Longer copulation duration increases the risk of injury during copulation in the male bell cricket Meloimorpha japonica

Recent studies have documented male traits that cause physical harm to their mates during copulation. However, whether or not males sustain injuries during copulation has not been investigated except in sexually cannibalistic species. We found that males were injured by females during copulation in the polygynous cricket Meloimorpha japonica . More than half of the males sustained injuries to their forewings and/or genitalia during copulation. The probability of injury increased with the duration of copulation. The remating rate of females was high when the copulation duration in the first mating was short. These results suggest that copulation duration will be affected by the compromise between the risk of injury for males and paternity assurance.     

Brain control of mating behavior in the male cricket Gryllus bimaculatus DeGeer: brain neurons responsible for inhibition of copulation actions

We localized brain neurons responsible for the inhibition of copulation actions in the male cricket, Gryllus bimaculatus. Males with one connective-cut between the brain and the terminal abdominal ganglion (TAG) were able to perform normal reproductive behaviors such as courtship, copulation and spermatophore protrusion. However, copulation response tests using a female dummy showed that they failed to exhibit a transient inhibitory state after application of noxious stimulation in the mating stage and to exhibit a tonic inhibitory state in the post-copulatory sexual refractory stage. A partial cut of a single connective (hemilateral connective-cut) combined with the copulation response test and an axonal backfilling demonstrated that approximately 40 brain neurons (per one connective), whose somata were located in the posterior region of the protocerebrum and mainly contralaterally to their descending axons running through the dorso-medial part of the connective, were candidates for the inhibition of copulation actions, at least in the mating stage. In addition, we report here for the first time, that copulation actions can be elicited shortly after copulation even in intact males. This unusual behavior was frequently observed in fledglings after the final molt. The results are discussed from the perspective of the operation of the brain inhibition system on the central pattern generator (CPG) for copulation.     

Reproductive biology and function of multiple mating in the mating system of a tree cricket,Truljalia hibinonis (Orthoptera: Podoscritinae)

A tree cricket,Truljalia hibinonis, is known to show a novel sperm removal during copulation. The pattern of copulations and ovipositions showed that the sperm removal functioned to increase reproductive success for sperm removing males. The sperm removal by males evolves under the system in which female accept multiple mating. The possible benefits of multiple mating for females are examined. Multiple mating did not seem to be necessary for avoiding sperm depletion, because females stored huge number of sperm in their sperm storage organ after finishing oviposition. The ingestion of metanotal secretion during copulation also had no effect on increasing fecundity and egg size. However, mating experience may have a positive effect on increasing fecundity slightly, though there were no differences between once- and twice-mated females. The other possible benefits for each male and female are discussed.     

Male copulatory patterns in the lesser bushbaby (Galago moholi) in captivity

I observed seven pairs of sexually mature, captiveGalago moholi during the females’ ovarian cycles to determine the pattern of male copulatory behavior and its relationship to female receptivity. In this species, a copulatory lock was not apparent, a variety of thrusting patterns was displayed, and the male was capable of ejaculating following a single mount and intromission and may have ejaculated more than once during the 30-min test. Male mating indices did not differ significantly on consecutive days of the mating period, though declines in certain mating latencies on the second and third days of mating suggested increased female receptivity. Mating indices did not differ notably between the first and the second copulation within a test, though both showed some differences compared to the third copulation. InG. moholi, male copulatory behavior is typical of a prosimian species with a dispersed — nongregarious — mating system, in which copulation has a single prolonged intromission that extends into the postejaculatory period. This is characteristic of species in which more than one male has access to an estrous female.     

The copulatory plug delays ejaculation by rival males and affects sperm competition outcome in house mice

Females of many species mate with multiple males (polyandry), resulting in male–male competition extending to post‐copulation (sperm competition). Males adapt to such post‐copulatory sexual selection by altering features of their ejaculate that increase its competitiveness and/or by decreasing the risk of sperm competition through female manipulation or interference with rival male behaviour. At ejaculation, males of many species deposit copulatory plugs, which are commonly interpreted as a male adaptation to post‐copulatory competition and are thought to reduce or delay female remating. Here, we used a vertebrate model species, the house mouse, to study the consequences of copulatory plugs for post‐copulatory competition. We experimentally manipulated plugs after a female's first mating and investigated the consequences for rival male behaviour and paternity outcome. We found that even intact copulatory plugs were ineffective at preventing female remating, but that plugs influenced the rival male copulatory behaviour. Rivals facing intact copulatory plugs performed more but shorter copulations and ejaculated later than when the plug had been fully or partially removed. This suggests that the copulatory plug represents a considerable physical barrier to rival males. The paternity share of first males increased with a longer delay between the first and second males' ejaculations, indicative of fitness consequences of copulatory plugs. However, when males provided little copulatory stimulation, the incidence of pregnancy failure increased, representing a potential benefit of intense and repeated copulation besides plug removal. We discuss the potential mechanisms of how plugs influence sperm competition outcome and consequences for male copulatory behaviour.     

A potential mechanism for cryptic female choice in a flour beetle

In polyandrous mating systems, events occurring during copulation can alter the fate of the mating male's sperm. These events may exert selective pressures resulting in the evolution of diverse reproductive behaviours, morphologies and physiologies. This study investigates the role of male and female copulatory behaviours on sperm fate in the red flour beetle, Tribolium castaneum. I describe and quantify copulatory behaviours for mating pairs, and examine sperm fate by quantifying sperm transfer, sperm storage and second male sperm precedence. Whereas female quiescence during copulation and male leg rubbing during copulation together account for significant variation (26%) in sperm precedence, female copulatory quiescence alone provides information about the timing and numbers of sperm transferred. These experiments show that a female copulatory behaviour predicts sperm fate, and emphasize the value of studying variation in both male and female copulatory behaviours for understanding factors affecting sperm use.     

Copulation in the cricket is performed by chain reaction

The male and female genitalia are finely designed to match each other for copulation in the cricket Gryllus bimaculatus. Copulatory acts of the male, stereotyped and time-fixed, are elicited by stimulation of mechanoreceptors on particular regions of the abdomen, cerci and genitalia. Sequential execution of each motor act proceeds as a chain reaction in which one act stimulates some receptors which in turn elicits another act and so on, while the female remains immobile on the male's back. Each key stimulus for a motor act appears as a result of the male's own act, except for copulatory papilla protrusion by the female. The final sequence of spermatophore extrusion and transfer are irreversible fixed motor actions which are triggered when the female copulatory papilla stimulates the epiphallic hairs. They proceed without continual central drive from the brain, and apparently without sensory feedback. In addition, they are well coordinated with movement and posture in the entire body. Some neural mechanisms of controlling mating behavior and switching the reproductive cycle are discussed.     

Observations of the mating behavior and dentition of the round stingray,Urolophus halleri

Synopsis The mating behavior and dentition ofUrolophus halleri, the round stingray was examined. Males frequently bite females during the mating period but most male biting does not result in copulation. In bites that do not lead to copulation, males bite the posterior (or occasionally the medial) portion of the females' disc but females often free themselves from these bites. In bites that precede copulation, males bite the anterior portion of the females' disc and females do not struggle to free themselves. Thus, females may exert some form of choice when they are bitten. Mature males have sexually dimorphic dentition that may aid in holding females. A principal component analysis (PCA) showed that in juvenile males, the relative size of the teeth decrease while the relative thickness of the disc increases as body size enlarges; adult males displayed no clear pattern. In adult females, there is a relative decrease in tooth size and increase in relative disc thickness as body size enlarges. The relative increase in females disc thickness in areas where they are bitten may function to minimize the amount of damage due to non-copulatory biting. There is no indication that biting functions to induce female receptivity but it may allow females and males to acquire information about potential mates. Thus, copulatory biting functions to maintain contact during copulation while the function of non-copulatory biting is less clear.     

Male Head Secretion Triggers Copulation in the Dwarf Spider Diplocephalus permixtus

The term nuptial feeding is applied for any possible form of transfer of substances during the course of mating. However, the underlying functions can range from increased coupling efficiency or increased male survival to various levels of sexual selection. Males of many dwarf spider species (Erigoninae: Linyphiidae; Araneae) produce secretions from conspicuous grooves or elevations on their prosoma that seem to play a role during the mating episode. We asked if these structures represent a survival strategy, a mating effort in that they function as a mate attractant or as a mechanism to secure matings or if they simply facilitate successful genital coupling. In Diplocephalus permixtus males possess a deep groove in the ocular area into which females sink their chelicerae before mating commences. Secretory glands in the area of the groove produce a substance, which is taken up by the female in the process of mating. We experimentally prevented females from taking up secretion by covering the head structures of one group of males and compared the behaviour with controls. We investigated effects in a single male setting as well as in a competitive two‐males setting. Under both experimental conditions the probability that copulation occurred was significantly reduced in head covered males: females bit into male head structures irrespective of male treatment but covered males did not commence pedipalp insertion despite optimal mating position. Our data suggest that in contrast to other gustatorial arthropod species investigated so far the head–mouth contact between the sexes primarily triggers the mating behaviour of the male and not that of the female. Clearly, the secretion does not function as a mate attractant as mating position was achieved with similar probability in covered and control males. A function as a survival strategy can be excluded as well, because sexual cannibalism never occurred. Potential stimulating or manipulative effects of the secretion during copulatory and post‐copulatory phases remain to be investigated.     

Males Respond to the Risk of Sperm Competition in the Sexually Cannibalistic Praying Mantis, Mantis religiosa

Although studies on sperm competition examined a wide range of taxa, little is known about the selection pressures on male traits in systems with simultaneous risk of sperm competition and sexual cannibalism. Here, we experimentally studied how the risk of sperm competition affects male copulatory behavior in the sexually cannibalistic praying mantis Mantis religiosa. We recorded the onset and duration of copulations following the introduction of virgin, adult praying mantises into mating arenas with three different sex ratio treatments: polyandrous, monogamous, and polygynous. We did not detect any female phenotypic trait predicting cannibalism. The chance of male survival was related to his condition, with males in better condition being cannibalized significantly less often. In contrast, we did not identify any male trait that would favor some males to obtain copulations. Our results on copulation duration support sperm competition theory in that the copulations in the male‐biased treatment, where the perceived risk of sperm competition was greatest, were significantly longer than those in single‐male treatments. Importantly, males in better condition copulated significantly longer regardless of sex ratio. Overall, our study suggests that males can adjust their copulation behavior in response to the risk of sperm competition even in a system with frequent sexual cannibalism.     

Sexual Dimorphisms in the Dermal Denticles of the Lesser-Spotted Catshark,Scyliorhinus canicula (Linnaeus, 1758)

The dermal layers of several elasmobranch species have been shown to be sexually dimorphic. Generally, when this occurs the females have thicker dermal layers compared to those of males. This sexual dimorphism has been suggested to occur as a response to male biting during mating. Although male biting as a copulatory behaviour in Scyliorhinus canicula has been widely speculated to occur, only relatively recently has this behaviour been observed. Male S. canicula use their mouths to bite the female’s pectoral and caudal fins as part of their pre-copulatory behaviour and to grasp females during copulation. Previous work has shown that female S. canicula have a thicker epidermis compared to that of males. The structure of the dermal denticles in females may also differ from that of males in order to protect against male biting or to provide a greater degree of friction in order to allow the male more purchase. This study reveals that the length, width and density of the dermal denticles of mature male and female S. canicula are sexually dimorphic across the integument in areas where males have been observed to bite and wrap themselves around females (pectoral fin, area posterior to the pectoral fin, caudal fin, and pelvic girdle). No significant differences in the dermal denticle dimensions were found in other body areas examined (head, dorsal skin and caudal peduncle). Sexually dimorphic dermal denticles in mature S. canicula could be a response to male biting/wrapping as part of the copulatory process.     

Male Performance and Body Size Affect Female Re-Mating Occurrence in the Orb-Web Spider Leucauge mariana (Araneae, Tetragnathidae)

Females can affect male probabilities of paternity success through behavioural, morphological and/or physiological processes occurring during or after copulation. These processes under female-control include the acceptance or rejection of mating attempts by subsequent males. Leucauge mariana is an orb weaving spider that shows male mate guarding of penultimate females, male–male competition on female webs and copulatory plugs, suggesting a polyandric mating system. The aim of the present study was to ascertain whether male behaviour during courtship and copulation in L. mariana relate with female re-mating decisions. Forty-three virgin females were exposed to up to three males until they mated. In 24 cases, the copulatory plug was absent after mating and females were exposed the next day to up to three other males. Eighteen females accepted a second mating. Relatively larger females were more receptive to second matings and were more likely to copulate if the second male was smaller. Longer duration of female tapping and abdominal bobbing during courtship, and first copulations with less short insertions and more flubs, were associated with increased female acceptance to second matings. The results indicate cryptic female choice on male courtship and copulatory performance and suggest female-control over the determination of male mating success in this spider species.     

A comparison of the mating behavior of adolescent and adult female rhesus macaques (Macaca mulatta)

This study compares adult and adolescent female rhesus macaques with regard to (1) characteristics of their copulatory partners, (2) their proceptive behaviors, and (3) adult male behaviors toward them during estrus. We conducted focal follows of 24 adolescent and 65 adult free-ranging estrous female rhesus macaques on Cayo Santiago during two mating seasons. Compared to adult females, adolescents presented sexually to males at higher rates; copulated more frequently with rankless young male, and extra-group males; and, in one of two mating seasons, were ignored more frequently by males to whom they presented sexually. Adolescents tended to copulate with ranked, resident males at higher frequencies on days when the operational sex ratio (adult males:estrous adult females) was high. Males directed “muzzle-up” signals to adolescents at lower rates than to adults in one of two mating seasons, although this effect vanished when males who might have fathered adolescent females were excluded from analysis. Adolescents did not differ consistently from adults in strength of the correlation between proximity maintenance (dyadic Hinde's Index) and copulation rate, or in approach rate to males. Adolescent females, relative to adult females, presented sexually more to rankless young males, but did not present more to ranked, resident males. Both proximate (e.g. endocrine) and ultimate (e.g. differential fecundity; female-female mate competition) explanations may account for the reported differences between adult and adolescent female rhesus macaque sexuality.     

Copulatory mechanism in a sexually cannibalistic spider with genital mutilation (Araneae: Araneidae: Argiope bruennichi)

Genitalia are among the fastest evolving morphological traits as evidenced by their common function as diagnostic traits in species identification. Even though the main function of genitalia is the successful transfer of spermatozoa, the presence of diverse structures that are obviously not necessary for this suggests that genitalia are a target of sexual selection. The male genitalia of many spider species are extremely complex and equipped with numerous sclerites, plates and spines whose functions are largely unknown. Selection on male genitalia may be particularly strong in sexually cannibalistic spiders, where mating success of males is restricted to a single female. We investigated the copulatory mechanism of the sexually cannibalistic orb weaving spider Argiope bruennichi by shock freezing mating pairs and revealed a complicated interaction between the appendices and sclerites that make up the male gonopods (paired pedipalps). The plate that covers the female genital opening (scape) is secured between two appendices of the male genital bulb, while three sclerites that bear the sperm duct are unfolded and extended into the female copulatory opening. During copulation, females attack and cannibalise the male and males mutilate their genitalia in about 80% of cases. Our study demonstrates that (i) genital coupling is largely accomplished on the external part of the female genitalia, (ii) that the mechanism requires an interaction between several non-sperm-transferring structures and (iii) that there are two predetermined breaking points in the male genitalia. Further comparative work on the genus Argiope will test if the copulatory mechanism with genital mutilation indeed is an adaptation to sexual cannibalism or if cannibalism is a female counter adaptation to male monopolisation through genital plugging.     

Mating Behavior, Sexual Selection, and Copulatory Courtship in a Promiscuous Beetle

The function of male movements during copulation is unclear. These movements may be a result of the necessary mechanics of insemination, or they may also have further function, for instance, stimulating or courting a female during mating, perhaps influencing female mate choice. We present data from three experiments exploring the mating behavior and copulatory movements of the highly promiscuous beetle Psilothrix viridicoeruleus. Male mating success in the struggle over mating was not related to male or female size (measured by weight) but successful males were more vigorous in terms of copulatory movements. These males took longer to mount females but copulated longer and remained mounted longer. We discuss these results in terms of the mating system of Psilothrix and also in terms of observations of the timing of insemination during copulation. We suggest that copulatory movements in this species are best understood as copulatory courtship.     

Female walking during copulation reduces the likelihood of sperm transfer from males in the sweet potato weevil,Cylas formicarius

Behaviour during copulation can alter the fate of sperm of the mating males. This behaviour may exert selective pressure, resulting in the evolution of diverse reproductive behaviour, morphology, and physiology. This study examined the role of female copulatory behaviour on sperm fate in the sweet potato weevil, Cylas formicarius (Fabricius) (Coleoptera: Brentidae). In this species, males mount the female during copulation. The female frequently walks during copulation, carrying the male on her back. Here, we describe and quantify the copulatory behaviour of mating pairs and examine the sperm fate. Insemination success, as determined by the presence of sperm in the spermatheca, was lower when females walked for longer periods during copulation. This result emphasizes the value of studying variation in female copulatory behaviour in order to understand the factors that influence sperm fate. We discuss the implications of these results on sexual selection and utility in programs applying sterile insect techniques.     

Sexual Inhibition in <Emphasis Type="Italic">Spalangia endius</Emphasis> Males After Mating and Time for Ejaculate Replenishment

Although males are often stereotyped as always sexually responsive, a previous study with the parasitoid wasp Spalangia endius showed that males exhibit a post-mating decrease in sexual responsiveness. The present study examined the duration of the decrease and whether waiting to remate increased the amount of ejaculate that a male provided to his second mate. Mated males were back to being as sexually responsive as virgin males by about 5 min after mating. The amount of ejaculate that females received from already mated males was not significantly related to the duration between the male’s first and second mating. The short-term decrease in sexual responsiveness of a mated male may prevent him from immediately trying to remate with the same female because it gives her time to get away.     

The secondary copulatory organ in female ground weta (Hemiandrus pallitarsis,Orthoptera: Anostostomatidae): a sexually selected device in females?

Secondary sexual devices in female insects, primarily abdominal modifications, appear to function as a means of thwarting coercive mating attempts by males or are, in rare cases, sexually selected adaptations. Female ground weta, Hemiandrus pallitarsis (Orthoptera: Anostostomatidae), have an elaborate elbowed device on the underside of the mid-abdomen. Experimental removal of this accessory organ shows that it does not increase the probability of mating as predicted by the thwart-copulation hypothesis. Instead, removal prevents copulation, thus demonstrating that it is a secondary copulatory device. The male attaches to the organ both at the beginning of copulation and at the end, when he positions himself to adhere a spermatophylax food gift onto the mid-ventral region of his mate. The female accessory organ does not function to manipulate eggs or larvae (females provide care to their single clutch of offspring) and is unlikely to be a copulatory structure that prevents hybridization. The great extent of the modification of the ventral abdominal segments of H. pallitarsis females compared to other Hemiandrus species is consistent with a history of sexual selection on the accessory organ. Taken together, these results and the finding that the length of the accessory organ of H. pallitarsis correlates with female fecundity, suggest that this structure evolved under sexual selection to acquire nuptial gifts from males.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 85 , 463–469.     

Female behaviour and the interaction of male and female genital traits mediate sperm transfer during mating

Natural selection and post‐copulatory sexual selection, including sexual conflict, contribute to genital diversification. Fundamental first steps in understanding how these processes shape the evolution of specific genital traits are to determine their function experimentally and to understand the interactions between female and male genitalia during copulation. Our experimental manipulations of male and female genitalia in red‐sided garter snakes (Thamnophis sirtalis parietalis) reveal that copulation duration and copulatory plug deposition, as well as total and oviductal/vaginal sperm counts, are influenced by the interaction between male and female genital traits and female behaviour during copulation. By mating females with anesthetized cloacae to males with spine‐ablated hemipenes using a fully factorial design, we identified significant female–male copulatory trait interactions and found that females prevent sperm from entering their oviducts by contracting their vaginal pouch. Furthermore, these muscular contractions limit copulatory plug size, whereas the basal spine of the male hemipene aids in sperm and plug transfer. Our results are consistent with a role of sexual conflict in mating interactions and highlight the evolutionary importance of female resistance to reproductive outcomes.