Influence of previous mating experience on future mating success in maleLucilia cuprina (Diptera: Calliphoridae)

Receptive and virginL. cuprina females were placed with a virgin male and an experienced male that had mated between one and five times previously. The experienced males secured significantly more matings than virgin males. Males with previous matings also gained experience in competing with males. Males directed mating attempts at each other, seemingly in the context of intrasexual competition. Experienced males directed more mating attempts at virgin males than vice versa. As their number of previous matings increased, experienced males made the first mating attempt at females more often and directed more mating attempts at females compared with virgin males. Females did not actively discriminate against experienced males, even though the proportion of matings secured on the first attempt by experienced males declined with increasing mating experience. Alternative behavioral explanations are discussed.     

Males mate with females even after sperm depletion in the two-spotted spider mite

Generally, males increase their reproductive success by mating with as many females as possible, whereas females increase their reproductive success by choosing males who provide more direct and indirect benefits. The difference in reproductive strategy between the sexes creates intense competition among males for access to females, therefore males spend much energy and time for competition with rival males for their reproduction. However, if they do not need to engage themselves into male competition and females are in no short supply, how many females can a male mate with and fertilize? We address this question in the two-spotted spider mite, Tetranychus urticae Koch. In this study, we investigated how many females a young, virgin male mated in 3 h, and checked whether the mated females were fertilized. We found that on average males mated with 12–13 females (range: 5–25). As latency to next mating did not change with the number of matings, the males are predicted to engage in even more matings if the mating trial were continued beyond 3 h. Copulation durations decreased with the number of matings and typically after 11 copulations with females any further copulations did not lead to fertilization, suggesting that males continued to mate with females even after sperm depletion. We discuss why spider mite males continue to display mating and copulation behaviour even after their sperm is depleted.

     

Size-assortative mating, male choice and female choice in the curculionid beetle Diaprepes abbreviatus

In the beetle Diaprepes abbreviatus (L.) females are larger on average than males, as indicated by elytra length. Size-assortative matings were observed in wild populations in Florida and in laboratory mating experiments. We tested three mechanisms for this size-assortative mating: (1) mate availability; (2) mating constraints; and (3) mate choice. We found that mate choice influenced size-assortative mating by: (1) large and small males preferring to mate with large females; (2) large males successfully competing for large females, leaving small males to mate with small females; and (3) females accepting large males as mates more readily than small males. Males increased their reproductive success by mating with larger, more fecund females. They transferred protein to females during mating. Copyright 1999 The Association for the Study of Animal Behaviour.     

The mating-induced refractoriness of Lucilia cuprina females: manipulating the male contribution

Abstract Three manipulations of the male contribution to copulation were used to investigate what determines the switch-off of female receptivity that follows mating in Lucilia cuprina. Multiple matings by males led to a reduction in their effectiveness at switching-off females: the first twelve matings caused more than 50% of females to be switched-off 1 day after mating, while the first four matings caused more than 50% of females to be switched-off for 7 days after mating. Males mated up to twenty-three times and on average ten times during 10 h of continuous access to virgin females. The number of sperm transferred declined logarithmically through a series of matings as did the quantity of material transferred from the male accessory gland (measured as radiolabeled material) through the first six matings. Overnight isolation of multiply-mated males led to considerable recovery of their ability to switch-off females, and to limited renewal of their ability to transfer sperm. Castrated males transferred similar quantities of accessory gland material to females as did sham-operated males during their first five matings and in their first three matings switched-off 95% of females for 1 day but only 48% for 7 days. When normal mating pairs were separated at increasing intervals after coupling, an increasing proportion of females were switched-off at 1 or 7 days afterward. 7 days after 2 min matings no females were switched-off though 40% of the number of sperm transferred in a full mating had been transferred by this time. The proposed explanation for these data is that both the initial switch-off and its duration are determined by the quantity of a receptivity-inhibiting substance that normally enters the female haemo-lymph after being injected by the male into the wall of the bursa copulatrix. It is proposed that when castrated males mate, an absence of sperm results in most accessory gland secretion entering the empty spermathecae (rather than the wall of the bursa as usually occurs) and hence being unable to reach the haemolymph and exert its influence. The effective dose of the receptivity-inhibiting substance is measured on a logarithmic scale.     

Mating Systems and Reproductive Strategies in Three Australian Gryllid Crickets: Bobilla victoriae Otte,Balamara gidya Otte and Teleogryllus commodus (Walker) (Orthoptera: Gryllidae: Nemobiinae; Trigonidiinae; Gryllinae)

The behaviour of B. victoriae and B. gidya in simulated natural habitats is described. Regular predictable sequences of behaviour are represented by ethograms. The mating systems of B. victoriae and B. gidya are compared with that of T. commodus which shows a sedentary call-and-reside strategy. B. victoriae shows an elaborated male call-search strategy while in B. gidya the male strategy is one purely of searching rather than advertising for mates. Males of B. victoriae and B. gidya were less territorial and less aggressive than males of T. commodus. Males of B. victoriae attracted females and monopolized them for repeated matings by providing what might be a nutritional resource in the form of multiple spermatophores which females were allowed to eat soon after copulation. Males of B. gidya produced fewer but much larger spermatophores than males of B. victoriae or T. commodus and females of B. gidya became unreceptive after mating. The reproductive strategies of the three species are analyzed in terms of environmental variables.     

Chastity belts in gartersnakes: the functional significance of mating plugs

Male red-sided gartersnakes (Tfiamnophis sirtalis parietalis) deposit a thick gelatinous plug that occludes the female cloaca after copulation. Previous workers have interpreted the plug as a sexually-selected adaptation to (1) physically prevent re-mating by the female, and/or (2) provide pheromonal cues to discourage courtship by rival males or to decrease receptivity by females. Our data support the former hypothesis, but not the latter. Plugs serve as effective physical barriers to additional copulation for <72 h, but this is long enough for most females to become unreceptive, and/or disperse from the mating aggregation. Experimental removal of plugs immediately after copulation results in some re-mating by females, but plug removal several hours later does not rekindle sexual receptivity. Contrary to previous work, our experiments show that fluids associated with copulation (rather than the plug per se) are responsible for the rapid decline of male interest in mated females. Thus, the plug's primary function is to physically prevent matings rather than as a source of pheromonal cues to manipulate the behaviour of females or rival males. Plug mass is determined not only by a male's body size, but by his prior mating history (plug mass decreases with repeated mating) and by the size of his partner (males allocate larger plugs to larger females). Gartersnakes are unusual not only in their production of mating plugs, but also in their brief duration of copulation compared to other snakes. Mating plugs may have evolved in gartersnakes to reduce mating times, because of the extremely high 'opportunity cost' of prolonged mating to a male gartersnake in a mating aggregation.     

DOES LARGE BODY SIZE IN MALES EVOLVE TO FACILITATE FORCIBLE INSEMINATION? A STUDY ON GARTER SNAKES

A trend for larger males to obtain a disproportionately high number of matings, as occurs in many animal populations, typically is attributed either to female choice or success in male-male rivalry; an alternative mechanism, that larger males are better able to coercively inseminate females, has received much less attention. For example, previous studies on garter snakes (Thamnophis sirtalis parietalis) at communal dens in Manitoba have shown that the mating benefit to larger body size in males is due to size-dependent advantages in male-male rivalry. However, this previous work ignored the possibility that larger males may obtain more matings because of male-female interactions. In staged trials within outdoor arenas, larger body size enhanced male mating success regardless of whether a rival male was present. The mechanism involved was coercion rather than female choice, because mating occurred most often (and soonest) in females that were least able to resist courtship-induced hypoxic stress. Males do physically displace rivals from optimal positions in the mating ball, and larger males are better able to resist such displacement. Nonetheless, larger body size enhances male mating success even in the absence of such male-male interactions. Thus, even in mating systems where males compete physically and where larger body size confers a significant advantage in male-male competition, the actual selective force for larger body size in males may relate to forcible insemination of unreceptive females. Experimental studies are needed to determine whether the same situation occurs in other organisms in which body-size advantages have been attributed to male-male rather than male-female interactions.     

Repeated matings offset costs of reproduction in female crickets

Summary Courtship food gifts can be a significant source of nutrition to females and costly for males to produce; hence, costs of reproduction should be reduced for multiple-mating females and increased for multiplemating males in a gift-giving species. We tested this hypothesis by experimentally manipulating mating opportunities of males and females of two cricket species,Gryllodes sigillatus, a gift-giving species andGryllus veletis, a non-gift-giving species. Females of both species consume the externally attached spermatophore after mating, but inG. sigillatus, the sperm-containing ampulla is accompanied by a large gelatinous spermatophylax. In both species, survival of mated females given limited access to males was reduced relative to virgin females, thus suggesting a cost of reproduction to females. However, females given unlimited mating opportunities lived as long as virgins and also produced significantly more offspring than limited-access females. These results suggest that benefits of repeated matings, particularly those arising through spermatophore consumption, offset costs of reproduction in females. Lack of a treatment by species interaction suggests that females of both species derive nutritional benefits through spermatophore consumption, and that any additional advantage to the consumption of the spermatophylax inG. sigillatus is offset by more frequent mating byG. veletis females. In contrast to females, varying mating opportunities had no effect on male survival, suggesting that mating effort is not very costly to males. Male survival increased linearly with body mass but only when males were food-deprived, suggesting that larger males possess greater initial energy reserves to sustain their longevity when food-stressed.     

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.     

Strategic male mating effort and cryptic male choice in a scorpionfly

In animal species with high male mating effort, males often find themselves in a dilemma: by increasing their mating effort, the gain from each copulation increases but simultaneously reduces available resources and, thus, the opportunity for future copulations. Therefore, we expect males to spend less reproductive resources on matings that provide low reproductive potential, thereby saving resources for future copulations, possibly with high-quality females, a sort of cryptic male choice. However, the strength of the trade-off between investment in a current mating and resources available for future matings must not be the same for all males. Males with relatively high mating costs should allocate their limited resources more cautiously than males with more plentiful resources. Here, we examine this prediction in the scorpionfly Panorpa cognata. Prior to copulation, males produce a large salivary mass on which females feed during copulation. We show that the production of larger salivary masses leads to longer copulations. Moreover, the size of the salivary gland and salivary mass increases with increasing male condition. However, males in poor condition make a relatively higher mating investment than males in good condition. We therefore expect male condition to influence cryptic male choice. In accordance with our hypothesis, only males in poor condition choose cryptically, producing larger salivary masses in copulations with females of high fecundity.     

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.     

Male distribution and interactions at female oviposition sites as factors affecting mating success in the flyDryomyza anilis (Dryomyzidae)

Summary Dryomyza anilis males gather around carcasses where females arrive to lay their eggs and mate with them before oviposition. Distribution and mating success of males of different size at these sites were studied. The position of males of different size on the carcass and in ten circular zones around it was recorded every 10 min during 2–3 h observation periods. The position of mating pairs, mating initiations and the number of take-overs in different zones were also recorded. It was expected that males would distribute themselves according to the ideal free distribution, i.e. their gain rate in different zones in terms of number of mating initiations would be equalized.Males in different zones reached a stable distribution after 30 min from the beginning of observations. Although males, females and mating pairs were distributed over several zones, most matings were started and take-overs took place on the carcass or in the first zone (C+Z1) because arriving females usually reached the centre before being intercepted by a male. Therefore, males in C+Z1 attained a much higher mating rate than males further away: the percentage of matings started there was significantly higher than the percentage of males present there at the same time. The frequency of male-male fights was highest in C+Z1 which is likely to prevent more males from entering the central area. In this respect the observed male distribution bears resemblance to the ideal despotic distribution.However, the distribution of males of different size did not differ, and there were many small males in C+Z1. Small males in C+Z1 had a higher likelihood of starting a mating than small males further away but suffered from frequent take-overs like small males in all zones. For example, small males started 38% of all matings but there were only 8% of them among males guarding an ovipositing female. The corresponding figures for large males were 35% and 77%, respectively, suggesting that kleptoparasitic interactions and not aggressions between males were the main reason for differences in mating success between males of different size. Since small males do not have any alternative mating sites where large males would be absent, being the second last male to mate with a female is the best small males can achieve. Large males may even benefit from the presence of small males in the centre: they intercept females which large males can then easily take over.     

Causes and correlates of loss and recovery of sexual receptivity inLucilia cuprina females after their first mating

After an initial mating, females of the Australian sheep blowfly, Lucilia cuprina(Diptera: Calliphoridae), rejected mating attempts by males for several days. Almost all (>98%) females were unreceptive 24 h after mating. When tested 9 days after mating receptivity had returned in 24% of females denied the opportunity to oviposit and 45% of females given repeated opportunities to oviposit on liver. Those mated females that regained receptivity mated as readily as virgins. Gravid first-, second-, or third-cycle females that did not oviposit when presented with liver had a higher receptivity than those that laid. These non-layers laid after remating. Injection of 1 male equivalent of an extract of male accessory reproductive glands into virgin females switched off their receptivity within 5 h for at least 8 days. An extract of testis reduced receptivity 5 h after injection but had no detectable effect 3 days after injection. Both extracts increased the tendency of virgin females to oviposit. There was no difference in the sperm stores of females that refused to lay a second egg mass and comprised a high percentage of receptive females and those given no opportunity to lay this egg mass.     

Effects of size and sex on the courting success and foraging behaviour of Embiotoca jacksoni

To test life-history theory that body size and sex should influence how animals allocate time to foraging versus reproductive activities, we measured the effects of size and sex on courting success and foraging behaviour of black surfperch Embiotoca jacksoni off Santa Catalina Island, southern California. Observations of focal fish were made while snorkelling, during which the length of each fish (estimated to the nearest cm), total duration of courting encounters and foraging rates were recorded. We made observations during and outside the mating season. Courtship occurred only between pairs and its duration increased with the size of both the male and female. Although males would court females that were smaller or larger than themselves, pairs that were closely matched in size had long courting sessions, whereas those that differed considerably in size courted only briefly. Small fish foraged more than larger fish, both during and outside the mating season. Males and females foraged at similar rates outside of the mating season, but during the mating season males reduced their foraging rates to less than half that seen outside of the mating season, whereas females continued to forage at the same rate. This decrease in foraging rate of males during the mating season was seen in all sizes of males but was proportionally greatest in the largest males. These observations indicate that males trade off time spent on foraging for time spent courting during the mating season, whereas females do not.     

Costly sexual harassment in a beetle

Abstract The optimal number of mating partners for females rarely coincides with that for males, leading to sexual conflict over mating frequency. In the bruchid beetle Callosobruchus maculatus, the fitness consequences to females of engaging in multiple copulations are complex, with studies demonstrating both costs and benefits to multiple mating. However, females kept continuously with males have a lower lifetime egg production compared with females mated only once and then isolated from males. This reduction in fitness may be a result of damage caused by male genitalia, which bear spines that puncture the female’s reproductive tract, and/or toxic elements in the ejaculate. However, male harassment rather than costs of matings themselves could also explain the results. In the present study, the fitness costs of male harassment for female C. maculatus are estimated. The natural refractory period of females immediately after their first mating is used to separate the cost of harassment from the cost of mating. Male harassment results in females laying fewer eggs and this results in a tendency to produce fewer offspring. The results are discussed in the context of mate choice and sexual selection.     

Directional selection on body size but no apparent survival cost to being large in wild New Zealand giraffe weevils

When an individual's reproductive success relies on winning fights to secure mating opportunities, bearing larger weapons is advantageous. However, sexual selection can be extremely complex, and over an animal's life the opportunity to mate is influenced by numerous factors. We studied a wild population of giraffe weevils (Lasiorhynchus barbicornis) that exhibit enormous intra and intersexual size variation. Males bear an elongated rostrum used as a weapon in fights for mating opportunities. However, small males also employ sneaking behavior as an alternative reproductive tactic. We investigated sexual selection on size by tracking individual males and females daily over two 30‐day periods to measure long‐term mating success. We also assessed how survival and recapture probabilities vary with sex and size to determine whether there might be a survival cost associated with size. We found evidence for directional selection on size through higher mating success, but no apparent survival trade‐off. Instead, larger individuals mate more often and have a higher survival probability, suggesting an accumulation of benefits to bigger individuals. Furthermore, we found evidence of size assortative mating where males appear to selectively mate with bigger females. Larger and more competitive males secure matings with larger females more frequently than smaller males, which may further increase their fitness.     

Remating, sperm transfer, and sperm displacement in the cactophilic species Drosophila buzzatii Patterson & Wheeler (Diptera: Drosophilidae)

The mating system of Drosophila buzzatii is characterized by short copulation duration, frequent remating in both males and females, and male ejaculate partitioning. Additional features of the system are strong sperm displacement and a high frequency of sterile matings. Remating frequencies and the effects of remating on various mating parameters were studied. In order to characterize variation, five isofemale lines from geographically distant localities in Australia (three localities), Brazil and the Canary Islands were used. Mating parameters studied were: premating time, copulation duration, interval between successive matings, and progeny number as a measure of sperm transfer. Variation for sperm displacement was studied in crosses between laboratory stocks and a number of isofemale lines from Australia. There were significant between‐line differences in female remating frequencies, premating time, copulation duration, interval between successive matings, and progeny numbers, indicating genetic variation for these traits. Females from the five lines mated on average 1.6 to 3.1 times in 4 h, with a maximum of eight matings for one female. The males were given a maximum of ten virgin females in sequence and more than one‐third of the males mated all ten females in the 2 h observation period. Copulation duration decreased and interval between matings increased with copulation number in multiply mated males. Mean copulation duration was c. 2 min. Sperm transfer, measured as the average number of progeny from a single mating, was low (c. 25) and multiply mated females gave more progeny than single mated females, although with much lower progeny numbers than observed in wild‐caught non‐virgin females. A surprisingly high proportion of observed matings gave no progeny, i.e. they were sterile matings. Sperm displacement was strong in most crosses and remained strong in multiply mated females. The results are discussed in relation to the evolution of mating patterns in Drosophila.     

Role of Contests in the Scramble Competition Mating System of a Leaf Beetle

Male competition for mates can occur through contests or a scramble to locate females. We examined the significance of contests for mates in the leaf beetle Chrysomela aeneicollis, which experiences a short breeding season. During peak mating season, 18–52% of beetles are found in male-female pairs, and nearly half of these are copulating. Sex ratios do not differ from parity, females are larger than males, and positive size-assortative mating occurs. Males fight (2–4% of beetles) over access to females, and disruption of mating usually follows these contests. In the laboratory, we compared mating and fighting frequencies for males found in mating pairs (field-paired) and single males placed into an arena with a field-paired female. Mating pairs were switched in half of arenas (new male-female pairs) and maintained in the other half. For 2 days, each male was free to move about and fight; thereafter males were tethered to prevent contests. Mating frequencies were significantly greater for field-paired than single males in both situations. Male size was not related to mating frequency; however, large females received more matings than small ones. These data suggest that males fight for high quality females, but otherwise search for as many matings as possible.     

First evidences of sexual selection by mate choice in marine zooplankton

Sexual selection is potentially important in marine zooplankton, presumably the most abundant metazoans on earth, but it has never been documented. We examine the conditions for sexual selection through mate choice and describe mating preferences in relation to size in a marine zooplankter, the pelagic copepod Acartia tonsa. Males produce spermatophores at a rate (~1 day⁻¹) much lower than known female encounter rates for most of the year and the decision to mate a particular female thus implies lost future opportunities. Female egg production increases with female size, and males mating larger females therefore sire more offspring per mating event. Similarly, females encounter males more frequently than they need to mate. Large males produce larger spermatophores than small males and the offspring production of female increases with the size of the spermatophore she receives. Additionally, large spermatophores allow females to fertilize eggs for a longer period. Thus, mating with large males reduces the female’s need for frequent matings and she may sire sons that produce more offspring because size is heritable in copepods. Finally, we show that both males and females mate preferentially with large partners. This is the first demonstration of sexual selection by mate choice in a planktonic organism.     

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.