Lifetime mating success of males in a natural population of the papilionid butterfly,Atrophaneura alcinous (Lepidoptera: Papilionidae)

Lifetime mating success of males in a natural population of the papilionid butterfly, Atrophaneura alcinous, was investigated and causes of the variation were examined. The most successful males mated with 5 females, whereas about 73% of the males failed to mate. Body size of males was not correlated with their eclosion date, longevity and lifetime mating success. There was no trade-off between mating success and longevity, and long-lived males had a disproportionately high mating success. Although number of available females per male per day was not variable among males with different longevities, long-lived males had higher mating efficiency. Time interval between matings by non-virgin males was shorter than that from eclosion to the first mating. High lifetime mating success of long-lived males was strongly related to their mating experience, not to their age per se.     

SIZE,SYMMETRY, AND SEXUAL SELECTION IN THE HOUSEFLY,MUSCA DOMESTICA

Relationships between measures of body size, asymmetry, courtship effort, and mating success were investigated in the housefly, Musca domestica (Diptera: Muscidae). A previous study indicated that both male and female flies with low fluctuating asymmetry enjoyed enhanced mating success. The aim of our investigations was to determine whether the greater success of symmetrical males is due to variation in male mating effort or to female choice and whether males exhibited mate choice. However, our study found directional rather than fluctuating asymmetry with both male and female flies having, on average, longer left wings than right. Also, asymmetry was not related to mating success in either sex. Rather, both males and females appeared to exhibit choice on the basis of the size of potential mates, with males preferring females with long bodies and females preferring heavy males. Possible benefits from choice of large mates are discussed. The initial mating strikes (in which the male leaps onto the back of the female) did not appear to be targeted according to female morphology, and their frequency did not vary according to male morphology. This indicates that mate choice by both sexes according to size probably occurs during the later stages of courtship, when the flies are in intimate contact. Possible reasons for the absence of choice according to asymmetry are discussed.     

Fluctuating Asymmetry and Male Mating Success in a Sphragis-Bearing Butterfly Luehdorfia japonica (Lepidoptera: Papilionidae)

The relationship between fluctuating asymmetry (FA) and mating success was studied within males of the sphragis-baring butterfly Luehdorfia japonica, which were collected at various periods during their mating season. FA was measured on the forewing and hindwing radius lengths of male butterflies. Mating frequency of males was estimated by assessing the degree of scale loss from their claspers. Males consume scales and use them to form sphragis on the female abdomen during copulation, sealing the ostium bursa for life. Age of males was scored as wing age 0 to 4 according to the wearing of the wing. FA was negatively correlated with mating frequency but positively correlated with wing age, and average FA decreased with mating season. As females have little chance to express mate choice, it is likely that FA is an indicator of male viability: symmetrical males live longer and/or fly more actively, resulting in a higher lifetime mating success compared to asymmetrical males.     

Male morphology,performance and female mate choice of a swarming insect

1. Females of many species select mates based on important morphological and performance traits. This behaviour likely facilitates reproductive success thus exhibiting sexual selection. Most such studies have evaluated a single morphological variable, and only a minority of them studies the effects of behaviour and performance (functional capacity: Irschick et al., Evolutionary Ecology Research, 10 , 177–196, 2008) at all.
2. This study compares male morphological and performance traits differing in three variables that may correlate with fitness: condition index, flight ascent speed and resistance to torpor in relation to male mating success in the parasitoid wasp species Alabagrus texanus (Braconidae), a species where males swarm about emerging virgin females that display both choosy and non-choosy behavioural phenotypes.
3. Males that successfully mated with choosy females exhibited higher condition indices and somewhat stronger resistance to torpor than other males. Conversely, males that mated with non-choosy females did not differ in any trait from other males we measured. Early-arriving males did not have higher condition indices or greater mating success than other males.
4. Thus, both morphological and performance traits contributed to male success, which acted through female choice, indicating a role for sexual selection. Patterns of choice further differed among females, independent of male traits.

     

Variation in energy reserves and role of body size in the mating system of Anopheles gambiae

Anopheles gambiae mates in flight. Males gather at stationary places at sunset and compete for incoming females. Factors that account for male mating success are not known but are critical for the future of any genetic control strategy. The current study explored variations in nutritional reserves (sugars, glycogen, lipids, and proteins) in wild‐caught swarming and resting males and evaluated the effect of body size and wing symmetry on male mating success. Our results showed that glycogen and sugar reserves are mobilized for flight. Males consume proportionally 5.9‐fold as much energy derived from sugars in swarming activities than when they are at rest. Mated males were on average bigger than unmated ones (P<0.0001). A strong correlation between the left and right wings in both mated and unmated males was found and additional analysis on fluctuating asymmetry did not show any indication of mated males being more symmetrical than unmated ones. The distribution of wing size of mated males was focused around a central value, suggesting that intermediate size of males is advantageous in the An. gambiae mating system. The results are discussed in the context of sexual selection.     

Greater Mating Success of Drosophila biarmipes Males Possessing an Apical Dark Black Wing Patch

Drosophila biarmipes males possess an apical black patch on their wings. In a laboratory stock of this species, males without such a patch have been found. Mating success of these two types of male was studied in an Elcns-Wattiaux mating chamber. Results show that males possessing the dark black patch have greater mating success than those without. This finding suggests that visual stimulus plays an important role in the mating behaviour of D. biarmipes.     

Delayed mating and reproduction in the autumn gum moth Mnesampela privata

• 1 The success of mating disruption using synthetic sex pheromones depends not only on preventing mating, but also on delaying mating in the target insect. Using the geometrid pest of Eucalyptus plantations, Mnesampela privata (Guenée), we determined the effect of delaying mating when imposed on males only, females only or on both sexes simultaneously, for 1, 3, 5 and 7 days.
• 2 Delayed mating had a significant negative impact on reproduction, with a 0.89‐fold decrease in the likelihood of mating and a 0.67‐fold decrease in the likelihood of that mating resulting in fertile eggs for every day that mating is delayed. A mating delay of 7 days reduced the mean number of viable eggs laid to 4–13% of that laid by moths paired immediately after emergence.
• 3 Male only imposed mating delays had a significantly lower effect on reducing the likelihood of pairs mating than when both sexes were delayed. A delay imposed on one sex only or on both sexes simultaneously, however, had a similar negative impact on the proportion of fertile matings as well as on the total number of fertile eggs laid.
• 4 Longevity of mated female and male M. privata was significantly different between mating delay treatments, with a significant decline in female longevity when they mated with older males.
• 5 The underlying mechanisms causing a decline in female reproductive output when a mating delay was imposed on males versus females are discussed in relation to the reproductive biology of M. privata and the potential of using mating disruption strategies to control populations in Eucalyptus plantations.

     

Substrate influences mating success and transmission of courtship vibrations for the parasitoid Cotesia marginiventris

The influences of artificial and natural rearing substrates on mating success were investigated for the parasitoid wasp Cotesia marginiventris (Cresson) (Hymenoptera: Braconidae), a candidate for augmentative biological control of various lepidopteran pests. Five rearing substrates were tested: plastic, glass, chiffon fabric, and leaves of two host plants, bean [Vigna unguiculata (L.) (Fabaceae)] and maize [Zea mays L. (Poaceae)]. Mating success was highest on chiffon, lowest on plastic and glass, and intermediate on maize and bean. The transmission characteristics of one component (buzz 1) of the courtship vibrations produced by male wing fanning were investigated using laser vibrometry. The duration of buzz 1 was longer on maize, bean, and chiffon than on plastic and glass. The fundamental frequency of buzz 1 (~300 Hz) was lowest on bean and highest on glass, and intermediate among other substrates. The relative amplitude of buzz 1 was higher on chiffon than on plastic, glass, or bean, and intermediate on maize. The relative importance of airborne sound and substrate vibration as courtship signals was also investigated with experiments that manipulated the production of courtship vibrations and the mating substrates. The amplitude of courtship vibrations on chiffon was significantly higher for winged males than for dealated males. The mating success of males was impacted by both the presence of wings and the mating substrate. These findings suggest that mating success and transmission of courtship vibrations are influenced by the rearing substrate, and that courtship vibrations are critical to mating success in C. marginiventris. Future efforts to mass rear this parasitoid and other insects should consider the potential influences of rearing substrates on mating.     

Multiple mating and its effect on the reproductive success of female Epiphyas postvittana (Lepidoptera: Tortricidae)

Abstract.

• 1 Multiple mating and its effect on reproductive performance of female Epiphyas postvittana (Walker) moths were studied under controlled conditions.
• 2 The age at which the moths mated for the first time ranged from the first to the tenth day after emergence, but 71% of first matings were during the first 3 days.
• 3 The majority (63%) of females had one or two spermatophores in the bursa copulatrix. Some (24%) were found with three to five spermatophores, whereas no successful mating occurred among 13% of individuals. The number of matings was partly dependent on the number of mates available to the female. Between the range of sex ratios of one male to one female and four males to one female maximal mating success occurred at the ratio of three males to one female.
• 4 Virgin females were capable of egg-laying, but mating stimulated and accelerated oviposition. Mated individuals laid twice as many eggs as unmated ones.
• 5 The level of copulatory activity did not influence the longevity of females irrespective of the number of males available to them.
• 6 Sex ratios with greater than one male to a female improved the reproductive success by marginally increasing fecundity and fertility.
• 7 It is concluded that multiple mating would enhance population growth, and is of particular benefit to populations with a preponderance of females, as is known to occur naturally in this species.

     

The contribution of multiple mating and spermatophore consumption to the lifetime reproductive success of female field crickets (Gryllus bimaculatus)

ABSTRACT.

• 1 The adaptive significance of multiple mating by female Gryllus bimaculatus (De Geer) was investigated.
• 2 Multiple mating prevented the depletion of sperm stores and, therefore, maintained high hatching success. This may not, however, explain the high frequency of remating in this species.
• 3 Male-derived egg stimulants known to be passed with sperm at mating increased the number of eggs produced only when females mated throughout their lifespans.
• 4 Spermatophore consumption appeared to provide nutrients which, while they did not increase the quantity of eggs, increased egg quality as indicated by weight. Females who consumed spermatophores had a greater hatching success.
• 5 While females may derive non-genetic benefits from mating, these are apparently long-term benefits; females must mate throughout their lives in order to accrue them.
• 6 Since the benefits of mating may not be derived from individual males, the spermatophores and their contents in this species are best considered as mating effort.

     

Fat and sassy: factors underlying male mating success in a butterfly

Theory predicts that male mating success depends on resource holding potential (RHP), which is reflected by proxies of condition, such as body mass, fat content, strength, or weaponry. In species lacking any physical means to inflict injuries upon combatants, such as butterflies, the factors determining mating success are less clear. Against this background, we explored the determinants of male mating success in the tropical butterfly Bicyclus anynana Butler (Lepidoptera: Nymphalidae: Satyrinae), by comparing physiological, immunological, and morphological traits between winning and losing males. Our results showed that successful males are characterized by a better flight performance, evidenced by having longer wings, a heavier thorax, a lighter abdomen, a higher fat content, and higher phenoloxidase expression levels than their unsuccessful counterparts, when being compared after their first mating. Males that won three consecutive trials against the same combatant were also characterized by a better flight performance, having larger forewings, a higher body mass, and a higher fat content. Thus, successful males were larger and in better condition than unsuccessful ones. Strikingly, many differences found indicated an enhanced flight performance for the former, which we suggest ultimately plays the key role for male mating success in B. anynana. As fat is the main energy source for flying insects, being crucial to flight endurance and in turn presumably to male mating success, it may represent a key determinant at the proximate level.     

Intergenerational paternal effect of adult density in Drosophila melanogaster

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Male mating success,sexual size dimorphism,and site fidelity in two species of Malacoctenus (Labrisomidae)

Synopsis In both Malacoctenus hubbsi and Malacoctenus macropus, males defended preferred oviposition sites from both other males and potential egg predators. In M. hubbsi, adult females were larger than adult males. Larger M. hubbsi males were not associated with territory parameters that were correlated with higher mating success, and male size was not correlated with mating success. Male size did affect mating success when territory parameters were statistically controlled for, but the failure of large males to associate with better territories eliminated any mating advantage for larger males. In M. macropus, males are larger than females. Larger males defended preferred oviposition sites, and had higher mating success than did smaller males. Male M. macropus also had much higher site fidelity than male M. hubbsi. These results suggest that the evolution of the differences in site fidelity and sexual size dimorphism between these two species may be due to sexual selection acting differentially in these two species.     

Scramble competition polygyny in the milkweed leaf beetle: combat, mobility, and the importance of being there

I examined the scramble competition mating system of the milkweedleaf beetle, Labidomera clivicollis(Coleoptera: Chrysomelidae),using data on lifetime mating success of two breeding cohortsin Austin, Texas, USA (30° N, 97° W) and one in Bridgeport,New York, USA (43° N, 76° W). Data from six daily censuseswere combined with focal animal samples to examine the importanceof combat between males and to evaluate the extent to whichthe characters elytron length, vagility, and emergence datewere correlated with male mating success. In both populations,emergence date was the only consistent determinant of the numberof mates a male obtained during his lifetime. Males emergingearlier in the season had higher mating success because theywere active for more of the breeding season. Male body size(elytron length) was not correlated with mating efficiency orlifetime mating success in either population. Males formed prolongedmating associations with females that lasted for up to 2.5 daysand paired males in the New York population had a strong advantageover interlopers attempting to mate with the female. Only 10%of takeover attempts were successful. Selection favored vagilemales in New York, where the sex ratio was male biased, butnot in Texas, where it was female biased. Males that were morevagile had higher mating efficiencies in New York but did nothave higher lifetime mating success because of a trade-off betweenmating efficiency and survival. Behavioral data demonstratedthat lone males spent more time walking and less time feedingthan lone females; mating males were unable to feed at all.The survival disadvantage that comes with increased vagilitymay be due to loss of time spent feeding. The characters examinedsuggest that the most important thing a male L. clivicolliscan do to increase his lifetime mating success is to be presentfor as much of the breeding season as possible.     

Beyond magic traits: Multimodal mating cues in Heliconius butterflies

Species coexistence involves the evolution of reproductive barriers opposing gene flow. Heliconius butterflies display colorful patterns affecting mate choice and survival through warning signaling and mimicry. These patterns are called “magic traits” for speciation because divergent natural selection may promote mimicry shifts in pattern whose role as mating cue facilitates reproductive isolation. By contrast, between comimetic species, natural selection promotes pattern convergence. We addressed whether visual convergence interferes with reproductive isolation by testing for sexual isolation between two closely related species with similar patterns, H. timareta thelxinoe and H. melpomene amaryllis. Experiments with models confirmed visual attraction based on wing phenotype, leading to indiscriminate approach. Nevertheless, mate choice experiments showed assortative mating. Monitoring male behavior toward live females revealed asymmetry in male preference, H. melpomene males courting both species equally while H. timareta males strongly preferred conspecifics. Experiments with hybrid males suggested an important genetic component for such asymmetry. Behavioral observations support a key role for short‐distance cues in determining male choice in H. timareta. Scents extracts from wings and genitalia revealed interspecific divergence in chemical signatures, and hybrid female scent composition was significantly associated with courtship intensity by H. timareta males, providing candidate chemical mating cues involved in sexual isolation.     

Genetic evidence for the mating system and reproductive success of black sea bream (Acanthopagrus schlegelii)

Understanding the mating system and reproductive success of a species provides evidence for sexual selection. We examined the mating system and the reproductive success of captive adult black sea bream (Acanthopagrus schlegelii), using parentage assignment based on two microsatellites multiplex PCR systems, with 91.5% accuracy in a mixed family (29 sires, 25 dams, and 200 offspring). Based on the parentage result, we found that 93.1% of males and 100% of females participated in reproduction. A total of 79% of males and 92% of females mated with multiple partners (only 1 sire and 1 dam were monogamous), indicating that polygynandry best described the genetic mating system of black sea bream. For males, maximizing the reproductive success by multiple mating was accorded with the sexual selection theory while the material benefits hypothesis may contribute to explain the multiple mating for females. For both sexes, there was a significant correlation between mating success and reproductive success and the variance in reproductive success of males was higher than females. Variation in mating success is the greatest determinant to variation in reproductive success when the relationship is strongly positive. The opportunity for sexual selection of males was twice that of females, as well as the higher slope of the Bateman curve in males suggested that the intensity of intrasexual selection of males was higher than females. Thus, male–male competition would lead to the greater variation of mating success for males, which caused greater variation in reproductive success in males. The effective population number of breeders (N_b) was 33, and the N_b/N ratio was 0.61, slightly higher than the general ratio in polygynandrous fish populations which possibly because most individuals mated and had offspring with a low variance. The relatively high N_b contributes to the maintenance of genetic diversity in farmed black sea bream populations.     

The mating strategy and reproductive performance of Agriphila aeneociliella (Lepidoptera: Crambidae), a new insect pest of wheat in China

1. The Eastern Grass‐veneer Agriphila aeneociliella (Eversmann) (Lepidoptera: Crambidae) is a serious stem‐feeding pest of wheat crops that has become established in China in recent years. A better understanding of the mating strategy and reproductive performance of A. aeneociliella could improve integrated pest management programmes against this newly established species by disturbing its behaviour and reducing its reproduction potential. Based on ethological and reproductive biological approaches, the mating and reproductive performances of A. aeneociliella were investigated.
2. Unlike the common nocturnal lepidopterans, the mating rhythm of A. aeneociliella moths showed a marked diurnal pattern. The female courtship rhythm and the mating rhythm reached peaks within the first 2 h after the onset of photocycle.
3. The mating success rate of monogamous pairs was 55.6%, whereas the male‐biased sex ratio (2♂:1♀) increased the mating rate (72.2%) and the female‐biased ratio (1♂:3♀) led to the lowest mating rate (27.8%).
4. Both females and males were able to mate twice. The duration of copulation decreased substantially with male mating frequency, whereas, when a previously mated female was paired with a virgin male, fecundity significantly increased. Monogamous couples who mated only once in their lives have the highest hatchability (97.13 ± 0.49%).

     

SEXUAL SELECTION,VIABILITY SELECTION,AND DEVELOPMENTAL STABILITY IN THE DOMESTIC FLY MUSCA DOMESTICA

Associations between developmental stability, sexual selection, and viability selection were studied in the domestic fly Musca domestica (Diptera, Muscidae). Developmental stability of the wings and tibia of flies of both sexes, measured in terms of their level of fluctuating asymmetry, was positively associated with mating success in free ranging populations and in sexual selection experiments. Mated individuals may have obtained indirect fitness benefits from sexual selection of two different kinds. First, the entomopathogenic fungus Enthomophthora muscae (Zygomycetes, Entomophthorales) infects and kills adult domestic flies, and flies dead from fungus infections had more asymmetric wings than flies dead for other reasons. Experimental deposition of fungus spores on uninfected flies demonstrated that flies with asymmetric wings were more susceptible to fungus infections than flies with symmetric wings. Second, domestic flies were frequently eaten by insectivorous barn swallows Hirundo rustica, and flies depredated by birds had more asymmetric wings and tibia than surviving flies.     

Male size,mating potential and lifetime reproductive success in the field cricket,Gryllus bimaculatus (De Geer)

Male size is known to influence short-term, competitive mating success in the field cricket, Gryllus bimaculatus. Short-term measures of mating success are, however, potentially misleading when considering the influence of sexual selection on male morphology and behaviour. The lifetime mating success of males was investigated in the absence of competition to determine the effects of male size per se on reproductive potential and any effects of female behaviour on male lifetime mating success. The cost of spermatophore production was relatively greater for small males who consequently had a longer refractory period between matings, thus limiting their reproductive potential. Large males had a mating advantage resulting from a differential response of females to the courtship attempts of large and small males. Furthermore, small males were ‘sexually incompetent’ in terms of their ability to attach spermatophores once mounted. There was, therefore, an overall positive relationship between male size and lifetime reproductive success.     

Coping with the cold: temperature and mating activity of male sagebrush crickets Cyphoderris strepitans (Orthoptera: Haglidae)

Abstract. The effect of temperature on male sexual displays is well documented but its role in mediating the differential mating success of males is less well established. Male sagebrush crickets Cyphoderris strepitans Morris and Gwynne occur in high‐elevation sagebrush meadows in mountainous areas of the western U.S.A., and often are observed singing at temperatures as low ?8 to ?4 °C to attract sexually receptive females. Males exhibit differential mating success based on their previous mating experience: virgin males have a higher probability of obtaining a mating than do non‐virgin males of securing an additional mating. This differential mating success is mediated in part by an unusual form of nuptial feeding behaviour. Females feed on males’ hind wings at mating and the costs imposed on males as a consequence of this behaviour constrain male mating success. In the present study, to determine whether sagebrush crickets are capable of some degree of thermoregulation, and also to investigate whether this ability is influenced by male mating experience, thoracic temperatures of males of varying mating status were measured in relation to ambient temperature throughout the breeding season. A full factorial model examining the effect of site of capture, mating status, body mass and ambient temperature reveals that only ambient temperature has a significant effect on thoracic temperature after controlling for all other factors. It is concluded that differences in the calling times of virgin and non‐virgin males and their ability to secure mates does not result from differences in their ability to cope physiologically with low temperatures.