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.     

Female Gryllus vocalis Field Crickets Gain Diminishing Returns from Increasing Numbers of Matings

Although female insects generally gain reproductive benefits from mating frequently, females do not mate unlimited numbers of times. This study asks whether the limit on female mating rate is imposed by trade‐offs between reproduction and survival. Female Gryllus vocalis were given the opportunity to mate 5, 10, or 15 times with novel males, and the effects on daily fecundity (egg production), fertility (proportion of eggs that were fertilized), and female post‐experimental longevity were measured. Females that mated 10 times laid more eggs and had a higher proportion of fertile eggs than females that mated 5 times. However, females that mated 15 times did not lay significantly more eggs or have a higher proportion of fertile eggs than females that mated 10 times. Although number of matings did not affect the date that females laid their last egg, mating more times was associated with a prolonged period of laying fertile eggs. Number of matings did not affect female post‐experimental longevity. Thus, there was no trade‐off between female reproductive effort and survival, even when females mated very large numbers of times. When females were allowed to mate ad libitum, the average number of times that females mated was greater than the number of times that confers maximal fitness. The lack of cost to mating explains why females might be willing to mate beyond the point of diminishing reproductive returns.     

The effect of adult feeding on the fecundity,weight loss and survival of the pine beauty moth,Panolis flammea (D&S)

Summary Adult females of the pine beauty moth, Panolis flammea, fed on a saturated honey solution, laid significantly more eggs than those individuals given water only or not fed at all.At 20°, the longevity of both male and female moths was significantly increased when they were supplied with a food source. At 10° fed male months were not significantly longer lived than unfed moths. Fed female moths at 10° were however, significantly longer lived than unfed individuals. At 20°, fed male moths lived longer than fed females (8.17 days as compared with 5.95 days). At 10°, fed female moths lived longer than fed males (16.00 days compared with 11.41 days).At 20°, weight loss in unfed females over the reproductive period was proportional to the number of eggs laid and the survival time of the individual. In fed females, no significant weight loss occurred over the reproductive period, irrespective of the number of eggs laid or the longevity of the individual.Egg production, even in fed females, over the life span of an individual was only a fifth of the total eggs present in the reproductive tract at eclosion.It is concluded that adult females of P. flammea are likely to require an extra external food source to achieve their full reproductive potential.     

Life history of breeding partners alters age‐related changes of reproductive traits in a natural population of blue tits

Reproductive senescence, an intra‐individual decline in reproductive function with age, is widespread, but proximate factors determining its rate remain largely unknown. Most studies of reproductive senescence focus on females, leaving senescence in male function and its implications for female function largely understudied. We constructed linear mixed models to explore the interactive effects of paternal and maternal age and a life‐history trait (i.e. age at first reproduction) on four fitness components (i.e. laying date, clutch size, number of fledglings and number of recruits) measured in a wild, breeding population of blue tits Cyanistes caeruleus ogliastrae where individual breeding success has been followed for over 30 years (our dataset spanned 29 years). Previous studies have shown that, across female lifespan, laying date decreases and subsequently increases; earlier laying dates result in higher fitness because hatchlings have greater access to a seasonal food source. Our analyses reveal that females that initiate reproduction early in life show a greater delay in laying date with old age. In addition to delayed laying dates, older females lay smaller clutches. However, the magnitude of female age effects was influenced by the age at first reproduction of their breeding partners. Senescence of laying date and clutch size was reduced when females mated with males that reproduced early in life compared to males that delayed reproduction. We confirmed that both laying date and clutch size were significantly correlated with reproductive fitness suggesting that these dynamics early in the breeding cycle can have long‐term consequences. These complex phenotypic interactions shed light on the proximate mechanisms underlying reproductive senescence in nature and highlight the potential importance of cross‐sex age by life‐history interactions.     

Effects of age at mating of both sexes on female longevity and fecundity performance in Ophraella communa LeSage (Coleoptera: Chrysomelidae)

Life-history strategies of animals are governed by fitness-related trait trade-offs. In particular, the age at which either one of the two sexes copulate has been shown to affect insect reproductive output and longevity. This suggests that trade-offs between longevity and reproduction might drive the choice of a mating partner based on their age and ultimately such choices might select for different life-histories. Although several studies indicate that female age at mating dictates subsequent insect longevity, fecundity, and egg hatch rate, it is unclear how male age at copulation affects these life-history traits. In this study, we simultaneously investigated the effects of female and male age on female fecundity, eggs hatch rate, and adult longevity in Ophraella communa LeSage (Coleoptera: Chrysomellidae), the primary biological control agent of the invasive common ragweed, Ambrosia artemisiifolia L. We found that young mature females lived significantly longer when they were mated with older males. Maximum female fecundity and subsequent egg hatch rate occurred when young females mated with 3-day-old males. On the other hand, females lived longer when mating with an older male. These findings are in accordance with the cost-of-reproduction concept.     

Geographical variation in reproductive ageing patterns and life‐history strategy of a short‐lived passerine bird

We investigated differences in ageing patterns in three measures of breeding performance in populations of barn swallows Hirundo rustica L. from Spain and Denmark differing in breeding latitude and hence migration distance and duration of the breeding season. We found differences in ageing patterns between populations. Generally, young (i.e. yearling) and old females (i.e. ≥ 5 years of age) laid their first eggs later and produced smaller clutches than middle‐aged females (i.e. 2–4 years of age) in both populations. The southernmost population (i.e. Spanish) showing the shorter migratory distance experienced a greater within‐individual increase in timing of breeding and clutch size in early life and a greater within‐individual decrease in laying date but not in clutch size during senescence compared with the northernmost population (i.e. Danish). We also found that the number of fledglings produced annually was related to the age of the two members of the breeding pairs with pairs composed of young and old females performing less well than breeding pairs composed of middle‐aged females. We did not find reproductive senescence for the age of the male while controlling for the age of the female on the number of fledglings produced annually by the breeding pair. Differential survival between individuals did not explain age effects on laying date or annual clutch size in neither population. However, the increase in the number of fledglings produced annually with age was partly explained by the disappearance of poor‐quality members of the pairs, mainly poor‐quality males. Age‐related breeding success (i.e. number of fledglings) was similar for barn swallows from Spain and Denmark. Therefore, the study of ageing patterns and life‐history strategies in free‐ranging animals from more than a single population can throw new light on life‐history theory, population dynamics and evolutionary studies of senescence.     

Influence of age at mating on the reproductive performance of Parthenium beetle,Zygogramma bicolorata (Coleoptera: Chrysomelidae)

Abstract Age-specific mating incidence, sexual maturation and effect of age at mating on reproductive performance of the Parthenium beetle, Zygogramma bicolorata Pallister, was studied. Based on 50% mating incidence the calculated age of sexual maturation of males and females was 10.5 and 11.1 days, respectively, which was not statistically significant. However, on the basis of age at first mating, that is, sexual maturity, females matured 2 days earlier than males. Fecundity, pre-oviposition, oviposition and post-oviposition period and female longevity appear to be influenced by female age at mating with reproductive performance peaking at 30 days. On the other hand, egg viability was influenced by male age and was highest when males mated at the age of 40 days. To summarise, egg production and timing of egg deposition was female age-dependent, whereas egg fertility was male age-dependent. It was also observed that females mated at a later age and laid a higher number of eggs immediately after mating than did earlier mated females. This was ostensibly in a bid to increase fitness by maximizing reproductive output in the reduced life span available. This is the first investigation on the effect of age of females at mating on reproduction in this beetle.     

Reproductive trade‐offs in a long‐lived bird species: condition‐dependent reproductive allocation maintains female survival and offspring quality

Life history theory is an essential framework to understand the evolution of reproductive allocation. It predicts that individuals of long‐lived species favour their own survival over current reproduction, leading individuals to refrain from reproducing under harsh conditions. Here we test this prediction in a long‐lived bird species, the Siberian jay Perisoreus infaustus. Long‐term data revealed that females rarely refrain from breeding, but lay smaller clutches in unfavourable years. Neither offspring body size, female survival nor offspring survival until the next year was influenced by annual condition, habitat quality, clutch size, female age or female phenotype. Given that many nests failed due to nest predation, the variance in the number of fledglings was higher than the variance in the number of eggs and female survival. An experimental challenge with a novel pathogen before egg laying largely replicated these patterns in two consecutive years with contrasting conditions. Challenged females refrained from breeding only in the unfavourable year, but no downstream effects were found in either year. Taken together, these findings demonstrate that condition‐dependent reproductive allocation may serve to maintain female survival and offspring quality, supporting patterns found in long‐lived mammals. We discuss avenues to develop life history theory concerning strategies to offset reproductive costs.     

Effect of multiple mating on the reproductive performance of the rice leaffolder moth,Cnaphalocrocis medinalis (Lepidoptera: Crambidae)

To clarify whether multiple mating of females and males affects the reproductive performance of the rice leaffolder moth, Cnaphalocrocis medinalis (Guenée), we examined the effect of the number of matings (once, twice, or three times) for females (female treatment) and males (male treatment) on the incidence of moth mating, number of eggs laid, egg hatchability, and adult longevity. We also compared the effect of multiple mating imposed on males or females separately with the effect of that imposed on both sexes simultaneously (both sexes treatment). The incidence of mating of females and males that mated three times (3-mated females and males) was significantly lower than for females and males that mated twice or once (2-mated or 1-mated females and males). The incidence of mating of 1-mated moths (both sexes) was significantly higher than for 2-mated or 3-mated moths (both sexes). Two-mated or 3-mated females laid significantly more eggs with significantly higher hatchability than 1-mated females. Females that mated with 1-mated males (second male mating) or 2-mated males (third male mating) laid significantly fewer eggs than those that mated with virgin males (first male mating). Females laid significantly more eggs after the second and third matings for moths of both sexes than after the first mating for moths of both sexes. The mechanisms of improvement and decline of female reproductive performance when multiple mating was imposed on males or females are also discussed in relation to the reproductive biology of C. medinalis.     

Geographical variation in reproductive traits and trade‐offs between size and number of eggs in the king ratsnake,Elaphe carinata

We collected gravid king ratsnakes (Elaphe carinata) from three geographically separated populations in Chenzhou (CZ), Lishui (LS) and Dinghai (DH) of China to study the geographical variation in female reproductive traits and trade‐offs between the size and number of eggs. Not all reproductive traits varied among the three populations. Of the traits examined, five (egg‐laying date, post‐oviposition body mass, clutch size, egg mass and egg width) differed among the three populations. The egg‐laying date, ranging from late June to early August, varied among populations in a geographically continuous trend, with females at the most northern latitude (DH) laying eggs latest, and females at the most southern latitude (CZ) laying eggs earliest. Such a trend was less evident or even absent in the other traits that differed among the three populations. CZ and DH females, although separated by a distance of approximately 1100 km as the crow flies, were similar to each other in most traits examined. LS females were distinguished from CZ and DH females by the fact that they laid a greater number of eggs, but these were smaller. The egg size–number trade‐off was evident in each of the three populations and, at a given level of relative fecundity, egg mass was significantly greater in the DH population than in the LS population. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 701–709.     

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.     

水稻二化螟的交配行为

在室内条件下,对水稻二化螟Chilo suppressalis的交配行为及能力进行了研究.结果表明：大多数二化螟雌蛾一生只交配一次,平均0.92次;而雄蛾具有多次交配能力,最多达4次,平均2.72次.二化螟雌蛾的日龄影响其交配率、交配起始时间和持续时间,随二化螟雌蛾日龄的增加,其交配率逐渐下降,交配起始时间逐渐提前,而交配持续时间逐渐上升.相反,二化螟雄蛾日龄对其交配率、交配起始时间和持续时间没有明显影响.交配日龄对二化螟雌蛾的生殖力也存在显著影响,随着二化螟雌蛾交配日龄的增加,雌蛾产卵量下降,卵孵化率降低,产卵期缩短,它们都与雌蛾交配日龄存在显著的负相关;而雌蛾产卵前期和雌蛾寿命随雌蛾交配日龄的增加而延长,与雌蛾交配日龄存在显著的正相关.但二化螟雄蛾交配日龄对雌蛾的生殖力没有明显影响,二化螟雄蛾一生都具有较强的交配繁殖能力.同时,不同交配史的雄蛾与雌蛾交配,对雌蛾的生殖力也没有显著影响.表明二化螟的交配活动是由雌蛾主导控制的.最后,对这些结果在二化螟性信息素防治中应用的可行性进行了探讨.在应用性信息素控制二化螟的实践中,可以在两方面取得实效,一是性信息素可以阻碍雌雄之间正常交配,降低交配率;二是可以推迟二化螟雌虫的交配,使其产卵量和卵孵化率降低.     

Laying, egg and hatchability characteristics in ostrich (Struthio camelus) at different age

The experimental material comprised 7 ostrich families (7 males and 14 females) of which five families were at the age of 7 and two at the age of 5 years. In the course of the entire reproductive season, the following parameters were analysed: length of the laying period, mean laying rate, number of eggs laid by one female, proportion ofhatching eggs, egg weight and shape, egg weight lost during incubation, egg fertilisation, percentage of dead embryos and unhatched chicks, hatchability from fertilised and set eggs. Seven year-old ostriches were characterised by shorter laying period (134 days) but, at the same time, by higher proportions of hatching eggs. This group was also characterised by high egg fertilisation (79.7%) as well as high hatchability indices at simultaneous highest embryo mortality during incubation (11.6 %). Five year-old ostriches exhibited a longer laying period (175 days) during which females laid more eggs (49 pcs.). In addition, this group was characterised by a smaller proportion of hatching eggs, better egg fertilisation indices (83.5%) and hatchability results. Moreover, the determined higher egg shape index indicates that the 5 year-old females laid eggs which were more spherical. Recapitulating, the obtained results indicate that, under Polish conditions, better indices of laying performance, egg fertilisation and hatchability were observed in the group of 5 year-old ostriches.     

Age‐dependent relationships between coloration and reproduction in a species exhibiting delayed plumage maturation in females

Males of many bird species exhibit delayed plumage maturation (DPM), a condition in which young individuals display an immature plumage. Several adaptive hypotheses have been suggested for the signaling utility of DPM in males. Tree swallows Tachycineta bicolor, however, are one of the few bird species to exhibit DPM in females, but not in males. Few studies have focused on the age‐dependent signaling function of female plumage traits due to the uncommon nature of DPM in females. Therefore, we used reflectance spectrometry and scanning electron microscopy of sub‐adult (melanin‐based brown) and adult (iridescent‐blue structural) female tree swallows to characterize plumage coloration. Next, we asked whether variation in plumage coloration in females reflects condition and reproductive performance between and within age classes. We found that older females were in better body condition and laid eggs earlier in the season compared to young females; however, average egg mass and reproductive success (number of offspring fledged and offspring condition) did not differ between age classes. There were significant relationships indicating that young females with more‐ornamented (darker brown melanin) plumage laid smaller eggs, but hatched eggs earlier in the season leading to nestlings in better condition compared to less‐ornamented young females. Older females that were more ornamented (brighter, greater blue chroma, and lower hues in iridescent plumage) laid heavier eggs, but ornamentation was negatively associated with immune function, health, and reproductive success. Together, these data suggest that female ornamentation reflects reproductive performance and that there are complicated relationships between plumage coloration, condition, and reproductive performance that ultimately influence reproductive success.     

Oviposition of potato tuber moth,Phthorimaea operculella zell. (Lepidoptera: Gelechiidae); fecundity in relation to mated state,age, and pupal weight

The fecundity of individually held potato tuber moths ranged from 0 to 236 eggs deposited over the total life span. The number of eggs laid was not correlated with pupal weight, but pupal weight and the number of mature eggs in the ovaries shortly after emergence from the pupa were positively correlated. This initial egg complement accounted for slightly more than half of the total number of eggs laid. Among moths held in groups at 25°c in the absence of host‐plant material, multiple‐mated females did not lay significantly more eggs (mean 98.4) than those mated only once (mean 91.0), but their life span was shorter (8.5 days cf. 14.4). Virgin females laid a small number of non‐viable eggs (mean 7.7), and lived about as long as single‐mated females. Males were significantly longer‐lived (23.6 days) than all groups of females. Peak oviposition of mated females occurred 2–5 days after emergence, and declined to low levels by age 7 days.     

A survival cost to mating in a polyandrous butterfly, Colias eurytheme

Adaptations that enhance fitness in one sex may be harmful to members of the opposite sex and lead to antagonistic coevolution between the sexes. In fruit flies, for example, selection for fertilization success has rendered the male ejaculate slightly toxic to females. Here we investigated whether mating imposes a cost upon female fitness in a polyandrous pierid butterfly ( Colias eurytheme ) by comparing life history traits between once-mated females and virgins. Mated females laid relatively more eggs early in their adult life, but suffered a reduction in longevity relative to virgins held under identical experimental conditions. The effect of mating on female survivorship was statistically independent of lifetime and early life fecundity. Moreover, lifetime fecundity co-varied positively with longevity across all females, and across females within each treatment group, hence there was no phenotypic trade-off between survival and reproduction. These results suggest that the observed longevity difference between virgin and mated females represents a true cost of mating, possibly arising from a toxic side effect of the male ejaculate. However, irrespective of this cost, virgin and mated females laid an equivalent lifetime number of eggs. Female C. eurytheme are also known to use nutrients from the male ejaculate to supplement their reproductive output, hence it is presently unclear how the observed longevity cost may have influenced the evolution of lifetime mating schedules in this polyandrous species.     

Seminal fluid reduces female longevity and stimulates egg production and sperm trigger oviposition in a moth

Previous studies suggest that a number of factors in relation to mating may reduce female longevity and stimulate egg production and oviposition. However, it is still not clear whether these factors act on these parameters independently or in a collective way. Here we carried out a series of experiments including mating trials and seminal fluid injection to determine the factors responsible for reducing female longevity and stimulating egg production and oviposition in relation to mating in the moth Ephestia kuehniella Zeller (Lepidoptera: Pyralidae). Results show that seminal fluid and sperm work collectively to allow females to achieve maximum realized fecundity (number of eggs laid) in E. kuehniella but these factors play different roles in the process and their actions are independent. Seminal fluid signals females to allocate resources to ova, resulting in shorter longevity and greater egg production while eupyrene (not apyrene) sperm in the spermatheca trigger females to lay maximum number of eggs. We suggest that the receptors for seminal fluid signal may be located in the female reproductive tract and haemolymph, and those for sperm signal may be in the spermatheca. Hypotheses that females prolong their longevity by oosorption, physical injuries by males reduce female longevity, and mechanical stimulation by males triggers oviposition, are not substantiated in the present study.     

Mating and social contact change egg production and longevity in adult females of the mirid Lygus hesperus

There is frequently a tradeoff between fecundity and longevity, but the relationship is inconsistent across species and influenced by various exogenous and endogenous factors. Previous studies of Lygus hesperus Knight (Hemiptera: Miridae) established that egg production is promoted by insemination, at least temporarily, but little is known about the long‐term effects of mating and nonsexual interactions with conspecifics on egg production and female lifespan. To elucidate these relationships, survivorship and oviposition rate were tracked daily in females that were isolated or paired with a fertile male or another female throughout their adult lives. Mating rates were determined by postmortem examination. Results indicate that male‐specific stimuli accelerate female reproductive maturation, and that mating elevates oviposition rate. However, females paired with either a female or male companion had shortened lifespans, suggesting that social contacts exact a significant cost in this solitary species. Despite the negative impact of conspecific interactions and the finding that a singly mated female has sufficient sperm to fertilize a lifetime supply of eggs, many females were found to have mated more than once. Multiply mated females had higher sustained oviposition rates, lived longer, and had greater lifetime fecundities. Collectively, no strong evidence was found of a direct physiological link between fecundity and longevity, but environmental factors and mating were found to significantly influence both traits.     

Egg formation and the pre-laying period of the Common guillemot Una aalge

The patterns of colony attendance of male and female Common guillemots, relative to calendar date and relative to the female's laying date during the three weeks prior to egg laying, are presented. The probability of a male being at the colony was consistently higher than that for females. Male attendance peaked in the three days before his mate laid: female attendance was lowest at this time. Examination of the yolk showed that the egg was formed over 14–15 days, with yolk deposition (of first eggs) taking 11–5 days, and a lag period (between the end of yolk deposition and laying) of 3–4 days. Yolk deposition occurred over a shorter period (9–3 days) in replacement eggs, and followed a different pattern from first eggs.     

Mating frequency and reproductive success in an income breeding moth,Mnesampela privata

The frequency of mating in insects is often an important determinant of female reproductive output and male sperm competition. In Lepidoptera that provide male nutrients to the female when mating, it is hypothesized that polyandry may be more prevalent. This is thought to be especially so among species described as income breeders; that is, in species who do not derive all their nutrients for reproductive output entirely from the resources obtained during the larval stage. We selected the geometrid moth, Mnesampela privata (Guenée) (Lepidoptera: Geometridae), to examine this hypothesis further. We found this species was best characterized as an income breeder with female weight on emergence positively correlated with total egg load but not with the number of eggs laid. Further, in accord with income breeders, females emerged with a partially developed egg load and lifetime fecundity was positively correlated with the number of oviposition days. However, in the laboratory we found that incidence of repeated matings or polyandry was rare. When moths were paired singly over their lifetime, only 4% of mated females multiple mated. When females were paired with three males concurrently, female mating success increased from 60 to 81% with multiple mating among mated females increasing to just 15%. Dissection of wild caught M. privata found that polyandry levels were also low with a maximum of 16.4% of females collected at any one time being multiple mated. In accord with theory, mating significantly increased the longevity of females, but not of males, suggesting that females acquire essential resources from male ejaculates. Despite this, multiple mated females showed a trend toward decreasing rather than increasing female reproductive output. Spermatophore size, measured on death of the female, was not correlated with male or female forewing length but was negatively correlated with the number of fertile eggs laid and female longevity. Smaller spermatophore width may be related to uptake of more nutrients by the female from a spermatophore. We discuss our findings in relation to income breeding and its relationship to polyandry in Lepidoptera.