, the costs associated with spermatophores appeared as a reduced ability to transfer spermatophores in successive matings. Furthermore, males ingested more food following spermatophore transfer than after matings without spermatophores, suggesting that spermatophore transfer depletes male nutritional reserves. This is to our knowledge the first report of an effect of variation in copulatory behaviour on male foraging behaviour. Overall, our study highlights the advantages of assessing mating costs using multiple currencies, and suggests that male A. bipunctata should exhibit mate choice.     

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.     

Female alpine newts (Triturus alpestris) mate initially with males signalling fertility benefits

Gametic asymmetry implies that females invest more per gamete than males do and thus sperm is considered to be a relatively cheap resource. However, contrary to this classic view, sperm has been shown to be frequently in short supply; hence, selection favouring females that mate for fertility benefits should occur. For this reason, we determined whether males signalling fertility are preferred by female newts of the species Triturus alpestris . We performed paired female–male trials using unmated and previously inseminated females to determine potential criteria for female interest in a courting male, to establish what factors lead to successful mating and to assess the importance of female choice for direct and indirect benefits. We found that female interest in any potential mate and mating success decreased once mating had occurred. Furthermore, we detected an increase in spermatophore deposition rate and rapid spermatophore transfer in encounters that resulted in a successful mating. The results obtained indicate that female alpine newts are attracted to males showing signs of relatively high fertility and that females exhibit a decreased propensity to mate once initial sperm reserves have been acquired. Our results support the theory of initial female choice for fertility benefits.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 483–491.     

Postcopulatory female choice increases the fertilization success of novel males in the field cricket, Gryllus vocalis

Although recent studies have demonstrated that female crickets prefer novel males to previous mates, the relative contribution of pre- and postcopulatory behaviors to this advantage remain unknown, as do the reproductive consequences to males. I paired females either with previous or novel mates, and recorded the latency to mating and the time after mating at which the female removed the male's spermatophore, terminating sperm transfer. Females that mated with familiar males removed their spermatophores sooner than females that mated with novel males. Females paired with novel males also mated more quickly than females paired with familiar males, but this difference was not statistically significant. A molecular-based paternity analysis was used to determine whether the postcopulatory preference of females for novel males influences a male's fertilization success. Females were assigned to either mate three times with the same male and then once with a novel male, or four times with four different males. The paternity of the last male was higher when the female previously had mated repeatedly with the same male than when she had mated previously with different males. These results suggest that female spermatophore removal behavior influences male paternity such that novel males receive a fertility benefit.     

Limits to Nuptial Gift Production by Male Fireflies, Photinus ignitus

Males of diverse insect species provide females with nuptial gifts, and limits on males' ability to produce these gifts may influence courtship behavior and mating systems. In the firefly Photinus ignitus, males transfer a complex spermatophore to females during mating. We provided firefly males unlimited access to responsive females to examine whether spermatophore production limits male mating success. Male spermatophore mass decreased significantly across sequential matings, and the percentage of successful matings declined during the second half of each male's life span. Male body mass explained a significant proportion of variation in size of the first spermatophore produced by P. ignitus males, but this relationship disappeared with second spermatophores. This study indicates that males' ability to produce spermatophores declines over their lifetime and that limits on nuptial giftproduction can constrain male mating success in Photinus fireflies.     

Behavioural shift from courtship solicitation to mate avoidance in female ringlet butterflies (Aphantopus hyperanthus) after copulation

In butterflies where nutrients contributed by males through mating are used by females for egg production and/or somatic maintenance, females may benefit from mating more than once. However, in species where sperm is used only for the fertilization of eggs and the sperm received in one copulation is sufficient for fertilization of all of the eggs, females should benefit from mating only once. In these species the reproductive success of females is likely to be proportional to the time they can allocate to egg-laying activities. Thus these females should be selected to minimize the time spent in an unmated condition, and to minimize time-consuming interactions, like courtship, with males after mating. As shown by spermatophore counts, females of the ringlet butterfly, Aphantopus hyperanthus, generally mate only once. These females exhibit behaviour consonant with the above view: they solicit courtship before copulation and actively avoid males after mating.     

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.     

Repeated matings offset costs of reproduction in female crickets

Summary Courtship food gifts can be a significant source of nutrition to females and costly for males to produce; hence, costs of reproduction should be reduced for multiple-mating females and increased for multiplemating males in a gift-giving species. We tested this hypothesis by experimentally manipulating mating opportunities of males and females of two cricket species,Gryllodes sigillatus, a gift-giving species andGryllus veletis, a non-gift-giving species. Females of both species consume the externally attached spermatophore after mating, but inG. sigillatus, the sperm-containing ampulla is accompanied by a large gelatinous spermatophylax. In both species, survival of mated females given limited access to males was reduced relative to virgin females, thus suggesting a cost of reproduction to females. However, females given unlimited mating opportunities lived as long as virgins and also produced significantly more offspring than limited-access females. These results suggest that benefits of repeated matings, particularly those arising through spermatophore consumption, offset costs of reproduction in females. Lack of a treatment by species interaction suggests that females of both species derive nutritional benefits through spermatophore consumption, and that any additional advantage to the consumption of the spermatophylax inG. sigillatus is offset by more frequent mating byG. veletis females. In contrast to females, varying mating opportunities had no effect on male survival, suggesting that mating effort is not very costly to males. Male survival increased linearly with body mass but only when males were food-deprived, suggesting that larger males possess greater initial energy reserves to sustain their longevity when food-stressed.     

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.     

Breeding in the crayfish, Austropotamobius pallipes: mating patterns, mate choice and intermale competition

1. The breeding behaviour of the white-clawed crayfish, Austropotamobius pallipes, which is in decline throughout Europe, was analysed. 2. Observations and experiments focused on: (i) several aspects of mating behaviour, showing the primary role of the female during courtship, the patterns of mature male receptivity and the possible existence of a mating stimulus produced by either a mating pair or a receptive female; (ii) the potential for mate choice either by males (males were not choosy, mating with the first receptive female they met) or by females (that rejected the smallest males and males with only one cheliped) and (iii) intermale fighting before copulation (larger males copulated more often, with the exception of some ’sneakers‘). 3. Sperm competition occurred when a new male was presented with a female after a previous male‘s spermatophore had been deposited; the new male cleaned the rival male‘s spermatophore by feeding on it before copulation. This behaviour has also been observed in some pseudoscorpions, millipedes and collembolans, but always in cases where the spermatophore is deposited on the substratum. Introduction     

Effect of Adult Feeding on Male Mating Behaviour in the Butterfly, <Emphasis Type="Italic">Bicyclus anynana</Emphasis> (Lepidoptera: Nymphalidae)

Many species of lepidopterans supplement their nectar diet with foods rich in nitrogen and minerals, which are present only in trace amounts in nectar. We examined the effect of adult diet on mating behaviour and spermatophore characteristics in male Bicyclus anynana (Butler, 1879) butterflies, which feed on rotten fruits as adults. We found little effect of adult diet on male reproduction in terms of mating rate and sperm production, although males fed on fruit produced larger spermatophores on their first mating compared to males fed sugar only. We also examined how males allocate sperm across matings. Males ejaculate larger spermatophores during their first mating, and produce spermatophores containing decreasingly fewer non-fertile sperm with number of matings performed. Males that produced more non-fertile sperm on their first mating had reduced lifespan possibly indicating a trade-off between sperm production and adult longevity. It is suggested that adult diet has little affect on male ejaculate production and males feed on fruit to supplement their energetic carbon requirements.     

Sperm competition and the significance of female multiple mating in the predatory mite Parasitus fimetorum

Females of the predatory mite Parasitus fimetorum (Gamasida; Parasitina) inhabiting animal manure indiscriminately copulate with many mates. The sperm competition between the males was estimated by electrophoresis of allozymes and the effects of multiple mating on female reproduction were investigated. When females were forced to mate only once, their fecundity decreased drastically compared to the case of multiple mating (but longevity was unaffected). When one female mated with two males, the outcome of sperm competition depended greatly upon the mating interval. When the second mating occurred immediately after the first, the female fecundity decreased as in the case of single mating and the second male fertilized only a few eggs. However, when there was an interval of 1 day between the two matings, the females achieved normal fecundity and the second male fertilized approximately half the eggs. This suggests that the spermatophore deposited by the first male may act as a short-term copulatory plug in the female's genital opening. When one female mated with several males with 1 day intervals, three or more males shared fertilization of the eggs. This study suggests that the multiple mating of females is a necessary stimulus to continue oogenesis and some physiological factors for this stimulation may exist in spermatophores.