Interactions of mating,egg production and death rates in females of the Mediterranean fruit fly,Ceratitis capitata.

Costs of reproduction include costs of producing eggs and of mating itself. In the present study, we made an experimental investigation of costs of reproduction in the Mediterranean fruit fly (medfly, Ceratitis capitata). We demonstrated that virgins live longer than non-virgin females. However, in strong contrast to most findings within the Diptera, non-virginity had no detectable effect on egg production. Therefore the increased longevity of the virgin females cannot be attributed to an increase in egg production in non-virgin females, and instead indicates a cost of mating. A comparison of the life spans of normal females and those sterilized by low doses of X-irradiation, revealed an additional cost of egg production. There were no significant differences in remating levels between females that did and did not lay eggs, showing that the cost of producing eggs is independent of mating frequency. Medfly females therefore suffer a decrease in survival as a result of egg production and of mating, and these costs are independent of one another. To put our results into context, we reviewed the existing literature on the effects of mating on longevity, egg production and sexual receptivity for 64 species of Diptera, and examined the pattern of mating effects that emerged.     

Low cost of reproduction in female stalk-eyed flies, Cyrtodiopsis dalmanni

We investigated the effect of exposure to males on female longevity and egg production in the stalk-eyed fly, Cyrtodiopsis dalmanni. In this species, females typically mate several times each morning over a lifetime that can span several months. We hypothesised that high costs of mating with males would be incompatible with the life history of this species. We predicted that 1) female longevity costs of exposure to males would be low, and 2) that the magnitude of mating-induced fecundity increases in C. dalmanni, if present, would also be low. We tested the predictions by comparing the longevity and lifetime fecundity of virgin and mated females. In line with prediction 1), the longevity difference between virgin and fully reproductive females was small and of borderline significance. In line with prediction 2), egg production was not significantly higher in females continually exposed to males than for virgin females. Our results suggest that costs of reproduction resulting from exposure to males are low in species that mate promiscuously at high rates.     

Copulation reduces male but not female longevity in Saltella sphondylli (Diptera: Sepsidae)

Mating more than once is extremely costly for females in many species, making the near ubiquity of polyandry difficult to understand. However, evidence of mating costs for males is much rarer. We investigated the effects of copulation on longevity of male and female flies (Saltella sphondylli). We also scrutinized potential fecundity and fertility benefits to females with differing mating history. Copulation per se was found to decrease the longevity of males but not that of females. However, when females were allowed to lay eggs, females that mated died earlier than virgin females, indicating costs of egg production and/or oviposition. Thus, although longevity costs of copulation are higher for males, reproduction is nevertheless costly for females. We also found no differences in fecundity or fertility relative to female mating history. Results suggest that polyandry may be driven by minor costs rather than by major benefits in this species.     

Single and multiple mating reduces longevity of female dumpling squid (Euprymna tasmanica)

For many species, mating is a necessary yet costly activity. The costs involved can have an important influence on the evolution of life histories and senescence. Females of many species mate multiply, and this behaviour can inflict a longevity cost. Most studies investigating the effects of multiple mating on female survival have been conducted on insects, and the effects in other taxa are largely unknown. We investigate the effects of both a single mating and a second mating on longevity in female dumpling squid (Euprymna tasmanica), a species in which both sexes mate multiply. Through comparing the longevity of virgin, once‐mated and twice‐mated females, we found that a single mating reduced female life span by 15 days on average. A second mating resulted in an additional 8 day (on average) longevity cost, despite no difference in total clutch mass, number of clutches, single egg mass or number of eggs per clutch between once‐mated and twice‐mated females. This demonstrates a cost to multiple mating which may be independent of the cost of egg production. Furthermore, total clutch mass and female life span were positively correlated, whereas female life span decreased with increasing average water temperature. The presence of an additive effect of reproduction on longevity suggests that multiple mating in cephalopods may have benefits that outweigh these costs, or that there is a conflict in optimal mating frequency between males and females.     

The effect of male mating history on paternal investment, fecundity and female remating in the seed beetle Callosobruchus maculatus

1. In many organisms, males provide nutrients to females via ejaculates that can influence female fecundity, longevity and mating behaviour. The effect of male mating history on male ejaculate size, female fecundity, female longevity and female remating behaviour in the seed beetle Callosobruchus maculatus was determined.
2. The quantity of ejaculate passed to females declined dramatically with successive matings. Despite the decline, a male's ability to fertilize a female fully did not appear to decline substantially until his fourth mating.
3. When females multiply mated with males of a particular mated status, the pattern of egg production was cyclic, with egg production increasing after mating. Females multiply mated to virgins had higher fecundity than females mated to non-virgins, and females mated to twice-mated males had disproportionately increased egg production late in their life.
4. Females that mated to multiple virgins, and consequently laid more eggs, experienced greater mortality than females mated only once or mated to non-virgins, suggesting that egg production is costly, and rather than ameliorating these costs, male ejaculates may increase them by allowing or stimulating females to lay more eggs.
5. Females mating with non-virgin males remated more readily than did females mated to virgins. Females given food supplements were less likely to remate than females that were nutritionally stressed, suggesting that females remate in part to obtain additional nutrients.     

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.     

Life Cycle and Mating Behavior of Belonolaimus longicaudatus in Gnotobiotic Culture

The life cycle of Belonolaimus longicaudatus was observed in vitro on excised roots of Zea mays. Roots were cultured on Gamborg''s B5 medium in petri dishes with 1.5% agar adjusted to pH 5.8 and incubated at 28 °C in darkness. Second-stage juveniles (J2) fed on the roots and started the second molt (M2) to the third-stage juveniles 2 days after inoculation (DAI). The third molt (M3) to the fourth-stage juveniles occurred 7 DAI, followed by the fourth molt (M4) to males 13 DAI or to females 14 DAI. Nematode gender differences were observed by the end of the fourth molt. The first male appeared 15 DAI and the first female 17 DAI, after which mating occurred. Males were attracted to females, and mating was observed. Mating was required for reproduction. Fertilized females began to lay eggs 19 DAI and continued egg laying without the further presence of males during a 90-day observation. All of the eggs hatched. Unfertilized females rarely laid eggs, and none of the eggs were able to hatch. Feeding took place between each molt and before egg deposition occurred. The first-stage juveniles molted in the eggs 4 days after deposition, and J2 hatched from eggs 5 days after egg deposition. The life cycle from J2 to J2 was completed in 24 days.     

The costs of egg production and incubation in great tits (Parus major).

The costs of egg production and incubation may have a crucial effect on avian reproductive decisions, such as clutch size and the timing of reproduction. We carried out a brood-size enlargement experiment on the great tit (Parus major), in which the birds had to lay and incubate extra eggs (full costs), only incubate extra eggs (free eggs) or did not pay any extra cost (free chicks) in obtaining a larger brood. We used female fitness (half the recruits produced plus female survival) as a fitness measure because it is the female which pays the costs of egg production and incubation, and because clutch size is under female control. Female fitness decreased with increasing costs (fitness of free chicks females is higher than that of free eggs females which is higher than that of full costs females). These fitness differences were due to differences in female survival rather than in the number of recruits produced. This is the first time that the costs of egg production and incubation have been estimated using such a complete fitness measure, including, as our measure does, the local survival to the following year of both the female and her offspring. Our results emphasize that reproductive decisions cannot be understood without taking egg production and incubation costs into account.     

Cost of immune priming within generations: trade-off between infection and reproduction

Immune priming is a new paradigm in innate immunity. However, most studies have focused on the benefits of priming (enhanced survival and parasite clearance after a second challenge), while little attention has been paid to the costs. In this study, both factors were investigated in Anopheles albimanus primed against Plasmodium berghei. As previously observed in other invertebrates, compared to un-primed mosquitoes, those primed better controlled a challenge from the same parasite, and had a higher survival rate. Although there was no difference in the number of oviposited eggs between primed and control females, hatching rate was lower in primed than in control mosquitoes and it was more likely for control females to produce eggs than for primed females. Furthermore, a trade-off between parasite elimination and egg production was observed among primed mosquitoes, as primed females that successfully fought the infection were unable to produce eggs, but primed females that produced eggs were similarly infected as control un-primed ones. These results concord with recent mathematical models suggesting that reproduction affects immune priming outcomes, and may explain why in some species and under some conditions it seems that immune priming is not occurring.     

Longevity cost of reproduction for males but no longevity cost of mating or courtship for females in the male-dimorphic dung beetle Onthophagus binodis

Life history theory predicts a trade-off between current and future reproduction. Despite a wealth of research on the cost of reproduction for females, there have been very few studies that have looked at the cost of reproduction for males. Longevity is closely related to the opportunity for future reproduction, and thus decreased longevity in response to current reproductive effort has been used as a measure of the cost of reproduction. Here we examine the cost of reproduction for males and females in the dung beetle Onthophagus binodis. Like many onthophagines, O. binodis exhibit dimorphic male morphology; major males develop a large pronotal horn while minor males remain hornless. Alternative morphologies are associated with alternative reproductive tactics. Thus, we ask whether major and minor males pay different costs of reproduction. We found that in contrast to previous work on Diptera, mating is not costly in terms of reduced longevity for female dung beetles. Despite a longevity cost of reproduction for males, we found no evidence for differential longevity costs associated with alternative reproductive tactics.     

Effects of multiple mating on the fecundity of an invasive pest (Octodonta nipae): the existence of an intermediate optimal female mating rate

Multiple mating is found in many insect taxa where both of the sexes can mate more than once. For males, this leads to the advantage of increasing their paternity by fertilizing more females. However, there is a trade‐off of resource allocation between reproduction and other life‐history characters. In the present study, the impact of increased mating rate on reproductive fitness of the invasive nipa palm hispid beetle Octodonta nipae Maulik (Coleoptera: Chrysomelidae) is investigated. A series of mating frequencies (i.e. 1, 5, 10, 15, 20 times) is selected from video frame playback, ranking from the minimum to maximum mating rate observed under laboratory conditions over a given time period. Fecundity parameters such as lifetime egg production, egg‐hatching rate, effective oviposition period and longevity are investigated for the evaluation of reproductive efficiency. For female O. nipae, increased fecundity is correlated with the mating frequency. Females mating 15 times lay the largest number of eggs (138.82 ± 6.87) and have a hatching rate of 47.43 ± 4.08%. After mating 20 times, females suffer significant declines in oviposition (90.31 ± 8.38 eggs) and egg‐hatching rate (34.16 ± 4.93%). Moreover, the population growth rate reaches a maximum in the females that mate 15 times. The results show that multiple matings in O. nipae have an intermediate optimal range within which female reproductive success is enhanced, providing empirical evidence for the existence of a trade‐off between costs and benefits during copulation based on resource allocation.     

The effect of ovariectomy on reproductive behaviour and physiology of adult female ring-legged earwigs, Euborellia annulipes

The effect of ovariectomy on feeding, mating, Juvenile Hormone (JH) production, and maternal behaviour was assessed in female ring-legged earwigs, Euborellia annulipes (Lucas) (Dermaptera: Carcinophoridae), during the first 16 days of adult life (the first gonadotrophic cycle and early brooding). Ovariectomy of 2-day-old adults did not affect weight gain, nor did it alter mating behaviour on day 7. Similarly, ovariectomy did not prevent the increase in JH biosynthesis that accompanies vitellogenesis in this species, which suggests a cycle of JH production that is not dependent on the presence of the ovaries. Both ovariectomy and mating status affected feeding behaviour. Most introduced eggs were consumed (i.e. disappeared) within 24 h, and younger (7-day-old) females consumed more eggs than did older ones. However, 12-day-old intact virgins and 16-day-old ovariectomized, mated females consumed fewest eggs, and allowed some hatching. Thus, ovariectomy did not abolish changes in feeding behaviour that normally accompany reproduction but, instead, appeared to delay the reduction in feeding that normally accompanies the completion of the cycle of egg development. By contrast, mating enhanced the reduction in feeding late in the reproductive cycle. Mating significantly enhanced maternal behaviour in both ovariectomized and sham-operated females. Hatching success from egg clutches introduced to day 16 virgin or mated females that had been ovariectomized or sham-operated on day 2, was significantly greater in the mated groups.     

Host plant effects on development and reproduction of the glassy-winged sharpshooter, Homalodisca vitripennis (Homoptera: Cicadellidae)

Development, survivorship, longevity, reproduction, and life table parameters of the glassy-winged sharpshooter, Homalodisca vitripennis (Germar), were examined in the laboratory using three host plants, sunflower (Helianthus annuus L.), Chrysanthemum morifolium L., and euonymus (Euonymus japonica Thurb.). Females deposited similar-sized egg masses on all three plants. Hatching was highest with eggs deposited on euonymus and lowest for those deposited on sunflower. Embryonic development time among host plants was similar while nymph development time was shortest on sunflower and longest on euonymus. Nymph survival to adulthood ranged from 32% on euonymus to 82% for those reared on sunflower. Adult females had similar life spans on sunflower and chrysanthemum. H. vitripennis completed a lengthy egg-to-adult development on euonymus, however, mating did not occur. The onset of mating was contingent on maturation of adult females. The majority of mating activity occurred within the first three days after onset. Premating periods ranged from 6 to 7 d on sunflower to 27 d on chrysanthemum, with overall mating rates of 77.4 and 19.8%, respectively. Females typically mated more than once and they had the longest oviposition period and highest egg production on sunflower; ≈ 50 and 67% of total number of eggs were deposited within first 45 d after the start of oviposition on sunflower and chrysanthemum, respectively. Adult size and weight related to which host plant was consumed throughout development. Greater intrinsic and finite rates of increase and net reproduction rate, and shorter population doubling time occurred when the sharpshooters were allowed to develop on sunflower. The overall developmental and reproductive parameters obtained in this study indicate that a mixed host plant system, composed of sunflower and euonymus or chrysanthemum plants, is an efficient means for optimizing egg production and colony maintenance of the glassy-winged sharpshooter.     

Multiple mating in the traumatically inseminating Warehouse pirate bug, Xylocoris flavipes: effects on fecundity and longevity

Optimal mating frequencies differ between sexes as a consequence of the sexual differentiation of reproductive costs per mating, where mating is normally more costly to females than males. In mating systems where sexual reproduction is costly to females, sexual conflict may cause both direct (i.e. by reducing female fecundity or causing mortality) and indirect (i.e. increased risk of mortality, reduced offspring viability) reductions in lifetime reproductive success of females, which have individual and population consequences. We investigated the direct and indirect costs of multiple mating in a traumatically inseminating (TI) predatory Warehouse pirate bug, Xylocoris flavipes (Reuter) (Hemiptera: Anthocoridae), where the male penetrates the female's abdomen during copulation. This study aimed to quantify the effects of TI on female fecundity, egg viability, the lifetime fecundity schedule, longevity and prey consumption in this cosmopolitan biocontrol agent. We found no difference in the total reproductive output between mating treatments in terms of total eggs laid or offspring viability, but there were significant differences found in daily fecundity schedules and adult longevity. In terms of lifetime reproduction, female Warehouse pirate bugs appear to be adapted to compensate for the costs of TI mating to their longevity.     

Repetitive mating and female fitness in Dysdercus cardinalis (Hemiptera: Pyrrhocoridae)

Using three mating-treatment groups–pairing with a male for life, pairing with a male from adult emergence to first oviposition, and no pairing (virgin)–of Dysdercus cardinalis females in a laboratory study, the following results were obtained. Most of the virgin females did not lay eggs; those that did oviposited for the first time at a considerably older age than females in the other two groups. In a lifetime, females pairing with a male for life and those pairing with a male up to first oviposition produced essentially the same number of eggs, and this was more than 3.5 times the number of eggs laid by a virgin female. Virgin females produced an average of 1.3 clutches in a lifetime, compared with about 4.5 clutches produced by females in the other two groups. Reproductive span was significantly shorter, and fecundity per day of reproductive span significantly greater, for females pairing with males for life than for those pairing with a male up to first oviposition. Age at death was significantly different amongst females in different mating-treatment groups. Virgin females had the longest life, followed by females pairing with a male up to first oviposition and females pairing with a male for life, in that order. There was a positive correlation between lifetime gross reproduction and age at death for females pairing with a male for life. There was no relationship between these two characters for females pairing with a male up to first oviposition. Both females pairing with a male for life and those pairing with a male to first oviposition exhibited a significant inverse relationship between fecundity per day and age at death. The results obtained indicate that (1) mating is a prerequisite for normal gonadal activity in Dysdercus cardinalis , and (2) repetitive mating increases the rate of reproduction. This would reduce the mean age of parenthood which is inversely related to the intrinsic rate of increase.     

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.     

Life cycle and mating behavior of <Emphasis Type="Italic">Helicotylenchus multicinctus</Emphasis> (Nematoda: Hoplolaimidae) on excised <Emphasis Type="Italic">Musa cavendishii</Emphasis> roots

The life cycle and mating behavior of Helicotylenchus multicinctus (Nematoda: Hoplolaimidae) were observed in vitro on excised roots of Musa cavendishii in gnotobiotic culture. Eggs hatched into juveniles whose appearance and structure were similar to those of the adults. Juveniles grew in size and each juvenile stage was terminated by a molt. H. multicinctus had four juvenile stages. The first molt occurred outside the egg shortly after hatching. After the final molt the juveniles differentiated into adult males and females. Mating was required for reproduction. After mating, fertilized females began to lay eggs. The life cycle from second stage juvenile to second stage juvenile was completed in 39 days.     

Size specific demography of three species of Anastrepha fruit flies

Individuals of three Anastrepha species: A. obliqua, A. ludens, and A. serpentina (Diptera: Tephritidae), were sorted according to pupal weight in cohorts of large and small flies. Demographic parameters and reproductive patterns and heterogeneity were determined for each cohort. Large flies of the three species presented greater expectation of life and gross fecundity rates. A. ludens was the species with the longest life span (expectation of life of large adults was 110 days) and the greatest gross fecundity rates (1597 eggs/female for the large flies). While, A. obliqua had the shortest mean age of reproduction (33 days), and the greatest daily egg production (14 eggs/female/day). Net fecundity was similar in these two. A. serpentina had lower fecundity rates.Reproductive information for each size and each species include: age-by-parity relations, fraction of sexually mature life in which females lay eggs, and frequency distribution of individual egg production. Results demonstrate that even under constant laboratory conditions and using standard artificial hosts, there is a great deal of life history variation among these Anastrepha species and among other tephritid fruit flies.     

Are there costs in the repeated mating activities of female Southern stink bugs Nezara viridula?

Mating is an energy demanding process, imposing risks to physical injuries, pathogen infection and predation. Nevertheless, repeated and multiple mating are widespread even in insect species where nuptial gifts are not involved. The effects of repeated mating, by the same male, are examined on the reproductive performance of female Southern green stink bug Nezara viridula (L.). Fecundity is reduced in females mated three or four times, although there is increased longevity. Females mated once or twice produce more egg clusters and concentrate egg‐laying activity in the early part of adult life, whereas those mating more often lay eggs throughout the life span, with fewer egg clusters. Although fecundity is negatively affected by the number of matings, egg fertility remains unaffected.