Effects of sequential mating by males on reproductive output of the stinkbug predator, Podisus nigrispinus

Reproductive output of the stinkbug predator Podisus nigrispinus (Dallas) (Heteroptera: Pentatomidae) was investigated as a function of the number of matings that the male had made with a range of females. After being placed with a female, virgin males were most likely to mate within 12 hours, while non-virgin males were most likely to mate within 12–24 hours. Although males lost weight during their first mating, the weights of mated and unmated males were not significantly different throughout their lifetime. Longevity was significantly greater for unmated males (36.0 days) than for mated males (29.8 days). Survival curves for both mated and unmated males were Type II. The capacity of males to transfer sperm to virgin females was not affected by previous matings. From 65.7 to 76.4% of eggs were viable and 206.7 to 274.6 nymphs were produced per female. Regardless of the number of matings that the male had made, females that had mated only once exhausted their stored sperm progressively and produced an increasing proportion of infertile eggs, which peaked at the end of their lives. These results show that P. nigrispinus females need more than one mating to maintain fertility, but their performance is not affected by the number of previous matings that the male has made or by male weight. Thus, the strategy of pairing with males multiple times improved production efficiency by increasing output and reducing food waste in mass production systems. This is achieved by temporarily pairing females at intervals of about 20 days during their entire lifetime.     

Variation in the benefits of multiple mating on female fertility in wild stalk‐eyed flies

Polyandry, female mating with multiple males, is widespread across many taxa and almost ubiquitous in insects. This conflicts with the traditional idea that females are constrained by their comparatively large investment in each offspring, and so should only need to mate once or a few times. Females may need to mate multiply to gain sufficient sperm supplies to maintain their fertility, especially in species in which male promiscuity results in division of their ejaculate among many females. Here, we take a novel approach, utilizing wild‐caught individuals to explore how natural variation among females and males influences fertility gains for females. We studied this in the Malaysian stalk‐eyed fly species Teleopsis dalmanni. After an additional mating, females benefit from greatly increased fertility (proportion fertile eggs). Gains from multiple mating are not uniform across females; they are greatest when females have high fecundity or low fertility. Fertility gains also vary spatially, as we find an additional strong effect of the stream from which females were collected. Responses were unaffected by male mating history (males kept with females or in male‐only groups). Recent male mating may be of lesser importance because males in many species, including T. dalmanni, partition their ejaculate to maintain their fertility over many matings. This study highlights the importance of complementing laboratory studies with data on wild‐caught populations, where there is considerable heterogeneity between individuals. Future research should focus on environmental, demographic and genetic factors that are likely to significantly influence variation in individual female fecundity and fertility.     

Mating Behavior and Reproductive Potential in the Turnip Moth Agrotis segetum (Lepidoptera: Noctuidae)

We investigated the lifetime mating potential and the reproductive behavior of male and female turnip moths Agrotis segetum (Schiff.) under field and laboratory conditions. The sex ratio was 1 : 1 in a lab-reared population as well as in two wild populations. Males were capable of mating repetitively a relatively large number of times (mean of 6.7 ± 2.7 matings) when given access to new virgin females throughout their lifetimes. Females seldom mated more than once (mean ± 1.3 ± 0.6 matings), indicating a male-biased operational sex ratio. The mean potential lifetime mating was five times higher in males, while the coefficient of variance was lower in males. There was no differences in longevity between animals that were allowed to mate and animals not allowed to mate, indicating no direct costs or benefits of mating in physiological terms. In males, the number of matings was positively correlated with longevity, but this was not the case in females. Nor was there a correlation between the number of female matings and the number of fertilized eggs. There was a negative correlation between the number of eggs fertilized and the number of times males had previously mated, indicating that male ejaculates were limited. Male spermatophore size also decreased with number of achieved matings. Laboratory-reared females attracted males in the field throughout their lifetimes, with a peak at 3–7 days of age. Wild males, allowed to choose between pairs of caged females in the field, were attracted in equal numbers to females of different ages. Females did not show any mate-rejection behavior in the field. They mated with the first male that courted them. No incidence of mate replacement by males arriving later to already courted females were recorded.     

The effect of mating frequency and mating pattern on female reproductive fitness in cabbage beetle,Colaphellus bowringi

It is generally thought that females can receive more of the material benefits from males by increasing mating frequency and polyandry can lead to greater reproductive success. The cabbage beetle, Colaphellus bowringi Baly (Coleoptera: Chrysomelidae), is a highly promiscuous species, in which females or males can readily mate repeatedly with a given partner or multiple partners at a very high frequency. In the present study, the effect of mating frequency (number of matings) and mating pattern (polyandry vs. monogamy) on female reproductive fitness was investigated by measuring fecundity, fertility, and female longevity. The results indicated that increased female mating frequency with the same male did not result in variation in lifetime fecundity, but significantly increased fertility and decreased female longevity. Moreover, five copulations were sufficient to acquire maximal reproductive potential. Female lifetime fecundity also did not differ between polyandrous and monogamous treatments. However, monogamous females exhibited a significant increase in fertility and significant prolongation of longevity compared with polyandrous females, further demonstrating that monogamy is superior to polyandry in this beetle.     

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.     

Forced Copulations and Female Contact Guarding at a High Male Density in a Calopterygid Damselfly

Territorial males of Calopteryx damselflies court females on territories that contain oviposition substrates. Nonterritorial males try to mate without courtship but very rarely obtain matings because females fail to bring up their abdomen to engage genitalia. Here I report the results of observations made on a very high-density population of Calopteryx haemorrhoidalis in central Italy. Mating activity was intense, and during 40 h of observation in an 8-m section of the stream, 209 matings were recorded (a maximum of 17 matings h ^–1 ). Males were continuously disturbing ovipositing females and tried to achieve tandem forcibly. Of 84 cases, males achieved forced tandem in 53, and 49 ended with copulation. Forced tandems were the most common method to obtain a mating in this population (55% of 65 matings). Males guarded females after forced or courtship copulations and, in some cases, maintained physical contact with their mate, by perching on her wings. Confusion was common and males guarded nonmates frequently, which suggests that they were unable to recognize their mate individually.     

No material benefits, and a fertilization cost, for multiple mating by female leaf beetles

When females mate multiple times it presents an intriguing problem for evolutionary biologists due to the high costs and not very apparent benefits. Yet, this behaviour must have higher benefits than costs to be maintained by natural selection. We studied possible benefits for multiple mating by female leaf beetles, Leptinotarsa decemlineata, both with multiple males (polyandry) and multiple times with the same male (repeated matings). For polyandry, we tested the material benefits hypothesis, as well as fertility insurance, by mating females to varying numbers of males and looking at sperm received and fecundity. Although there was a relationship between amount of stored sperm and number of matings, we did not find the predicted positive relationship between fecundity and number of matings. Thus, the material benefits hypothesis was not supported. For repeated matings, we tested the sperm and nonsperm material benefit hypotheses by mating females to a single male varying numbers of times on one mount. For this set of tests neither sperm number nor fecundity was positively correlated with number of matings, therefore neither hypothesis was supported. Interestingly, in both experiments, there was a significant decrease in hatch rate with an increase in matings, demonstrating a cost of polyandry. These findings, along with previous research, suggest a cost of multiple mating with a possible role of seminal fluids in the reduction in female fecundity.     

Effects of multiple matings on reproductive fitness of male and female Diaeretiella rapae

Mating frequency and the amount of sperm transferred during mating have important consequences on progeny sex ratio and fitness of haplodiploid insects. Production of female offspring may be limited by the availability of sperm for fertilizing eggs. This study examined multiple mating and its effect on fitness of the cabbage aphid parasitoid Diaeretiella rapae McIntosh (Hymenoptera: Aphidiidae). Female D. rapae mated once, whereas males mated with on average more than three females in a single day. The minimum time lag between two consecutive matings by a male was 3 min, and the maximum number of matings a male achieved in a day was eight. Sperm depletion occurred as a consequence of multiple mating in D. rapae. The number of daughters produced by females that mated with multiple‐mated males was negatively correlated with the number of matings achieved by these males. Similarly, the proportion of female progeny decreased in females that mated with males that had already mated three times. Although the proportion of female progeny resulting from multiple mating decreased, the decrease was quicker when the mating occurred on the same day than when the matings occurred once per day over several days. Mating success of males initially increased after the first mating, but then males became ‘exhausted’ in later matings; their mating success decreased with the number of prior matings. The fertility of females was affected by mating with multiple‐mated males. The study suggests that male mating history affects the fitness of male and female D. rapae.     

Sperm Numbers and Female Fertility in the Moth Plodia interpunctella (Hübner) (Lepidoptera; Pyralidae)

I investigated two possible reasons for remating in female Plodia interpunctella: i) females remate to obtain sufficient sperm to maintain fertility; and ii) male investment in non-sperm components increases female fecundity and longevity. The number of sperm and the mass of the spermatophore transferred by males decreases on successive matings. Sperm numbers and potential male investment were varied by allowing females to mate either once or twice with males either on their first or second mating. Females receiving a single small spermatophore containing few sperm (from a male on his second mating) had sufficient sperm to fertilize all their eggs. Females did not show increased fecundity or longevity as a result of obtaining more spermatophore material. I discuss why females remate when they already have sufficient sperm to fertilize all their eggs.     

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.     

Females Remate for Sperm Replenishment in a Seed Bug: Evidence from Offspring Viability

Females of many insects mate multiply but why they do so remains controversial. Here we investigated the effects of multiple matings on female reproductive success of a New Zealand seed bug, Nysius huttoni. We found little evidence for females to gain material (nutritional) benefits through multiple matings because the number of matings did not have significant effect on female fecundity. Females remated to the same males or different males produced similar number of viable offspring, suggesting that females do not obtain genetic benefit from remating in terms of offspring viability. With the increase of the number of matings, however, overall fertility rate significantly increased and daily fertility rate declined significantly slower over time. These results suggest that females remate for the replenishment of sperm. Five matings are sufficient for females to maximize their reproductive success, and additional matings appear to be superfluous. However, the females of this bug mate as many as 68 times if males and females are paired for lifetime. This can be explained by the convenience hypothesis, i.e., females remate superfluously to minimize the costs of harassment by promiscuous males.     

FEMALES RECEIVE A LIFE-SPAN BENEFIT FROM MALE EJACULATES IN A FIELD CRICKET

Abstract.— Mating has been found to be costly for females of some species because of toxic products that males transfer to females in their seminal fluid. Such mating costs seem paradoxical, particularly for species in which females mate more frequently than is necessary to fertilize their eggs. Indeed, some studies suggest that females may benefit from mating more frequently. The effect of male ejaculates on female life span and lifetime fecundity was experimentally tested in the variable field cricket, Gryllus lineaticeps. In field crickets, females will mate repeatedly with a given male and mate with multiple males. Females that were experimentally mated either repeatedly or multiply lived more than 32% longer than singly mated females. In addition, multiply mated females produced 98% more eggs than singly mated females. Because females received only sperm and seminal fluid from males in the experimental matings, these life‐span and fecundity benefits may result from beneficial seminal fluid products that males transfer to females during mating. Mating benefits rather than mating costs may be common in many animals, particularly in species where female mate choice has a larger effect on male reproductive success than does the outcome of sperm competition.     

INTERSEXUAL SELECTION IN THE MEDITERRANEAN FRUIT FLY: DOES FEMALE CHOICE ENHANCE FITNESS?

Adaptive hypotheses of female choice predict that females use male courtship displays as an indicator of male quality. A test of whether female choice is adaptive measuring direct and indirect effects of mate choice on females was made using a laboratory population of a lek-mating species, the Mediterranean fruit fly. The nonrandom mating observed in this species is thought to be strongly influenced by female choice. Whether female choice acts to increase fecundity or offspring quality was assessed using two different statistical tests. Multiple regression showed that females generally do not receive direct benefits as a result of mating with males which are successful in copulating with many females. However, in one trial the relationship between male quality and female benefit was nonlinear. Females which mate with males that obtain few matings (<2), and females which mate with males that obtain many matings (>6) enjoy increased fecundity. Mate choice does not, however, appear to enhance offspring quality as father/son correlation and sibling analysis showed no heritable component to male copulatory success.     

Reproductive costs of mating with a sibling male: sperm depletion and multiple mating in Cephalonomia hyalinipennis

Polygynous parasitoid males may be limited by the amount of sperm they can transmit to females, which in turn may become sperm limited. In this study, I tested the effect of male mating history on copula duration, female fecundity, and offspring sex ratio, and the likelihood that females will have multiple mates, in the gregarious parasitoid Cephalonomia hyalinipennis Ashmead (Hymenoptera: Bethylidae: Epyrinae), a likely candidate for sperm depletion due to its local mate competition system. Males were eager to mate with the seven females presented in rapid succession. Copula duration did not differ with male mating history, but latency before a first mating was significantly longer than before consecutive matings. Male mating history had no bearing on female fecundity (number of offspring), but significantly influenced offspring sex ratio. The last female to mate with a given male produced significantly more male offspring than the first one, and eventually became sperm depleted. In contrast, the offspring sex ratio of first‐mated females was female biased, denoting a high degree of sex allocation control. Once‐mated females, whether sperm‐depleted or not, accepted a second mating after a period of oviposition. Sperm‐depleted females resumed production of fertilized eggs after a second mating. Young, recently mated females also accepted a second mating, but extended in‐copula courtship was observed. Carrying out multiple matings in this species thus seems to reduce the cost of being constrained to produce only haploid males after accepting copulation with a sperm‐depleted male. I discuss the reproductive fitness costs that females experience when mating solely with their sibling males and the reproductive fitness gain of males that persist in mating, even when almost sperm‐depleted. Behavioural observations support the hypothesis that females monitor their sperm stock. It is concluded that C. hyalinipennis is a species with a partial local mating system.     

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.     

Sperm‐limited fecundity and polyandry‐induced mortality in female nematodes Caenorhabditis remanei

In many sexually reproducing species, females are sperm limited and actively mate more than once which can lead to sperm competition between males. However, the costs and benefits of multiple matings may differ for males and females leading to different optimal mating frequencies and consequent sexual conflict. Under these circumstances, male traits that reduce females' re‐mating rates are likely to evolve. However, the same traits can also reduce, directly or indirectly, female survival and/or manipulate female fecundity. Evidence of this sexual conflict is common across several taxa. Here, we examine the evidence for this form of conflict in the free‐living nematodes of the Caenorhabditis genus. Members of this group are extensively used to describe developmental and physiological processes. Despite this, we understand little about the evolution of selfing, maintenance of males and sexual conflict in these species, particularly those with gonochoristic mating strategies. In this study, we demonstrate experimentally sexual conflict in the gonochoristic of C. remanei cultured under laboratory conditions. In our first experiment, we found that female fecundity increased with the number of males present which suggests that females' reproduction may be sperm limited. However, increasing the number of males present also reduced female survival. A second experiment ruled out the alternative explanation of density‐dependent reduction in female survival when more males were present as increasing female density correspondingly did not affect female survival. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 362–369.     

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

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

Ejaculate investment and attractiveness in the stalk‐eyed fly,Diasemopsis meigenii

The phenotype‐linked fertility hypothesis proposes that male fertility is advertised via phenotypic signals, explaining female preference for highly sexually ornamented males. An alternative view is that highly attractive males constrain their ejaculate allocation per mating so as to participate in a greater number of matings. Males are also expected to bias their ejaculate allocation to the most fecund females. We test these hypotheses in the African stalk‐eyed fly, Diasemopsis meigenii. We ask how male ejaculate allocation strategy is influenced by male eyespan and female size. Despite large eyespan males having larger internal reproductive organs, we found no association between male eyespan and spermatophore size or sperm number, lending no support to the phenotype‐linked fertility hypothesis. However, males mated for longer and transferred more sperm to large females. As female size was positively correlated with fecundity, this suggests that males gain a selective advantage by investing more in large females. Given these findings, we consider how female mate preference for large male eyespan can be adaptive despite the lack of obvious direct benefits.     

Female and Male Moths Display Different Reproductive Behavior when Facing New versus Previous Mates

Multiple mating allows females to obtain material (more sperm and nutrient) and/or genetic benefits. The genetic benefit models require sperm from different males to fertilize eggs competitively or the offspring be fathered by multiple males. To maximize genetic benefits from multiple mating, females have evolved strategies to prefer novel versus previous mates in their subsequent matings. However, the reproductive behavior during mate encounter, mate choice and egg laying in relation to discrimination and preference between sexes has been largely neglected. In the present study, we used novel and previous mate treatments and studied male and female behavior and reproductive output in Spodoptera litura. The results of this study do not support the sperm and nutrient replenishment hypotheses because neither the number of mates nor the number of copulations achieved by females significantly increased female fecundity, fertility and longevity. However, females showed different oviposition patterns when facing new versus previous mates by slowing down oviposition, which allows the last male has opportunities to fertilize her eggs and the female to promote offspring diversity. Moreover, females that have novel males present called earlier and more than females that have their previous mates present, whereas no significant differences were found on male courtship between treatments. These results suggest that S. litura females can distinguish novel from previous mates and prefer the former, whereas males generally remate regardless of whether the female is a previous mate or not. In S. litura, eggs are laid in large clusters and offspring competition, inbreeding and disease transfer risks are thus increased. Therefore, offspring diversity should be valuable for S. litura, and genetic benefits should be the main force behind the evolution of female behavioral strategies found in the present study.     

Female choice for heterozygous mates changes along successive matings in a lizard

Female mate choice and female multiple mating are major focuses of studies on sexual selection. In a multiple mating context, the benefits of mate choice can change along successive matings, and female choice would be expected to change accordingly. We investigated sequential female mate choice in the moderately polyandrous common lizard (Zootoca vivipara, synonym Lacerta vivipara). Along successive mating opportunities, we found that females were relatively unselective for the first mate, but accepted males of higher heterozygosity for subsequent mating, consistent with the trade-up choice hypothesis. We discuss the evidence of trade-up mate choice in squamates and generally trade-up for mate heterozygosity in order to motivate new studies to fill gaps on these questions.