Swarm Site Fidelity in the Sex Role-Reversed Dance Fly Empis borealis

In the dance fly species Empis borealis (L.), females (1–40) gather to swarm at landmarks (swarm markers, like trees and bushes), and males carrying an insect prey visit these swarms for mating. We noticed earlier that some swarm sites were used for several years and that they appeared to be frequented by a similar number of swarming females in each year, although the numbers of females varied greatly among swarm sites and certain sites attracted more swarming individuals than others. To explore swarm site fidelity in this mating system, in 1993 we monitored the same swarm sites that we studied in 1989, addressing the questions, Would the same swarm sites still attract the same number of females and males after 4 years? and Why do some swarm sites attract more displaying females than others? The number of females swarming at the different markers in 1993 was approximately the same as 4 years earlier. Some of these swarm sites are known to have been used for 18 years. The swarm sites with the largest number of flies had a high sun exposure during the day and were found at coniferous swarm marker trees and in a mixed forest habitat. A swarm site with few females attending and with a low amount of insolation during the day can be predicted to be abandoned as a swarming site soon. Empis borealis swarm sites thus persist over many years and are attended by a similar number of individuals each year. To our knowledge, such site fidelity has not been demonstrated for any swarming insect species earlier.     

The effects of copulation duration in the bruchid beetle Callosobruchus maculatus

Control over copulation duration is a potentially importantgenerator of sexual conflict that has received little empiricalattention. The copulatory behavior of the bruchid beetle Callosobruchusmaculatus may reflect a sexual conflict over copulation duration.Males have spines on their intromittent organs that puncturethe female reproductive tract, and females kick their matesduring copulation. If females are prevented from kicking, copulationslast longer and the injuries females sustain are more severe.Males supposedly use the spines as anchors to prolong copulationduration, and females kick to terminate copulations. We manipulatedcopulation duration experimentally and quantified its effectson male and female fitness components to test whether or notthere is a conflict over copulation duration in C. maculatus.Females did not suffer from long copulations but instead experiencedincreased lifetime fecundity. Ejaculate size increased withcopulation duration, and females apparently derive materialbenefits from the ejaculates. Males that mated first and hadlong copulations were relatively unsuccessful when competingwith sperm from other males. However, there was a trend forfemale remating propensity to decrease with long copulationdurations, and first males may therefore also benefit from longcopulations. The copulation duration of the second male to matedid not have a significant effect on sperm precedence. We concludethat even though it seems likely that the male spines have evolvedto act as an anchor during copulation, there seems to be littleconflict over copulation duration per se in C. maculatus.     

Sex‐specific repeatabilities and effects of relatedness and mating status on copulation duration in an acridid grasshopper

In species with direct sperm transfer, copulation duration is a crucial trait that may affect male and female reproductive success and that may vary with the quality of the mating partner. Furthermore, traits such as copulation duration represent the outcome of behavioral interactions between the sexes, for which it is important—but often difficult—to determine which sex is in phenotypic control. Using a double‐mating protocol, we compared copulation durations between (1) virgin and nonvirgin and (2) sibling and nonsibling mating pairs in rufous grasshoppers Gomphocerippus rufus. Nonvirgin copulations took on average approximately 30% longer than virgin copulations, whereas relatedness of mating partners was not a significant predictor of copulation duration. Longer nonvirgin copulations may represent a male adaptation to sperm competition if longer copulations allow more sperm to be transferred or function as postinsemination mate guarding. The absence of differences between pairs with different degrees of relatedness suggests no precopulatory or preinsemination inbreeding avoidance mechanism has evolved in this species, perhaps because there is no inbreeding depression in this species, or because inbreeding avoidance occurs after copulation. Controlling for the effects of male and female mating status (virgin vs. nonvirgin) and relatedness (sibling vs. nonsibling), we found significant repeatabilities (R) in copulation duration for males (R = 0.33; 95% CI: 0.09–0.55) but not for females (R = 0.09; 95% CI: 0.00–0.30). Thus, copulation durations of males more strongly represent a nontransient trait expressed in a consistent manner with different mating partners, suggesting that some aspect of the male phenotype may determine copulation duration in this species. However, overlapping confidence intervals for our sex‐specific repeatability estimates indicate that higher sampling effort is required for conclusive evidence.     

Determinants of sperm transfer in the scorpionfly Panorpa cognate: male variation, female condition and copulation duration

Recent studies suggest that sperm production and transfer may have significant costs to males. Male sperm investment into a current copulation may therefore influence resources available for future matings, which selects for male strategic mating investment. In addition, females may also benefit from actively or passively altering the number of sperm transferred by males. In the scorpionfly Panorpa cognata, the number of sperm transferred during copulation depended on copulation duration and males in good condition (residual weight) copulated longer and also transferred more sperm. Moreover, sperm transferred and stored per unit time was higher in copulations with females in good condition than in copulations with females in poor condition. Males varied greatly and consistently in their sperm transfer rate, indicative of costs associated with this trait. The duration of the pairing prelude also varied between males and correlated negatively with the male's sperm transfer rate, but no other male character correlated significantly with male sperm transfer rate. The results are consistent with strategic mating effort but sperm transfer could also be facilitated by the physical size of females and/or females in good condition may be more cooperative during sperm transfer.     

Males do not prolong copulation in response to competitor males in the polyandrous fly Drosophila bifasciata

Males in many taxa exhibit behavioural plasticity in response to the perceived threat of sperm competition. Drosophila males prolong mating in response to the presence of competitor males before copulation. The benefits of this behaviour to males are evident in Drosophila melanogaster. However, the adaptive nature of the trait is challenged by the observation that it is present in four other Drosophila species, two of which are largely monandrous, raising the possibility that this plasticity is not evolutionarily labile. In the present study, behavioural plasticity and the mating system in Drosophila bifasciata Pominini (Diptera, Drosophilidae) are analyzed. By contrast to other Drosophila species, male D. bifasciata do not exhibit plasticity in copulation duration when competitor males are present before mating. Thus, plasticity in mating duration is not fixed in the genus Drosophila. The mating system of D. bifasciata is also examined. The species is polyandrous but, uncommonly for the genus Drosophila, males transfer a mating plug composed of sperm to females, which forms very shortly after copulation and fills the female uterus. The absence of plasticity observed in D. bifasciata may arise from the sperm plug.     

Why it is difficult to model sperm displacement in Drosophila melanogaster: the relation between sperm transfer and copulation duration

Abstract.— We have investigated the effects of experimental manipulation of copulation duration on sperm displacement in Drosophila melanogaster . Both spermless and normal males were used as second (displacing) males in the experiments. Displacement induced in the absence of sperm, that is, by males that pass accessory gland fluid alone, was a relatively inefficient process and produced much lower levels of displacement than normal males. Therefore, the presence of second-male sperm is necessary (but unlikely sufficient) for the high levels of displacement commonly observed in D. melanogaster . Furthermore, when second matings were interrupted at various times after the initiation of copulation, the distribution of displacement was strongly bimodal. We conclude that sperm transfer is relatively rapid, beginning shortly after the initiation of copulation, and is essentially complete before the midpoint of copulation. Therefore, sperm transfer bears no simple relation to copulation duration. Because it would be difficult to manipulate the numbers of sperm transferred by manipulating copulation duration, methods used to study sperm displacement in other insect species are unlikely to be appropriate for D. melanogaster . We also investigated why males mate for more than twice the duration that appears to be necessary to complete sperm transfer. Experimental interruption of first matings indicated that the extra copulation time serves to delay female remating, rather than to increase that rate at which of offspring are sired before remating.     

Nuptial gift in the spider Pisaura mirabilis maintained by sexual selection

The nuptial prey gift in the spider Pisaura mirabilis has been suggested to function as a male protection against sexual cannibalismduring courtship and mating. This hypothesis together withtwo alternatives—male mating effort and paternal investment hypotheses—were tested in a laboratory experiment withsexually inexperienced males and females. One group of malesoffered no gift to the female while three groups of males offeredsmall, medium, or large sized gifts, respectively. No malewas cannibalized among 82 trials. Aggression was observed onlyin encounters where a gift was presented. Males without a gift courted females, and 40% of these males managed to copulate,compared to 90% of males offering a gift. The copulation durationwas positively correlated with gift size. In general, the femaleterminated the copulation and ran away with the gift. The proportionof eggs fertilized increased with copulation time. Presenceor size of the nuptial gift did not affect female fecundityor spiderling size significantly. The results refute the hypothesesof sexual cannibalism and paternal investment. The nuptialgift represents a male mating effort; it entices the femaleto copulate, facilitates coupling during copulation, and byprolonging copulation it may increase the amount of sperm transferred.I conclude that the nuptial prey gift in Pisaura mirabilisis maintained by sexual selection.     

Adjustment of copula duration and ejaculate size according to the risk of sperm competition in the golden egg bug (Phyllomorpha laciniata)

Several hypotheses have been proposed to explain the adaptivesignificance of prolonged copulations in insects, which includemate guarding and sperm loading functions. We have exploredthe adaptive significance of the prolonged copulations in thegolden egg bug (copulations up to 50 h) and the effect of anincreased risk of sperm competition on ejaculate investment.Our data support predictions derived from sperm competitiontheory, which posits that males are expected to increase ejaculateexpenditure in response to an increased risk of sperm competition.Results show a combined response by males that has not beenpreviously described: males in the presence of rivals increasecopulation duration and the rate of sperm transfer. No relationshipwas found between male or female size and copulation durationor ejaculate size. Golden egg bug males transfer sperm slowlyand gradually throughout copulation; thus an increase in theamount of sperm transferred and the corresponding increase inthe male's numerical representation in the female's storageorgans could be particularly important in a system in whichso few sperm are transferred and in which so few sperm are storedby females. In addition, copulation duration may not only serveto increase the total amount of sperm transferred, but it mayalso increase the chances that the female will lay an egg soonafter copulation has ended. This could explain why males tendto accept eggs after copulation, since they could be maximizingthe chances that such eggs are fathered by them, and in thisway they would substantially increase the survival rates oftheir offspring because eggs laid on plants suffer high mortalityrates.     

The effect of female mating history on sperm precedence in the two-spot ladybird, Adalia bipunctata (Coleoptera, Coccinellidae)

Effects of two different mating regimes on sperm precedencein the two-spot ladybird, Adalia bipunctata, were studied usingthe polymorphic gene for melanism as a marker for paternity.Virgin nonmelanic females (homozygous recessive) were matedto nonmelanic male(s) and then, after laying fertilized eggs,were mated to a melanic male of known genotype. The resultsafter the two successive single matings showed a highly variabledegree of paternity of the second male. Initial multiple matingwith nonmelanic males did not alter the pattern of paternityafter the subsequent single mating with a melanic male, butit had two other effects: (1) the female showed an increasein rejection behavior, and (2) a longer copulation was requiredfor high success of the melanic male. Additional observationsin which families were reared from beetles collected in copulain the field demonstrated that sperm competition also occursunder natural conditions. The outcome of the competition wasvariable with frequent sperm mixing.     

Adaptive control of copulation duration by males under sperm competition in the mite,Macrocheles muscaedomesticae

In a manure-inhabiting predatory mite, Macrocheles muscaedomesticae (Gamasida, Macrochelidae), when the female mates with two males, the first male takes nearly perfect fertilization priority (Yasui, 1988). The present study examined whether the first-male's sperm precedence is influenced by the copula-duration of the first and second males mating with the same female, and whether males control their copulation duration by assessing the probability that the mate has been inseminated by other males. Results of the artificial interruption of copulation showed that sperm precedence value, P₂ (the proportion of the offspring fathered by the second male), was negatively correlated with the copulation duration of the first male but positively correlated with that of the second male. There was a threshold (ca. 180–300 seconds) in the first-male's copulation duration beyond which P₂ decreased drastically; when length of the first copulation exceeded this threshold, the second males did not fertilize eggs, whereas they fertilized more than half of the eggs when the first-copulation duration was shorter than the threshold. Almost all males copulated for a longer period (average 509.8 seconds) than this threshold if the copulation duration of the previous male had not exceeded the threshold, but if it was longer than the threshold, second males had shortened their copulation (67.6 seconds). These results suggest that males are able to assess the insemination status of their mates and to adjust their copulation duration depending on the probability of fertilizing eggs by their own sperm. A mechanistic explanation for sperm precedence (i.e., plug-formation within sperm receptive organ of the females) is proposed.     

Numbers and size of sperm storage tubules and the duration of sperm storage in birds: a comparative study

Estimates of the numbers of sperm storage tubules (SSTs) in the utero-vaginal junction of 11 bird species are presented. Numbers of SSTs varied by a factor of 40 between species, and ranged from 500 to 20000. Body mass accounted for over 50% of the variation in SST mumbers. SST length was positively correlated with the length of spermatozoa across species. The duration of sperm storage was not correlated with the number of SSTs or the volume of sperm storage tissue. However, the number of 'active' SSTs appears to vary between species and it was not possible to make allowance for this. Sperm storage duration was weakly, positively correlated wth clutch size, but showed a significant positive relationship with the number of days over which laying occurred. The number of SSTs was also positively correlated with the number of sperm per ejaculate. The best predictor of sperm storage duration was a multiple regression equation using the spread of laying and the length of sperm storage tubules. The duration of sperm storage in birds which remain together during the pre-laying period is such that a single insemination immediately before the start of laying could fertilize the entire clutch.     

Decoy presentations as a means to manipulate the risk of extrapair copulation: an experimental study in a semicolonial raptor, the Montagu's harrier (Circus pygargus)

Mate guarding and frequent copulations are two alternative paternity assurance strategies found in birds. In species with intensecourtship feeding, like raptors, the "frequent copulation"strategy is expected because male food provisioning conflictswith mate guarding. We evaluated experimentally the paternityassurance behavior of a semicolonial raptor, the Montagu'sharrier Circus pygargus, using decoy presentations to simulateterritorial intrusions. Breeding pairs were exposed to maleand female decoys at different periods during the female's reproductive cycle. Agonistic responses to decoys were intra-sexual,and the timing and intensity of male attacks toward male decoyssupported responses related to the risk of extrapair copulation(EPC): Male aggression peaked during the presumed fertile periodand almost disappeared after clutch completion. During thefertile period, copulation rate was significantly higher, andcopulations lasted longer, during male decoy presentations than during controls. Males also spent more time close to the femaleduring male decoy presentations compared to controls, bothduring the early prelaying and fertile periods, but not duringincubation. In the fertile period, males also increased presencetime close to the female in the hour following the removal of the male decoy. Conversely, female decoy presentations hadno significant effect on copulatory behavior or male presencetime. These results showed that the risk of EPC can be experimentallymanipulated by the means of decoy presentations, simulatingmale territorial intrusions, and that male Montagu's harriersincrease their short-term copulation frequency and female surveillancewhen they perceive themselves at an increased EPC risk.     

Repeated copulations as a strategy to maximize fertilization in the fly, Dryomyza anilis (Dryomyzidae)

In the fly, Dryomyza anilis, males copulate repeatedly withthe same female during oviposition. Each copulation bout consistsof intromission followed by several tapping sequences duringwhich the male touches the external genitalia of the femalewith his claspers. The relative fertilization success of thelast male to mate with the female increases with the numberof tapping sequences. In this study, male benefits of repeatedcopulation bouts were examined by comparing matings with thesame number of tapping sequences in one and several copulationbouts. The relative fertilization success for the last maleincreased with successive copulation bouts. Simulated ovipositionpatterns showed that fertilization success in matings with manycopulation bouts was higher than in matings with one bout onlyif eggs were distributed between bouts in a certain way. Thesepredicted oviposition patterns were compared with natural ones.Although males may benefit from repeated copulation, femalesare likely to prefer matings with quick oviposition. This intersexualconflict may also affect natural oviposition patterns. It issuggested that repeated copulation in D. anilis may have evolvedin connection with males trying to secure their paternity becausefemales can discharge sperm at any moment during mating.     

Postmating sexual selection: the effects of male body size and recovery period on paternity and egg production rate in a water strider

The role of male body size in postmating sexual selection wasexplored in a semiaquatic insect, the water strider Gerris lateralis.To separate effects of male size per se from those due to numericsperm competition, male recovery period (shown here to be proportionalto ejaculate size) was manipulated independently of body sizein a factorial experiment where virgin females were mated firstwith sterile males and then with focal males. Both relativemale fertilization success and female reproductive rate were measured.The number of sperm transferred increased with male recoveryperiod, an effect that was mediated by longer copulation duration,but there were no effects of body size on ejaculate size. Neithermale size nor recovery period had any significant direct effectson male fertilization success. However, copulation durationinfluenced relative fertilization success, suggesting that malesable to transfer more sperm also achieved higher fertilizationsuccess. Females exercised cryptic female choice by modulatingtheir reproductive rate in a manner favoring large males andmales that were successful in terms of achieving high relativefertilization success. Thus, successful males gained a twofoldadvantage in postmating sexual selection. This study has important implicationsfor previous estimates of sexual selection in this group of insectsbecause pre- and postmating sexual selection will be antagonisticdue to limitations in male sperm production: males mating frequently(high mating success) will on average transfer fewer sperm ineach mating and will hence tend to fertilize fewer eggs permating (low fertilization success).     

Sexual conflict over copula timing: a mathematical model and a test in the yellow dung fly

Sexual conflict over mating occurrence, timing, or durationis common in animals. This explains conspicuous female materejection behavior in many species, often involving shaking,fighting, and occasional forced copulations. We present a simplemodel that generates predictions about whether and when copulationoccurs in such conflict situations and how much female rejectionbehavior should be observed. Predictions depend on 2 underlyingparameters affecting female resistance and male persistence.We supply 2 qualitative tests of the model using the yellowdung fly Scathophaga stercoraria (Diptera: Scathophagidae).We manipulated adult age, body size (large and small), and adultfood availability (low and high), independently in males andfemales, staging replicate pairings of all treatment combinations.In agreement with predictions of our model, shaking durationfirst increased to a maximum at intermediate age, when the averagefemale copulated, and then decreased again. Contrary to expectation,body size did not affect copulation timing, female resistance,or male persistence. As predicted, adult food limitation delayedsexual maturity and hence prolonged female resistance, resultingin later copulations after more shaking. However, although foodlimitation equally delayed the increase in male persistencewith age, copulation also occurred later after more shaking,opposite to the model prediction. We conclude that shaking isdriven primarily by female age and male responses to it. Althoughfemale shaking can initially successfully deter males in S.stercoraria, this behavior is subtle and has apparently shiftedfunction from an effective means of mate choice to a signalof nonreceptivity, though its importance in nature remains unclear.     

Correlates of the duration of the egg collecting phase in the three-spined stickleback

Individual male three-spined sticklebacks in the field, often collected eggs for longer (up to 10 days) than had been reported for sticklebacks in captivity (3–6 days). The probability that a male stopped collecting eggs increased with the number of eggs already present, and possibly with the age of the eggs. Males with nests hidden under a plant were more likely to continue collecting than males with exposed nests. These results are discussed in the light of theoretical considerations that predict when males should stop collecting further eggs.     

Functional shifts in the use of parasitized hosts by a tephritid fly: the role of host quality

Superparasitism, a phenomenon in which parasitic insects layeggs in already-exploited hosts, provides a useful context inwhich to examine the dynamics of parental investment. This studyexplored conditions under which female Mediterranean fruit flies(Ceratitis capitata) shift from avoiding superparasitism ofhost fruit to preferring it, even placing eggs direcdy intoexisting egg-laying cavities. An a priori hypothesis of costsand benefits was used to predict how use and avoidance of parasitizedfruit would change in -response to changes in fruit size andripeness. We predicted that avoidance would decrease with increasingfruit size, while use would increase with decreasing ripeness.Using a field-cage assay, ripeness was held constant and thesize of host coffee berries manipulated. Avoidance of parasitizedberries was significandy less pronounced on large berries thanon small ones. In a second experiment, size was held constantand ripeness manipulated. On unripe berries, females depositedthe majority of clutches directly into existing egg-laying cavities.On ripe berries, by contrast, the same females deposited mostclutches in previously unparasitized fruit. Parallel patternsin the frequency of female-female contests were observed, supportingthe notion that a fruit's value is determined by an interactionbetween fruit size or ripeness, on one hand, and the prior occurrenceof eggs, on the other. Laboratory assays suggested that useof existing sites had advantages in terms of time savings; femalebehavior thus constitutes a relatively uncommon example of adaptivesuperparasitism in which parasitized hosts are actually preferredover unparasitized ones.[Behav Ecol 7: 235-242 (1996)]     

Nest size affects clutch size and the start of incubation in magpies: an experimental study

Nest size has been suggested to be a sexually selected traitindicating parental ability of both males and females. To testwhether a female's reproductive decisions (e.g., clutch sizeand starting incubation) change in relation to experimentalmanipulation of nest size, as would be predicted if nest sizeis a sexually selected signal reflecting the male's parental quality, we manipulated nest size in a population of monogamousmagpies before laying by adding or removing about 20 cm oflarge sticks in the roof of magpie nests. On the one hand,we found that clutch size of reduced nests was smaller thanthat of control or enlarged nests. Moreover, clutch size was significantly related to nest size after manipulation, whichindicates that females adjust clutch size to the final sizeof the nest, nest size thereby being a good candidate for asexually selected trait. On the other hand, number of eggshatched during the first day is hypothesized to be related to the expected available resources during nestling growth, andsubsequent nestlings hatched are likely to die due to broodreduction if resources are not sufficient to raise well-developednestlings. Nest size is hypothesized to inform females abouta male's willingness to invest in reproduction, and we foundthat in broods of experimentally reduced nests, females startedto incubate earlier in the laying sequence than they did inbroods of control or enlarged nests. Moreover, in experimentallyreduced nests, fewer nestlings hatched during the first day,and the difference in body mass between the first and the fourthnestling hatched increased. This result is in accordance withthe hypothesis that the female's decision of when to start incubationin the laying sequence is mediated by nest size, a sexuallyselected trait signaling parental quality. We discuss alternativeexplanations for the results such as the possibility that nestsof different treatments may differ in their thermoregulationproperties or in their protection against predators.