The influence of female age on male mating preference and reproductive success in cabbage beetle,Colaphellus bowringi

The influence of female age on male mating preference and reproductive success has been studied using a promiscuous cabbage beetle, Colaphellus bowringi Baly （Coleoptera： Chrysomelidae）. In a simultaneous choice test, middle-aged females had significantly greater mating success than young and old females. In single pair trials, when paired with middle-aged virgin males, middle-aged females mated faster, copulated longer, and had greater fecundity and fertility than young or old females, while the longevity of males was not significantly affected by female age. This study on C. bowringi suggests that middle-aged females are more receptive to mating, which can result in the highest male reproductive success.     

Impact of male condition on his spermatophore and consequences for female reproductive performance in the Glanville fritillary butterfly

In butterflies, male reproductive success is highly related to the quality and the size of the spermatophore transferred to the female. The spermatophore is a capsule produced by the male during copulation, which in many species contains sperm in addition to a nuptial gift, and which is digested by the female after copulation. The nuptial gift may contribute to egg production and offspring quality, and in some cases also to female body maintenance. The production of the spermatophore, however, represents a cost for the male and, in polyandrous species, ejaculates are sometimes allocated adaptively across matings. Nonetheless, although the ecological factors affecting the reproductive success of female butterflies have been the topic of numerous studies, little information exists on the factors affecting males’ contribution to reproduction, and the indirect impacts on female fecundity and fitness. We used the Glanville fritillary butterfly, Melitaea cinxia (Linnaeus, 1758) (Nymphalidae), in order to assess variation in male allocation to matings. In this species, smaller males produce smaller spermatophores, but variation in spermatophore size is not correlated with female reproductive success. We show that spermatophore size increases with male age at first mating, decreases with mating frequency and adult food‐deprivation, and is not influenced by developmental food‐limitation. The length of copulation period does not influence the spermatophore size nor influences the polyandrous mating behavior in this species. Male contribution to his spermatophore size is clearly influenced by his condition and adult‐resource at the time of mating. Despite this variation, spermatophore size does not seem to have a direct impact on female reproductive output or mating behavior.     

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.     

Nuptial gifts and sexual selection in photinus fireflies

The phenomenon of nuptial gift transfer during mating occursacross a remarkably wide range of taxa, and such male donationsare likely to influence both pre-copulatory and post-copulatorysexual selection. This paper reviews what is known about nuptialgifts in Photinus fireflies (Coleoptera: Lampyridae), and discussesthe adaptive significance of spermatophores in firefly matingsystems. During copulation Photinus males transfer a spiral,gelatinous spermatophore to the female: sperm are released intothe female's spermatheca for storage, while the remainder ofthe spermatophore disintegrates within a specialized gland.Radiolabelling studies indicate that male-derived protein isused to help provision the female's developing oocytes, andmultiply-mated females show increased fecundity. As most Photinusadults do not feed, these studies suggest that females shouldcontinue to forage for matings to supplement their diminishinglarval reserves, even after they have gained sufficient spermto fertilize their eggs. Male spermatophore mass declines acrosssequential matings, and smaller spermatophores are associatedwith lower paternity success in situations where males competefor fertilizations. Declining spermatophore size across sequentialmatings may thus lead to diminishing reproductive returns forfirefly males. Taken together, these results suggest that seasonalchanges in nuptial gift availability may contribute to reversalsof traditional courtship roles, with male choice and female-femalecompetition occurring as spermatophore availability declines.     

Age at mating and male quality influence female patterns of reproductive investment and survival

The trade‐off between the allocation of resources toward somatic maintenance or reproduction is one of the fundamentals of life history theory and predicts that females invest in offspring at the expense of their longevity or vice versa. Mate quality may also affect life history trade‐offs through mechanisms of sexual conflict; however, few studies have examined the interaction between mate quality and age at first mating in reproductive decisions. Using house crickets (Acheta domesticus), this study examines how survival and reproductive trade‐offs change based on females’ age at first reproduction and exposure to males of varying size. Females were exposed to either a large (presumably high‐quality) or small male at an early (young), middle (intermediate), or advanced (old) age, and longevity and reproductive investment were subsequently tracked. Females mated at a young age had the largest number of eggs but the shortest total lifespans while females mated at older ages produced fewer eggs but had longer total lifespans. The trade‐off between age at first mating and eggs laid appears to be mediated through higher egg‐laying rates and shorter postmating lifespans in females mated later in life. Exposure to small males resulted in shorter lifespans and higher egg‐laying rates for all females indicating that male manipulation of females, presumably through spermatophore contents, varies with male size in this species. Together, these data strongly support a trade‐off between age at first reproduction and lifespan and support the role of sexual conflict in shaping patterns of reproduction.     

Fitness advantage from nuptial gifts in female fireflies

Abstract 1. In many insects, males provide nuptial gifts to females in the form of spermatophores, sperm-containing structures produced by male accessory glands.
2. The work reported here examined the influence of both spermatophore number and spermatophore size on female reproductive output in two related firefly beetles, Photinus ignitus and Ellychnia corrusca (Coleoptera: Lampyridae). Based on differences in adult diet, male spermatophores were predicted to increase female reproductive output to a greater extent in P. ignitus than in E. corrusca .
3. Female fecundity was significantly higher in triply mated females than in singly mated females in both species, with no difference between mating treatments in female lifespan or egg hatching success. No effects of second male spermatophore size on fecundity, lifespan, or egg hatching success were detected in either species.
4. These results suggest a direct fitness advantage from multiple mating for females in both species, although enhanced fecundity may be due either to allocation of spermatophore nutrients to eggs or to other substances transferred within the spermatophore acting as oviposition stimulants.     

Cost of reproduction in males of a satyrine butterfly Lethe diana

Adult butterflies feed largely on floral nectar and tree sap, both of which consist mainly of carbohydrates and include little nitrogen. They depend on the larval diet for nitrogenous resources. Consequently, there is a trade‐off between the reproductive and somatic nitrogenous investments of adults. Furthermore, male butterflies invest a considerable amount into spermatophores, containing nitrogen, which they give to their sexual partners. One way in which male butterflies could potentially replenish their spermatophores is by flight muscle histolysis, which may reduce locomotor ability and lifespan. In the present study, the effect of mating experience on nitrogen dynamics and the lifespan of males is investigated in the satyrine butterfly Lethe diana (Butler). Mated males do not have less thoracic nitrogen than virgin males, suggesting that mating experiences do not induce spermatophore recovery through flight muscle histolysis. Mated males possess less abdominal nitrogen than virgin males at death, indicating that they cannot recover the lost nitrogenous resource used for a single mating. Lifespan does not differ between mated and virgin males. Thus, reproduction and longevity are not fuelled necessarily by the same shared resource pools. Spermatophore mass increases as males get older. Considering that resources for producing spermatophores are limited, males may adjust the spermatophore mass strategically, depending on their chances of future reproduction.     

Male age: effect on mating behaviour and success in the cockroach Diploptera punctata

Older cockroach, Diploptera punctata, males (28 days old) transferred more sperm and larger spermatophores to females than 8-or 14-day-old males. In one male: one female behavioural tests, no significant correlations were found between either how rapidly males initiated courtship with females (response time), or intensity or duration of courtship, and the number of sperm or size of spermatophore subsequently transferred. Eight-day males were slower to initiate courtship than 28-day males. There was no significant differences in the duration of courtship for 8-, 14- and 28-day males, even though the mean duration of courtship was more than twice, as long for 8-day males compared with 14- and 28-day males. In competitive mating tests with one female and three males, one of each age, 28-day males were significantly more successful than 8- and 14-day males. It is concluded that the greater success of 28-day males is due to differences in male behaviour rather than to female discrimination.     

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

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

The role of male age, sperm age and mating history on fecundity and fertilization success in the hide beetle

Models of age-related mate choice predict female preference for older males as they have proven survival ability. However, these models rarely address differences in sperm age and male mating history when evaluating the potential benefits to females from older partners. We used a novel experimental design to assess simultaneously the relative importance of these three parameters in the hide beetle, Dermestes maculatus. In a two-part experiment we first explored age-related male mating success and subsequently examined the consequences of male age, sperm age and male mating history on female fecundity and fertilization success. In a competitive mating environment, intermediate-age males gained significantly higher mating success than younger or older males. To test the consequences for females of aged-related male mating success, a second set of females were mated to males varying in age (young, intermediate-age and old), in numbers of matings and in timing of the most recent mating. We found that male age had a significant impact on female fecundity and fertilization success. Females mated to intermediate-age males laid more eggs and attained consistently higher levels of fertilization success than females with young and old mates. A male's previous mating history determined his current reproductive effort; virgin males spent longer in copula than males with prior mating opportunities. However, differences in copulation duration did not translate into increased fecundity or fertilization success. There was also little evidence to suggest that fertilization success was dependent on the age of a male's sperm. The experiment highlights the potential direct benefits accrued by females through mating with particular aged males. Such benefits are largely ignored by traditional viability models of age-related male mating success.     

Sex differences in the effects of juvenile and adult diet on age‐dependent reproductive effort

Sexual selection should cause sex differences in patterns of resource allocation. When current and future reproductive effort trade off, variation in resource acquisition might further cause sex differences in age‐dependent investment, or in sensitivity to changes in resource availability over time. However, the nature and prevalence of sex differences in age‐dependent investment remain unclear. We manipulated resource acquisition at juvenile and adult stages in decorated crickets, Gryllodes sigillatus, and assessed effects on sex‐specific allocation to age‐dependent reproductive effort (calling in males, fecundity in females) and longevity. We predicted that the resource and time demands of egg production would result in relatively consistent female strategies across treatments, whereas male investment should depend sharply on diet. Contrary to expectations, female age‐dependent reproductive effort diverged substantially across treatments, with resource‐limited females showing much lower and later investment in reproduction; the highest fecundity was associated with intermediate lifespans. In contrast, long‐lived males always signalled more than short‐lived males, and male age‐dependent reproductive effort did not depend on diet. We found consistently positive covariance between male reproductive effort and lifespan, whereas diet altered this covariance in females, revealing sex differences in the benefits of allocation to longevity. Our results support sex‐specific selection on allocation patterns, but also suggest a simpler alternative: males may use social feedback to make allocation decisions and preferentially store resources as energetic reserves in its absence. Increased calling effort with age therefore could be caused by gradual resource accumulation, heightened mortality risk over time, and a lack of feedback from available mates.     

Seasonal Variation in Mate Choice of Photinus ignitus Fireflies

Mate choice by either sex may vary with changes in the associated costs and benefits, determined by factors such as the availability of potential mates and variation in mate quality. We examined seasonal variation in operational sex ratio, courtship behavior, spermatophore mass, egg count, and the relationship between morphological traits and mating success in Photinus ignitus fireflies to determine if mate choice in either sex varied with the availability and relative reproductive investment of fertilizable females and sexually active males. Successfully mating males had larger lanterns than unsuccessful males when the operational sex ratio was male‐biased. In addition, female responsiveness to male signals increased as the number of courting males decreased, and male spermatophore mass decreased with body size across the mating season. Successfully mating females had larger body mass than unsuccessful females. Female body mass predicted egg count and female rejection by males increased as the season progressed and female size decreased. These results suggest that both male and female P. ignitus exhibit mate choice, and that such choice is influenced by seasonal variation in the abundance and quality of potential mates.     

Old‐male paternity advantage is a function of accumulating sperm and last‐male precedence in a butterfly

Old‐male mating advantage has been convincingly demonstrated in Bicyclus anynana butterflies. This intriguing pattern may be explained by two alternative hypotheses: (i) an increased aggressiveness and persistence of older males during courtship, being caused by the older males' low residual reproductive value; and (ii) an active preference of females towards older males what reflects a good genes hypothesis. Against this background, we here investigate postcopulatory sexual selection by double‐mating Bicyclus anynana females to older and younger males, thus allowing for sperm competition and cryptic mate choice, and by genotyping the resulting offspring. Virgin females were mated with a younger virgin (2–3 days old) and afterwards an older virgin male (12–13 days old) or vice versa. Older males had a higher paternity success than younger ones, but only when being the second (=last) mating partner, while paternity success was equal among older and younger males when older males were the first mating partner. Older males produced larger spermatophores with much higher numbers of fertile sperm than younger males. Thus, we found no evidence for cryptic female mate choice. Rather, the findings reported here seem to result from a combination of last‐male precedence and the number of sperm transferred upon mating, both increasing paternity success.     

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.     

Costs of Reproduction in Male Bicyclus anynana and Pieris napi Butterflies: Effects of Mating History and Food Limitation

Life history theory predicts trade‐offs arising from the costs of reproduction: individuals investing more into reproduction should have less energy available for their own somatic demands. Despite a wealth of research on such costs for females, very few studies have looked at the male side. We investigated the costs of reproduction in males of two butterfly species, Bicyclus anynana and Pieris napi. The males of these species differ tremendously in their allocation of nutrients to reproduction: virgin male spermatophores were 18.5 times larger in the latter compared with those in the former. Based on this striking difference, we expected to find a longevity cost of mating in P. napi, but not necessarily in B. anynana. In line with our predictions, we found no evidence for a survival cost of mating in B. anynana, while there was some evidence for such a cost in P. napi. Here, virgin males lived longer than once‐ or twice‐mated ones. However, this overall trend did not hold for all treatment groups, and there was no linear relationship between the number of copulations and longevity. Adult food limitation reduced longevity throughout, but had otherwise very little impact. Spermatophore dry mass decreased in later compared with first spermatophores, while water content increased, indicating that spermatophore production is indeed physiologically costly. Additionally, adult food limitation tended to reduce spermatophore fresh and dry mass, and tended to increase spermatophore water content. Based on our results we tentatively conclude that longevity and reproduction may not always be traded off against each other, as in our study organisms’ longevity seems to depend in first place on adult carbohydrate income, while male reproduction seems to be primarily (although not entirely) fuelled by larval storage reserves.     

Determinants and long‐term costs of early reproduction in males of a long‐lived polygynous mammal

In long‐lived polygynous species, male reproductive success is often monopolized by a few mature dominant individuals. Young males are generally too small to be dominant and may employ alternative tactics; however, little is known about the determinants of reproductive success for young males. Understanding the causes and consequences of variability in early reproductive success may be crucial to assess the strength of sexual selection and possible long‐term trade‐offs among life‐history traits. Selective pressures driven by fluctuating environmental conditions may depend on age class. We evaluated the determinants of reproduction in male bighorn sheep (Ovis canadensis) aged 2–4 years using 30 years of individual‐level data. These young males cannot defend estrous ewes and use alternative mating tactics. We also investigated how the age of first detected reproduction was correlated to lifetime reproductive success and longevity. We found that reproductive success of males aged 3 years was positively correlated to body mass, to the proportion of males aged 2–4 years in the competitor pool, and to the number of females available per adult male. These results suggest that reproductive success depends on both competitive ability and population age–sex structure. None of these variables, however, had significant effects on the reproductive success of males aged 2 or 4 years. Known reproduction before the age of five increased lifetime reproductive success but decreased longevity, suggesting a long‐term survival cost of early reproduction. Our analyses reveal that both individual‐level phenotypic and population‐level demographic variables influence reproductive success by young males and provide a rare assessment of fitness trade‐offs in wild polygynous males.     

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.     

Condition-dependent spermatophore size is correlated with male's age in a bushcricket (Orthoptera: Phaneropteridae)

During mating, male bushcrickets transfer a spermatophore that consists of a sperm-containing ampulla and a product of the accessory glands, the spermatophylax, which females directly ingest. In the present study, we demonstrate male spermatophore allocation in the bushcricket Poecilimon zimmeri . Males of this species show condition-dependent spermatophore investment. This investment depended upon the age at first mating of males, with older individuals transferring larger spermatophores than younger ones of the same body mass. Independently of age, heavier males transfer larger spermatophores, but the size of males (as measured by femur length) was not a good predictor. Heavier males allocate a lower proportion of their mass to spermatophores and reach their maximal investment point earlier than less heavy males. Spermatophylax production levelled off to a species specific maximum earlier than that of sperm investment (measured as ampulla mass), suggesting that males face high levels of sperm competition.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 354–360.     

Effects of diapause on post‐diapause reproductive investment in the moth Ostrinia scapulalis

Diapause is a strategy used by many insect species to survive adverse environmental conditions. However, diapause incurs costs that may have adverse effects on post‐diapause development and reproduction. We herein investigated the effects of diapause on the post‐diapause reproductive investment of males and females in a multivoltine moth, the adzuki bean borer, Ostrinia scapulalis (Walker) (Lepidoptera: Crambidae). We found that (1) post‐diapause males and females were smaller and had lower mating success than non‐diapause individuals, (2) post‐diapause females had lower fecundity and shorter longevity than non‐diapause females, (3) post‐diapause males transferred similar numbers of eupyrene and apyrene sperm as non‐diapause males, (4) the fecundity and longevity of non‐diapause females mated with post‐diapause males and those mated with non‐diapause males were not significantly different, and (5) no significant relationship was found between diapause duration (short and long) and post‐diapause reproductive investments in both males and females. These results suggest that post‐diapause males did not reduce reproductive investment in spite of the cost of diapause, and the significant decline in reproductive output in post‐diapause females was due to their reduced body weight and longevity, which appeared to be direct consequences of the cost of diapause.     

Factors affecting differential mating success in male crested newts, Triturus cristatus

This study investigates differences in mating success in male crested newts, Triturus cristatus. In experiments, mating success varies considerably between individual males. Male size affects mating success, but the maximal height of the male's crest is more important. Male size and crest height are correlated, but in partial correlation analysis only crest height is significantly correlated with mating success. Motivation is also an important factor in determining male mating success as males that had been isolated for a week were more successful than freshly caught males. Newts might be limited in their ability to produce spermatophores, and isolation gives the male time to produce more spermatophores. It is likely that motivational state is less important in a natural population than size and crest height, because wild males are often interrupted during courtship before spermatophore deposition and hence are highly motivated to mate.