Male traits under cryptic female choice in the spotted cucumber beetle (Coleoptera: Chrysomelidae)

Males of the spotted cucumber beetle (Diabrotica undecimpunctata howardi) rhythmically stroke females with their antennae during copulation. Males that stroke quickly have a higher probabilityof being accepted as a mate. We determined (1) the mechanismby which females prevent unattractive males from passing spermatophores,(2) whether antennal stroking signals to females the likelihoodof receiving a nuptial gift, and (3) if other male traits inaddition to stroking are subjected to sexual selection fromfemale preference. Dissections of pairs flash-frozen in copuladuring and after antennal stroking showed musculature that,when contracted, folded the vaginal duct leading to the female'sbursa copulatrix in a way that prevented complete penetrationby the aedeagus. These muscles were always contracted whilemales were stroking and always relaxed after stroking had ceased.Males accepted as mates did not differ from males that failedto pass a spermatophore in either absolute or relative bodyweight, aedeagus length, or the amount of cucurbitacins (potentialnuptial gifts) sequestered in their spermatophores. Although99% of the beetles that came to cucurbitacin-rich Cucurbitafruits in the field were males, males that had sequestered cucurbitacins did not stroke females faster than males withno cucurbitacins, and fast-stroking males were not more likelyto find and sequester cucurbitacins than were males that strokedmore slowly. Males with a cucurbitacin slurry painted on theirantennae had no mating advantage over controls. We concludethat females discriminate among males after copulation hasbegun on the basis of antennal stroking displays (or some traitcorrelated with stroking speed) that males perform to enticefemales to relax their bursal sphincter.     

Male genotype influences female reproductive investment in Drosophila melanogaster

In many species, males can influence the amount of resources their mates invest in reproduction. Two favoured hypotheses for this observation are that females assess male quality during courtship or copulation and alter their investment in offspring accordingly, or that males manipulate females to invest heavily in offspring produced soon after mating. Here, we examined whether there is genetic variation for males to influence female short-term reproductive investment in Drosophila melanogaster, a species with strong sexual selection and substantial sexual conflict. We measured the fecundity and egg size of females mated to males from multiple isofemale lines collected from populations around the globe. Although these traits were not strongly influenced by the male's population of origin, we found that 22 per cent of the variation in female short-term reproductive investment was attributable to the genotype of her mate. This is the first direct evidence that male D. melanogaster vary genetically in their proximate influence on female fecundity, egg size and overall reproductive investment.     

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.     

Characteristics of brown trout males influence growth and survival of their offspring

Half-sib groups of juvenile brown trout Salmo trutta (0+ years) having fathers with different characteristics were compared in several respects. Two kinds of experiments were performed, aquarium observations in the laboratory and stocking in a semi-natural stream. It was found in the aquarium experiments that the offspring of fathers with larger adipose fins were more active and were probably better at defending territories. Offspring of males chosen by females had higher growth rate. The feeding rate was higher in juveniles having a less aggressive father or a father that was chosen by the mother. The stream experiments showed that offspring of dominant males had higher growth rates, and that offspring of less aggressive fathers had higher survival. Thus, characteristics of the males (fathers) influences the quality of the offspring, and there seem to be benefits for the females both to choose among potential mates and to spawn with the most dominant male. As these male characteristics are never regarded during the process of artificial breeding in hatcheries, the results indicate a basis for divergence between hatchery and wild populations.     

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.     

Alternative Female Choice Tactics in the Scorpionfly Hylobittacus apicalis(Mecoptera) and Their Implications

Females of the scorpionfly Hylobittacus apicalis choose mateson the basis of material benefits (nuptial arthropod prey size)and probably on the basis of genetic benefits males deliverat mating. Females feed on the male's prey throughout copulation.They prefer males with large prey as mates and often refusemales who present small prey. That females may value male geneticquality is suggested by differences in ability of males to obtainlarge prey, which if inherited would influence offspring fitness,and by females often terminating mating with males with smallprey before they transfer any sperm or a complete ejaculate.Females hunt only when males with prey are not available becausehunting exposes individuals to predators. Female Hylobittacusapicalis exhibit alternative mate choice tactics, which arecondition-dependent in expression and probably comprise a conditionalstrategy. Body size, recent feeding history, and male availabilitydetermine how discriminating an individual female actually is,and these conditions may determine the value of material andgenetic benefits in mate choice decisions. The results suggestthat female choice controls male behavior. When females becomechoosy, males are forced to obtain rare large prey despite theincreased risks to males associated with this behavior. The implications of the findings on H. apicalis are discussedin relation to condition dependent female choice patterns inother species and the evolutionary maintenance of female choice.     

Harm to females increases with male body size in Drosophila melanogaster

Previous studies indicate that female Drosophila melanogaster are harmed by their mates through copulation. Here, we demonstrate that the harm that males inflict upon females increases with male size. Specifically, both the lifespan and egg-production rate of females decreased significantly as an increasing function of the body size of their mates. Consequently, females mating with larger males had lower lifetime fitness. The detrimental effect of male size on female longevity was not mediated by male effects on female fecundity, egg-production rate or female-remating behaviour. Similarly, the influence of male size on female lifetime fecundity was independent of the male-size effect on female longevity. There was no relationship between female size and female resistance to male harm. Thus, although increasing male body size is known to enhance male mating success, it has a detrimental effect on the direct fitness of their mates. Our results indicate that this harm is a pleiotropic effect of some other selected function and not an adaptation. To the extent that females prefer to mate with larger males, this choice is harmful, a pattern that is consistent with the theory of sexually antagonistic coevolution.     

The repeatability of mating failure in a polyandrous bug

Mating failure, characterized by the lack of production of offspring following copulation, is relatively common across taxa yet is little understood. It is unclear whether mating failures are stochastic occurrences between incompatible mating partners or represent a persistent, meaningful phenotype on the part of one or other sex. Here we test this in the seed bug Lygaeus simulans, by sequentially mating families of males with randomly allocated unrelated females and calculating the repeatability of mating outcome for each individual male and family. Mating outcome is significantly repeatable within individual males but not across full‐sib brothers. We conclude that mating failure represents a consistent male‐associated phenotype with low heritability in this species, affected by as yet undetermined environmental influences on males.     

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.     

Influence of Adult Size on Mate Choice in the Solitary and Gregarious Parasitoids, Cotesia marginiventris and Cotesia flavipes

Female and male mate choice in relation to adult size were examined for the solitary and gregarious parasitoids, Cotesia marginiventris (Cresson) and Cotesia flavipes Cameron, respectively. In addition, male precopulatory behaviors were observed for evidence of male competition or a large-male advantage in mate acquisition. Male parasitoids are not known to offer female mates direct benefits, but females that mate high quality males may obtain indirect benefits, such as offspring that are more successful in obtaining mates. Female choice experiments for C. marginiventris found that large males approached females first more frequently than small males, and that females mated large males significantly more often than small males. Male choice experiments for C. marginiventris did not demonstrate a male preference for female size. In contrast, female choice experiments with C. flavipes found that females mated equally with large or small males, while male choice experiments showed that males attempted copulation and mated more frequently with smaller females. Male competition was not observed in the gregarious species C. flavipes, but competition in this gregarious parasitoid could be moderated by dispersal.     

BENEFITS OF MULTIPLE MATES IN THE CRICKET GRYLLUS BIMACULATUS

Despite the importance of polyandry for sexual selection, the reasons why females frequently mate with several males remain poorly understood. A number of genetic benefits have been proposed, based on the idea that by taking multiple mates, females increase the likelihood that their offspring will be sired by genetically more compatible or superior males. If certain males have intrinsically “good genes,” any female mating with them will produce superior offspring. Alternatively, if some males have genetic elements that are incompatible with a particular female, then she may benefit from polyandry if the sperm of such males are less likely to fertilize her eggs. We examined these hypotheses in the field cricket Gryllus bimaculatus (Orthoptera: Gryllidae). By allocating females identical numbers of matings but different numbers of mates we investigated the influence of number of mates on female fecundity, and both short- and long-term offspring fitness. This revealed no effect of number of mates on number of eggs laid. However, hatching success of eggs increased with number of mates. This effect could not be attributed to nongenetic effects such as the possibility that polyandry reduces variance in the quantity or fertilizing ability of sperm females receive, because a control group receiving half the number of copulations showed no drop in hatching success. Offspring did not differ in survival, adult mass, size, or development time with treatment. When males were mated to several different females there were no repeatable differences between individual males in the hatching success of their mate's eggs. This suggests that improved hatching success of polyandrous females is not due to certain males having genes that improve egg viability regardless of their mate. Instead, our results support the hypothesis that certain males are genetically more compatible with certain females, and that this drives polyandry through differential fertilization success of sperm from more compatible males.     

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.     

Long-term study of female multiple mating indicates direct benefits in Tribolium castaneum

Polyandry or female mating with several different partners in a single fertile period is a widespread phenomenon possibly involving both costs and benefits. This study tested whether remating after weeks of initial copulation (periodic multiple mating) has fitness consequences for females of red flour beetle, Tribolium castaneum (Herbst) (Coleoptera: Tenebrionidae), a cosmopolitan storage pest. We hypothesize that females benefit from higher mating frequency and more mates through sperm replenishment and/or compatible sperm. Thus, offspring production and survivorship were examined of females that were mated to multiple males or the same male repeatedly at variable intervals (every 2 weeks, 1, 3, and 5 months). Our results suggest that remating, after months of initial copulation, confers direct benefits to females, likely by providing additional sperm or through an alternative mechanism such as better ability of fresh sperm to fertilize eggs, stimulation of oviposition from copulation itself, and/or hydration benefit of the ejaculate. We did not detect any additional benefit of female multiple mating.     

Reproductive strategies of the larval parasitoid Microplitis croceipes

Mate choice may have important consequences for offspring sex ratio and fitness of haplodiploid insects. Mate preference of females of the solitary larval parasitoid Microplitis croceipes (Cresson) (Hymenoptera: Braconidae) for virgin and mated males, and vice versa, and the reproductive consequences (i.e., the sex ratio expressed as the proportion of male offspring) were examined in choice and non‐choice experiments. In addition, the effect of repeated rapid and daily copulation of an individual male on the sex ratio of offspring of the female mates was assessed. Males preferred virgins over mated females, whereas females copulated with a male irrespective of his mating status. In both the rapid and daily copulation assay, females copulating with a male that had copulated five times or more produced a higher sex ratio than females that had copulated with a virgin male. Females that copulated with virgin males once or twice produced a significantly and considerably lower sex ratio than females that first copulated with a sperm‐depleted male followed by a virgin male. This indicates that copulating with a sperm‐depleted male has costs and limits acquisition by the female of sperm from virgin males.     

Effects of parasitic infection on mate sampling by female wild turkeys (Meleagris gallopavo): should infected females be more or less choosy?

Investigations of parasite-mediated sexual selection have concentratedon the effects of parasites on males. Differences in femalesusceptibility to parasitic infection may also cause variationin reproductive behavior. I propose two alternative hypothesesto explain how infected females may alter their mate samplingbehavior. In the first hypothesis, infected females sample fewerprospective mates because chronic parasitic infection imposesenergetic costs that limit the time and calories that a femalecan expend in mate searching. A novel alternative hypothesisis that females recognize their own susceptibility to infectionand thus invest more time searching for a male phenotype thatindicates he offers genes complementary to her genome. In recombination,these good genes would allow her offspring to better resistparasites despite their mother's susceptibility. I examinedthe mate sampling behavior of experimentally infected wild turkeyhens when presented with an array of males, and compared themto control hens. Infected females did not invest more time assessingindividuals, did not wait longer to choose a male, nor werethey less likely to solicit during the trial. They did differfrom control females in that they visited more males beforesoliciting copulation and exhibited different preference functionsfor snood length. These results suggest that females are notso energetically restricted by latent coccidia infection thatthey must hurry to find a mate. Instead, it appears that infectedfemales assess a larger set of males as prospective mates, perhapsto increase the opportunity to obtain complementary genes forparasite resistance.     

Consequences of sperm displacement for female dung flies,Scatophaga stercoraria

Displacement of stored sperm during copulation occurs in many insects. This process provides direct benefits for males via increased fertilization success, but the fitness consequences of sperm displacement for females are less clear. Here we investigate potential benefits of sperm displacement for female yellow dung flies, Scatophaga stercoraria. We find no evidence that female dung flies gain direct benefits from displacement of previously stored sperm in terms of increased fertility or fecundity. There was no difference in the relative survival rate, development time, size or fluctuating asymmetry of offspring produced by females that had previously stored sperm displaced before oviposition and those that did not. Females using previously stored sperm to fertilize their eggs produced significantly higher ratios of male to female offspring. These novel findings have important implications for understanding the evolutionary dynamics of male–female interactions in sperm competition.     

Polyandry in grain beetles, Tenebrio molitor, leads to greater reproductive success: material or genetic benefits?

Females that mate with more than one male may derive both materialand genetic benefits, and differentiating between the two benefitsis often difficult. We tested for both material and geneticeffects associated with multiple mating in the highly promiscuousyellow mealworm beetle, Tenebrio molitor. Females that matedfour times to the same male laid more eggs and produced morelarvae than females that mated only once. Whether copulationsoccurred on the same day or over several days, the result wasan immediate increase in the production of eggs by females.Some females were kept on a restricted diet to test whethernutrients in the spermatophore disproportionately benefittedfood-deprived females. Although females on poor diets producedfewer and smaller offspring, diet did not significantly affect the proportional benefit of mating treatment on female fecundity.By controlling for male mating history, we were able to separatethe effects of mating with different males from the effectsof receiving multiple spermatophores from the same male. Femalesthat mated with four different males achieved substantial gainsin numbers of eggs produced (32% increase) beyond those offemales that mated an identical number of times with the same male. We found no evidence that males allocate fewer sperm toprevious mates. Egg hatchability was unaffected by mating behavior,suggesting that genetic incompatibility at that stage is notresponsible for the low reproductive success of females matedwith a single male. These results suggest that females maydelay or reduce oviposition or may be incapable of achieving maximal fecundity until they have gained the material and/orgenetic benefits of mating with multiple males.     

Effects of female and male size on female mating and remating decisions in a bean beetle

The body sizes of individuals of the choosing and chosen sexes in a mate choice may affect sequential mating of females. We examined the effects of the body sizes of females and their mates on attributes of female first mating, and the effects of body sizes of females and their previous and potential future mates on female remating in the adzuki bean beetle, Callosobruchus chinensis. Large- and small-sized adults were derived from larvae reared under conditions of low and high density in a bean, respectively. The speed of first mating of large females was not affected by the size of courting males, whereas small females initiated mating more rapidly when courted by small males. The remating probability of large females was not affected by first male size, whereas small females that mated first with smaller males were more likely to remate. These data suggest that pre- and post-copulatory female choices for male size depend on the female’s size, and the small females might be more willing to copulate with smaller males but prefer larger males to sire their offspring after copulation. A possible explanation for this preference is that small females may suffer greater harm from copulating with larger males.     

Mating with large males decreases the immune defence of females in Drosophila melanogaster

Mating has been widely reported to be a costly event for females. Studies indicate that female cost of mating in terms of fecundity and survivorship can be affected by their mates, leading to antagonistic coevolution between the sexes. However, as of now, there is no evidence that the female cost of mating in terms of immune defence is affected by their mates. We assess the effect of different sized males on antibacterial immune defence and reproductive fitness of their mates. We used a large outbred population of Drososphila melanogaster as the host and Serratia marcescens as the pathogen. We generated three different male phenotypes: small, medium and large, by manipulating larval densities. Compared to females mating with small males, those mating with large males had higher bacterial loads and lower fecundity. There was no significant effect of male phenotype on the fraction of females mated or copulation duration (an indicator of ejaculate investment). Thus, our study is the first clear demonstration that male phenotype can affect the cost of mating to females in terms of their antibacterial immune defence. Mating with large males imposes an additional cost of mating to females in terms of reduced immune defence. The observed results are very likely due to qualitative/quantitative differences in the ejaculates of the three different types of males. If the phenotypic variation that we observed in males in our study is mirrored by genetic variation, then, it can potentially lead to antagonistic coevolution of the sexes over immune defence.     

Effects of mating patterns on reproductive performance and offspring fitness in Cryptolaemus montrouzieri

We examined the female mating choice of Cryptolaemus montrouzieri Mulsant (Coleoptera: Coccinellidae) when exposed to a male with which she had mated before vs. a novel male which she never encountered. After first mating, most females chose to remate with the same male and showed a shorter mating latency period to copulation with the familiar male as compared to a new male. Then, we allowed a female to mate once, 5× with the same, or 5× with different males to evaluate how different mating patterns influence the females’ reproductive performance and offspring fitness. Mating 5× with the same male or different males improved female egg production, but had no effect on hatching rate or offspring fitness. Our findings suggest that female C. montrouzieri are able to distinguish between familiar and unfamiliar males and prefer to mate with the former. Further, multiple mating appears to offer material benefits to female C. montrouzieri specifically by increasing fecundity.