Virgin male mating advantage in a primitive acoustic insect (Orthoptera: Haglidae)

Male reproductive behavior in the relict flightless haglid, Cyphoderris strepitans,entails the generation of sound signals and the provision of nuptial gifts to mates. These food gifts take two forms: (1) a gelatinous mass (spermatophylax) augmenting the spermatophore and (2) fleshy metathoracic wings adapted to be eaten. The female consumes a portion of the male's underwings during courtship and copulation and the spermatophylax afterward. The incidence of wing-feeding wounds can be used to monitor the mating success of field-caught males. If, when a male mates, he compromises his ability to provide subsequent nutritive gifts, females would benefit by mating with virgin rather than nonvirgin males. To test this, mating success of virgin and nonvirgin males was compared in a field population. Virgins were found to obtain more matings than explained by their numbers relative to nonvirgins in the population. We conclude that, having mated, a male is at a disadvantage, relative to his virgin competitors, in securing further matings.     

Female mate choice and mating costs in the polyandrous butterflyPieris napi (Lepidoptera: Pieridae)

Males of the green-veined white butterfly (Pieris napi L.) transfer large ejaculates that represent on average 15% of their body mass when mating for a first time. Shortly after mating a male is able to transfer only a small ejaculate when mating a second time. Male ejaculate production plays a crucial role in the mating system ofP. napi because females use male-derived nutrients for egg production and somatic maintenance. Here we study how timing of female rematings and copulation duration are influenced by the mating history of their mates and, also, study if females exert mate choice to minimize their mating costs. Mating with a recently mated male increased female mating costs by increasing time in copula and mating frequency. Virgin females that mated with virgin males remated after an average of 6 days, whereas virgin females that mated with recently mated males remated after an average of 2 days. Moreover, copulations involving recently mated males lasted on average almost 7 h, whereas copulations involving virgin males lasted on average 2 h. Recently mated males were eager to remate, in spite of the fact that the size of the ejaculate they transfer is small and that they remain in copula for a long time. Hence it seems that males are more successful in the sexual conflict over mating decisions and that females do not minimize mating costs by choosing to mate preferentially with virgin males.     

Adaptive female choice for middle-aged mates in a lekking sandfly

Most theoretical models of age-related mate choice predict that females should prefer older males because they have proven survival ability. An alternative view is that older males represent inferior mates because of negative genetic correlations between early and late fitness components, or because older males have traded off longevity against other fitness components, have accumulated deleterious germ-line mutations, or are less well adapted to current conditions than more recently born individuals. While numerous studies have reported female choice for older males, few have explicitly examined the fitness consequences of such a preference. We present evidence from a lekking sandfly, Lutzomyia longipalpis, showing that choosy females discriminate against older males and gain a fitness benefit from their choice. When permitted free choice from an aggregation consisting of males aged zero to two days (young), four to six days (middle-aged) and eight to ten days (old), females preferentially mated with middle-aged males, but all measures of female reproductive success were independent of male age. In contrast, when a second set of females was randomly assigned single virgin males of known age, the eggs of those paired to old mates exhibited lower hatching success than the eggs of females mated to young or middle-aged males. These results suggest that females avoid mating with older males because they represent poorer quality mates. Age-related differences in male quality may have a genetic basis, but could equally well arise through a phenotypic decline in sperm quality or sperm transfer ability with male age. The lack of evidence of female discrimination against older males from other studies may be because these did not explore the reproductive success of the full age range of males.     

Reduction in the attraction of males to females after mating in the rice leaf bug Trigonotylus caelestialium

The influence of mating on the extent to which males are attracted to females in Trigonotylus caelestialium (Heteroptera: Miridae) was examined. No differences in attraction of males to mated and virgin females were observed within 3–5 h of mating, but males became less attracted to females 1 to 2 days after the first mating. The difference in male attraction to mated vs virgin females disappeared at 4 days after mating. These results indicate that reduced attraction of males to mated females occurs after a certain time interval, and persists for a few days. Furthermore, males were less attracted to females that had mated with virgin vs recently mated males, i.e. males that had just mated with another female at 1 and 2 days after mating. The ejaculate expenditure of recently mated males was less than that of virgin males. Hence, the amount of male ejaculate transferred to females during mating, rather than the act of mating, might influence the attraction of males to females. The results demonstrate that mating reduces the attraction of males to females in T. caelestialium on the basis of direct observation of male behavior.     

Virgin females compete for mates in the male lekking species Ceratitis capitata

Abstract. Aggressive behaviour occurring in intrasexual competition is an important trait for animal fitness. Although female intrasexual aggression is reported in several insect species, little is known about female competition and aggressive interactions in polygynous male lekking species. The interactions of female Mediterranean fruit flies, Ceratitis capitata (a male lekking species), with other females and mating pairs under laboratory conditions are investigated. Mature, unmated (virgin) females are aggressive against each other and against mating pairs, whereas immature females are not. Female aggression against other females decreases dramatically after mating; however, mated females maintain aggression against mating pairs. In addition, higher intrasexual aggression rates are observed for mature, virgin females than for virgin males of the same age. The results show that female aggressiveness is virginity related, suggesting female competition for mates. These findings have important implications for understanding the physiological aspects of a complex social behaviour such as aggression and should stimulate further research on female agonistic behaviour in male lekking mating systems.     

Age,Experience and Sex – Do Female Bulb Mites Prefer Young Mating Partners?

In species where advancing sire age is associated with decreased progeny fitness, female resistance to mating with old partners can be expected to evolve. In polyandrous species, such resistance may be contingent on female mating experience: virgins should be relatively indiscriminate to ensure egg fertility, whereas non‐virgins can be expected to base their re‐mating decisions on the age of their previous versus potential new partners, and ‘trade‐up’ if previously mated with old males. Here, we tested these predictions using a promiscuous and relatively long‐living bulb mite (Rhizoglyphus robini), in which old sire age is associated with decreased fecundity of daughters. In a fully factorial design, we applied two male treatments, young and old, and three female treatments, virgin, previously mated to an old male and previously mated to a young male. Consistent with earlier studies, we observed a reduced mating success of old males. However, we found no support for attributing this result to female discrimination, as female behavior in response to male mounting attempts was not affected by the age of the suitor, or by its interaction with the age of the female’s previous mate. Interestingly, females were passive during 93% of male mounting attempts observed, suggesting that once they are located by a male, they exert little control over copulation. Old males had lower mate‐searching activity and were less efficient in obtaining matings (lower success rate per mounting attempt), suggesting a decreased mate‐securing ability because of aging. Overall, our results suggest that in bulb mites, male ability to secure mates declines with age, whereas they do not support the prediction that females actively discriminate against old partners.     

Mate Choice by Non-Virgin Females Contributes to Reproductive Isolation between Populations of the Harlequin Beetle-Riding Pseudoscorpion

Determining whether reproductive isolation evolves through mate choice and/or gametic factors that prevent fertilization or through the post‐zygotic mechanisms of hybrid sterility or inviability is fundamental to understanding speciation. Investigation of the pre‐ and post‐zygotic components of reproductive isolation is facilitated in the pseudoscorpion, Cordylochernes scorpioides, by its indirect method of sperm transfer and viviparous embryonic development. Previous research on this species, in which mate discrimination was assessed in virgin females, suggested that female choice played only a minor role in reproductive isolation between populations from French Guiana and Panamá. Here, in a study of three allopatric populations of C. scorpioides from Panamá, we assessed mating‐stage isolation in both virgin and once‐mated females, and found that female discrimination depends critically on mating status. Virgin females were almost invariably receptive, showing no tendency to discriminate against males from allopatric populations. By contrast, non‐virgin females were significantly more likely to reject foreign males than males from their own population. Male sexual motivation could not account for differences in either female sexual receptivity or male success in sperm transfer. Allopatric and sympatric males did not differ in number of spermatophores deposited as either a female’s first or second mate. Nonetheless, allopatric males achieved significantly lower sperm transfer success not only with choosy, non‐virgin females but also with virgin females. Given the lack of behavioral discrimination by virgin females, female receptivity was not the only factor influencing differences in sperm transfer success. Multivariate analysis of spermatophore morphology suggests that the higher failure rate of matings between allopatric males and virgin females resulted from population differences in sperm packet architecture. Overall, our findings indicate that assessment of discrimination against allopatric males that is limited to virgin females may seriously underestimate the contribution of female mate choice to reproductive isolation between populations.     

Reproductive strategies of the larval parasitoid Microplitis croceipes

Mate choice may have important consequences for offspring sex ratio and fitness of haplodiploid insects. Mate preference of females of the solitary larval parasitoid Microplitis croceipes (Cresson) (Hymenoptera: Braconidae) for virgin and mated males, and vice versa, and the reproductive consequences (i.e., the sex ratio expressed as the proportion of male offspring) were examined in choice and non‐choice experiments. In addition, the effect of repeated rapid and daily copulation of an individual male on the sex ratio of offspring of the female mates was assessed. Males preferred virgins over mated females, whereas females copulated with a male irrespective of his mating status. In both the rapid and daily copulation assay, females copulating with a male that had copulated five times or more produced a higher sex ratio than females that had copulated with a virgin male. Females that copulated with virgin males once or twice produced a significantly and considerably lower sex ratio than females that first copulated with a sperm‐depleted male followed by a virgin male. This indicates that copulating with a sperm‐depleted male has costs and limits acquisition by the female of sperm from virgin males.     

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.     

Chemosensory cues allow male Tenebrio molitor beetles to assess the reproductive status of potential mates

Males of many insect species, including beetles, choose their mates according to their reproductive status. However, the ways in which male beetles evaluate female reproductive status have received little attention. We tested the existence of male mate choice in the mealworm beetle, Tenebrio molitor, by observing mating and courtship behaviour of males given simultaneous access to pairs of females differing in their reproductive status: (1) mature versus immature; (2) virgin versus previously mated; (3) familiar (mated with the experimental male) versus unfamiliar (mated with a different male). Males courted and mated preferentially with mature and virgin females. To determine whether chemical cues played a role in these discriminations, we exposed males to filter paper squares bearing chemical cues from different types of females: (1) virgin versus mated; (2) mature versus immature. Males were significantly more attracted to those squares bearing chemical cues from virgin and mature females, suggesting that males can assess female reproductive status on the basis of chemical cues alone.     

Male Mating Success in the Presence of a Conspecific Opponent in a Madagascar Hissing Cockroach,Gromphadorhina portentosa (Dictyoptera: Blaberidae)

The influence of a conspecific competitor on male mating behavior was examined in a Madagascar hissing cockroach, Gromphadorhina portentosa. Previous studies have suggested that both male-male competition and female discrimination during courtship interactions may influence male mating success. Familiar pairs of males with a known social association were placed in an arena with a single virgin female and observed. As expected, subordinate males mated significantly less often than their dominant opponents. In pairs in which one male mated, dominant individuals limited the access of subordinates to females. Dominant males displayed an increased frequency and duration of interaction with the female. However, in pairs where both males remained unmated, the mating behavior of dominant and subordinate males did not differ significantly. As interactions progressed, as in the case of males that remained unmated, subordinate males gained increased access to the female. Mated males tended to be larger than their opponent although within a rank, males that mated were no larger than those that remained unmated. These results are discussed in light of the possible roles of male-male competition and female discrimination during courtship interactions.     

Sexual History Affects Mating Behavior and Mate Choice in the Crayfish Orconectes limosus

Because mating entails both costs and potential benefits to both sexes, males and females should be under selection to make optimal choices from among available potential mates. For example, in some cases, individuals may benefit by using information on potential mates' previous sexual histories to make mate choices. In such cases, the form and direction of these benefits may vary both between the sexes and based on the sexual history of the choosing individuals themselves. We investigated the effects of recent previous sexual history on the mate choice and mating behavior of both males and females of the crayfish Orconectes limosus. In one experiment, we found that opposite‐sex dyads comprising crayfish that had both mated 7–8 d previously with other conspecifics were significantly less likely to mate than dyads in which at least one crayfish was unmated. In a second experiment, we found that, when presented with a choice of tethered (but free to move) opposite‐sex conspecifics, only virgin females discriminated between males based on sexual history, showing a preference for virgin males over recently mated males. Mated females, mated males, and virgin males showed no preferences based on the sexual histories of potential mates. We discuss the implications of these inferences in the context of what was previously known about mating behavior and potential sperm limitation in crustaceans and other taxa.     

Effects of delayed mating and male mating history on the reproductive potential of Leucoptera coffeella (Lepidoptera: Lyonetiidae)

Abstract 1 Despite the importance of Leucoptera coffeella (Guérin‐Mèneville) in coffee production worldwide, there is a lack of information on its reproduction. This knowledge will help in mass rearing, and support the development of behavioural control techniques for this insect. The present study determined the effects of delayed mating and previous matings of male L. coffeella on fecundity, egg viability and frequency of female remating. 2 The highest levels of fecundity and egg viability were obtained from matings of 1–3‐day‐old females. When females mated at 5 days of age, there were reductions of 40% in oviposition and of 43% in egg viability. 3 Females mated with 2‐day‐old virgin males were more fecund than those mated with older males; egg viability was also low (18%) from females mated with older males. 4 Virgin females that mated with virgin males laid a greater number of eggs than those mated with previously copulated males. Egg viability decreased with the increase in the number of previous male matings. 5 Five‐day‐old females remated in greater proportion than 2–3‐day‐old females. Females that copulated with males that had previously mated three times had higher rates of remating than those that copulated with virgin males. 6 The results obtained indicate that 1–3 days after emergence is the optimum age for mating. The implications of these findings for the control of L. coffeella by synthetic sex pheromone are discussed.     

Mating strategy in Aleochara bilineata: sperm storage and allocation

Abstract By contrast to females that can maximize reproductive success with only one or a few copulations, males generally increase their fitness with frequency of mating. Sperm storage and allocation is therefore crucial for both male and female fitness. Sperm storage in Aleochara bilineata (Coleoptera; Staphylinidae) is investigated by measuring the number of spermatozoa stored in the female spermatheca after single, double or triple successive copulations with different males. The potential advantages of polyandry are studied in terms of the number of sperm stored by females mated twice with the same male (i.e. repeated copulation), compared with females mated twice with two different virgin males (i.e. polyandry). Level of polygyny is also estimated by measuring sperm allocation when ten successive mates are offered to a virgin male. Aleochara bilineata females store the sperm of the same or different males additively, suggesting no advantage for polyandry in terms of the number of sperm stored. A virgin male is able to inseminate ten different females but the number of sperm transferred decreases linearly. Finally, the latencies and durations of copulations are measured in all experiments to estimate changes according to the male or female status (i.e. virgin or mated). The latency before mating is higher when females are virgin than when females have already mated.     

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.     

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.     

Context‐ and State‐Dependent Male Mate Choice in a Sexually Cannibalistic Spider

High male mating investment may favor selection on male mate choice particularly if females vary in quality. Terminal investment strategies constitute a maximal mating effort and have evolved independently in the absence of paternal investment in several spider taxa including the genus Argiope. To test for male mate preferences in the above context, we used the sexually cannibalistic spider A. bruennichi. We varied male state (mating status and post‐maturation age) as well as the competitive context and quantified male mate choice decisions between females of different states and developmental stages in binary choice tests. We found an overall adaptive preference for the virgin against the mated female regardless of male mating state. Furthermore, we demonstrated that older males paid more attention to female fecundity‐related traits than to mating status. In a second set of experiments, we offered males a choice between a virgin and a subadult female and varied the competitive context which had no effect on male decisions. Curiously, a preference for the virgin adult female was only apparent after exclusion of females that matured <3 d prior to the test. Repeated tests of males supported the hypothesis that males do not distinguish between a freshly matured virgin female and a subadult female. Our results show that male spiders execute mate choice based on information collected from female silk strands and that they integrate their own state into mating decisions.     

Copulation patterns in the golden orb-web spider Nephila madagascariensis

A consequence of multiple mating by females can be that the sperm of two or more males directly compete for the fertilisation of ova inside the female reproductive tract. Selection through sperm-competition favours males that protect their sperm against that of rivals and strategically allocate their sperm, e.g., according to the mating status of the female and the morphology of the spermatheca. In the majority of spiders, we encounter the otherwise unusual situation that females possess two independent insemination ducts, both ending in their own sperm storage organ, the spermatheca. Males have paired mating organs, but generally can only fill one spermatheca at a time. We investigated whether males of the African golden orb-web spider Nephila madagascariensis can prevent rival males from mating into the same spermatheca and whether the mating status of the female and/or the spermatheca causes differences in male mating behaviour. There was no significant difference in the duration of copulations into unused spermathecae of virgin and mated females. We found that copulations into previously inseminated spermathecae were generally possible, but shorter than copulations into the unused side of mated females or with virgins. Thus, male N. madagascariensis may have an advantage when they mate with virgins, but cannot prevent future males from mating. However, in rare instances, parts of the male genitals can completely obstruct a female genital opening.     

Silk‐ and volatile‐based male mate choice in the genital plug‐producing spider

Female mating history can have a strong effect on male fertilization success. Although males often prefer to mate with virgin females, they often also engage with mated females. As the intensity of sperm competition can differ among mated females, males are expected to evolve means to identify their status. In spiders, males often use female silk to gather information about female quality. Males of many spider species deposit mating plugs into female genitalia to hinder further copulations. We tested whether males of the foliage‐dwelling, plug‐producing spider Philodromus cespitum, which is an important natural enemy of pests, discriminate between females of different mating status and whether they can determine the extent of genital plugging in mated females solely on the basis of cues gained from deposited female silk. We presented males with draglines of females that varied in either mating status (virgin vs. mated), the extent of plugging (small vs. big plug), or the age of the plug (fresh vs. old plug) and examined their mate preferences. Additionally, we tested whether males were attracted to volatile cues produced by female bodies. Our experiments revealed that males preferred draglines of virgin females to those of mated females, and mated females with small plugs to those with large plugs. They were also attracted to female volatile cues. This study suggests that males are able to extract fine‐scale information on mating status from female draglines.     

Male and Female Mate-Choice Decisions by Evarcha culicivora, An East African Jumping Spider

There has been a considerable recent interest in the criteria by which animals choose mates and in the extent to which mating systems tend to be based on mutual mate choice. In this study, we consider Evarcha culicivora, a salticid spider from East Africa. This species has some unusual characteristics, including active display by females as well as males, males that kill females more frequently than females kill males and wide intrasexual variation in body size. For females, larger males are especially dangerous. Here, we demonstrate, using two experimental designs (live‐mate choice and mount choice), that virgin males, virgin females and previously mated males prefer larger opposite‐sex individuals as potential mates, but mated females prefer smaller, safer males as potential mates.