Silent or Vocalizing Rats Copulate in a Similar Manner

Both male and female rats produce 50 kHz ultrasonic vocalizations (USVs) in the presence of a sexual partner and during copulation. Previous studies showed that USVs have no incentive value for rats. In this study, we evaluated the role of USVs in behavior during copulation. Three groups of rats were used: sham males paired with sham females, devocalized females paired with sham males, and sham females paired with devocalized males. During the copulation test, the USVs emitted by the sham rat were recorded and the sexual behavior of both the male and the female were observed. The results revealed that devocalized and sham females showed similar patterns of sexual behavior and no difference was found in the copulatory behavior of devocalized and sham males. Also the behavior of the partner of a sham rat was comparable to the partner of a devocalized rat. In addition, almost no changes in USVs emission were found in the 5 seconds before and/or after a copulatory behavior. It can be concluded that USVs play no important role in rat copulatory behavior at least in sexually naïve rats.     

Acoustic mate copying: female cowbirds attend to other females' vocalizations to modify their song preferences

We conducted a tutoring experiment to determine whether female brown-headed cowbirds (Molothrus ater) would attend to vocalizations of other females and use those cues to influence their own preferences for male courtship songs. We collected recordings of male songs that were unfamiliar to the subject females and paired half of the songs with female chatter vocalizations—vocalizations that females give in response to songs sung by males that are courting the females effectively. Thus, chatter immediately following a song provided a cue indicating that the song was sung by a male who was of high-enough quality to court a female successfully. Using a cross-over design, we tutored two groups of females with song–chatter pairings prior to the breeding season. In the breeding season, we placed the tutored females into sound-attenuating chambers and played them the same songs without the chatter. Females produced significantly more copulation solicitation displays in response to the songs that they had heard paired with chatter than to songs that had not been paired with chatter. This experiment is the first demonstration that females can modify their song preferences by attending to the vocal behaviour of other females.     

Female bonobos use copulation calls as social signals

During mating events, females of many primate species produce loud and distinct vocalizations known as 'copulation calls'. The adaptive significance of these signals is considered to be in promoting the caller's direct reproductive success. Here, we investigated copulation calling in bonobos (Pan paniscus), a species in which females produce these vocalizations during sexual interactions with partners of both sexes. Females were more likely to call when mating with males than with females. We also observed a positive relationship between the likelihood of calling and partner rank, regardless of partner sex. Sexual activity generally increased with swelling size (an indicator of reproductive state) and, during their peak swelling, females called more with male than with female partners. Female bonobos are unusual among the non-human primates in terms of their heightened socio-sexuality. Our results suggest that in this species, copulation calls have undergone an evolutionary transition from a purely reproductive to a more general social function, reflecting the intrinsic evolutionary links between vocal behaviour and social cognition.     

Inter-sexual multimodal communication during mating in wild geladas: the leading role of females

Female primates can emit vocalizations associated with mating that can function as honest signals of fertility. Here, we investigated the role of mating calls and visual signals in female geladas (Theropithecus gelada). Because females have a central role in the gelada society and seem to solicit sexual interactions, we answered whether they emit vocalizations in conjunction with gazing to increase mating success probability. Before and during copulations, females can emit pre-copulation calls and copulation calls. For the first time, we identified a new female vocalization emitted at the final stage of copulations (end-copulation call), possibly marking the occurrence of ejaculation. We found that longer pre-copulation call sequences were followed by both prolonged copulations and the presence of end-copulation calls, thus suggesting that females use pre-copulation calls to ensure successful copula completion. Moreover, we found that different combinations of female vocal types and gazing had different effects on male vocal behavior and motivation to complete the copula. The analysis of the vocal and visual signals revealed a complex inter-sexual multimodal chattering with the leading role of females in the signal exchange. Such chattering, led by females, modulates male sexual arousal, thus increasing the probability of the copula success.     

Sexual and maternal behaviour at the postpartum oestrus: The role of experience in time-sharing

The postpartum sexual and maternal behaviour of Norway rats (Rattus norvegicus) was observed in a seminatural habitat. Behavioural decision-making by the female was examined by disrupting the nest and newborn offspring during mating. In experiment 1, multiparous females responded to nest disruption with an increase in maternal behaviour that was largely restricted to the post-ejaculatory intervals. The temporal and frequency measures of copulation were largely unaffected, implicating behavioural time-sharing. In experiment 2, following nest disruption primiparous females did not switch efficiently between maternal and sexual behaviour, showed poor retrieval of pups, and returned to the nest during ejaculatory series. They did not display the characteristics of behavioural time-sharing, indicating that previous experience may play a role in the ability to time-share.     

Female mating status affects male mating tactic expression in the wolf spider Rabidosa punctulata

Males and females have conflicting interests on the frequency and outcomes of mating interactions. Males maximize their fitness by mating with as many females as possible, whereas choosy females often reduce receptivity following copulation. Alternative male mating tactics can be adaptive in their expression to a variety of mating contexts, including interactions with a relatively unreceptive mated female. Male Rabidosa punctulata wolf spiders can adopt distinctive mating tactics when interacting with a female, a complex courtship display, and/or a more coercive direct mount tactic that often involves grappling with females for copulation. In this study, we set up female mating treatments with initial trials and then paired mated and unmated females with males to observe both female remating frequencies and the male mating tactics used during the interactions. Males adopted different mating tactics depending on the mating status of the female they were paired with. Males were more likely to adopt a direct mount tactic with already-mated females and courtship with unmated females. Already-mated females were considerably less receptive to males during experimental trials, although they did remate 34% of the time, the majority of which were with males using a direct mount tactic. Whereas males adjusting to these contextual cues were able to gain more copulations, the observation of multiple mating in female R. punctulata introduces the potential for sperm competition. We discuss this sexual conflict in terms of the fitness consequences of these mating outcomes for both males and females.     

Male mating behavior in relation to spermatophore transfer in the white cabbage butterfly

The lifetime mating frequency of female butterflies is believed tobe dependent on the reproductive status of the males which they have mated. This report assesses those status usingPieris rapae L. Multiple mating females mated males with a short time interval after the last mating or males with many mating records. Such males, like small ones, produced small spermatophores during copulation, which may have resulted in high mating frequency of those females. The males with short time interval after the last mating or those with many mating records also showed a long mating duration. Alternative interpretations of the adaptive significance of this behavior for males are discussed.     

The function of Barbary macaque copulation calls.

In a wide variety of animal species, females produce vocalizations specific to mating contexts. It has been proposed that these copulation calls function to incite males to compete for access to the calling female. Two separate advantages of inciting male-male competition in this way have been put forward. The first suggests that as a result of calling, females are only mated by the highest ranking male in the vicinity (indirect mate choice hypothesis). The second proposes that copulation calling results in a female being mated by many males, thus promoting competition at the level of sperm (sperm competition hypothesis). In this paper, I give results from the first experimental study to test these hypotheses. Playback was used to examine the function of copulation calls of female Barbary macaques (Macaca sylvanus) in Gibraltar. Although rank did not affect lone males'' likelihood of approaching copulation calls, when playbacks were given to pairs of males only the higher ranking individual approached. Moreover, females were mated significantly sooner after playback of their copulation call than after playback of a control stimulus. These results suggest that the copulation calls of female Barbary macaques play a key role in affecting patterns of male reproductive behaviour, not only providing an indirect mechanism of female choice, but also promoting sperm competition by reducing the interval between copulations. Potential fitness benefits of inciting male-male competition at these two levels are discussed.     

Female presence and estrous state influence mouse ultrasonic courtship vocalizations

The laboratory mouse is an emerging model for context-dependent vocal signaling and reception. Mouse ultrasonic vocalizations are robustly produced in social contexts. In adults, male vocalization during courtship has become a model of interest for signal-receiver interactions. These vocalizations can be grouped into syllable types that are consistently produced by different subspecies and strains of mice. Vocalizations are unique to individuals, vary across development, and depend on social housing conditions. The behavioral significance of different syllable types, including the contexts in which different vocalizations are made and the responses listeners have to different types of vocalizations, is not well understood. We examined the effect of female presence and estrous state on male vocalizations by exploring the use of syllable types and the parameters of syllables during courtship. We also explored correlations between vocalizations and other behaviors. These experimental manipulations produced four main findings: 1) vocalizations varied among males, 2) the production of USVs and an increase in the use of a specific syllable type were temporally related to mounting behavior, 3) the frequency (kHz), bandwidth, and duration of syllables produced by males were influenced by the estrous phase of female partners, and 4) syllable types changed when females were removed. These findings show that mouse ultrasonic courtship vocalizations are sensitive to changes in female phase and presence, further demonstrating the context-sensitivity of these calls.     

Comparing the relative amount of testosterone required to restore sexual arousal, motivation, and performance in male rats

It is well established that male rat reproductive behaviors including sexual arousal, motivation, and performance are dependent on circulating levels of testosterone (T). The present study was designed to (1) compare the relative amount of T required to restore these different aspects of behavior in castrated rats, and (2) create an animal model for clinical populations with sexual impairments. Twenty-nine male Long–Evans rats were tested before and after castration for sexual performance (copulation), motivation (partner preference), and arousal (50 kHz ultrasonic vocalizations; measured together with scent marking). Sexual arousal was also inferred from copulation data. Rats were then assigned to one of four groups, and T was re-introduced via Silastic capsule implants varying in length and content: No T (empty capsules), Low T (2 mm capsules), Medium T (5 mm capsules), or High T (two 10 mm capsules). The highest dose was intended to restore physiological levels. Results indicate that High T is required for 50 kHz vocalizations, while Medium T was sufficient for the restoration of copulation, partner preference, and scent marking. These data suggest that sexual arousal may be most sensitive to reductions in testosterone. The role of T levels in measures of generalized and specific (sexual) arousal is discussed in the context of other reproductive behaviors. Furthermore, because the Low T group showed impairments across all behaviors during post-implant tests, we propose that these animals may provide a good animal model for studying clinical conditions marked by reduced motivation and arousal, including Hypoactive Sexual Desire Disorder.     

Influence of Female Copulation Calls on Male Sexual Behavior in Captive Macaca fascicularis

We investigated the function of copulation calls—vocalizations by females during mating—in captive groups of long-tailed macaques. We tested predictions of the contest-competition, sperm competition, synchronized orgasms, mate again, alpha-male notification and graded-signal hypotheses. We observed 371 copulations of 36 females wherein the presence or absence of a copulation call was clear. Females call equally often with different males and shortly after ejaculation. Copulation calls occurred equally with copulations with and without ejaculation. Calls did not incite disruptions of the mating. Following calls females mated again, more often than expected, with their mating partner. Both pregnant and fertile females uttered copulation calls. Two females conceived and mated mainly with the alpha male then. We conclude that copulation calls do not incite male contest competition for sexual access to females and that it is unlikely that calls synchronize male and female orgasms. Several hypotheses remain plausible, but not all predictions are borne out unequivocably. This alerts us to the possibility that the calls could have multiple beneficial effects; natural selection might strike a compromise among functions. Investigation of the mate again, sperm competition and alpha-male notification hypotheses, and of hypotheses not tested in our study concerning female breeding overlap and female-female agonism, is required.     

Acoustic characterization of ultrasonic vocalizations by a nocturnal primate Tarsius syrichta

This preliminary study characterizes the ultrasonic vocalizations produced by Philippine tarsiers, Tarsius syrichta. Data were collected at the Philippine Tarsier Foundation Sanctuary in Corella, Bohol, Philippines, from July through October 2010. Recordings were made on a Wildlife Acoustics Ultrasonic Song Meter 2 BAT from 29 wild, free-living adult resident T. syrichta (23 females and six males). A total of 10,309 USVs were recorded. These vocalizations fell into three main categories: chirps, twitters, and whistles. Chirps were the most frequent, followed by twitters and whistles. Whereas chirps and twitters were emitted by both male and female Philippine tarsiers, whistles were only emitted by adult males. Given that vocalizations reported in this study were exclusively recorded during capture and handling, it is very likely that these vocalizations function as distress calls. However, as the long whistle was only given by adult males who were captured at the same time as the female or the group’s infant, the function of the long whistle might be slightly different than the function of the other relatively lower-frequency USVs.     

Do Female Red-winged Blackbirds Engage in a Mixed Mating Strategy?

Parentage analyses of broods of nestling red-winged blackbirds (Agelaius phoeniceus) revealed that extra-pair fertilizations (EPFs) accounted for 24% of the offspring. 8% of attempted copulations and 13% of male courtship displays during observations of focal females were by extra-pair males. In addition, mates and non-mates often chased and occasionally made physical contact with females; 34% of those chases in which contact was made were extra-pair chases. Females behaved variably during both within-pair and extra-pair events; females crouched less and resisted more frequently during extra-pair courtship than during within-pair courtship. All extra-pair events, whether natural or induced by male removal, were either resisted or accepted by the female. In 318 focal female-hours of observation during the fertilizable period, no female was ever seen in another male's territory soliciting a copulation. In addition, removal of females' mates resulted in frequent extra-pair courtship and copulation; all of these occurred on the removed male's territory. Some females left their mates' territories on occasion — these forays were nearly always off the study area, no female was ever seen copulating with an extra-pair male while on these forays, and neither the frequency nor the duration of female forays correlated with the frequency of extra-pair fertilizations within broods. There were no associations between extra-pair fertilizations and female age, settlement order, nest order, or clutch size. The number of fledglings produced from a nest was significantly positively associated with the number of sires of the brood. Fewer offspring apparently starved in broods that were multiply sired, yet males did not provide courtship feedings during either within-pair or extra-pair copulations, nor was any paternal care provided to young sired through extra-pair matings. The frequency of infertile eggs was low (< 1%); in those instances of infertile eggs the territory owner sired some young in the same nest or another nest on his territory. Fewer broods were a mixture of within-pair and extra-pair paternity than expected by chance. Clear evidence implicating a mixed strategy on the part of females could not be gathered. Because females behaved variably and because not all costs and benefits to females of extra-pair copulations could be measured, it remains possible that female behavior patterns are either (1) part of a mixed strategy, or (2) part of a strategy minimizing the costs of copulation.     

Mating Behavior and Performance in the Two Cosmopolitan Mealybug Species Planococcus citri and Pseudococcus calceolariae (Hemiptera: Pseudococcidae)

Mating strategies of mealybugs were investigated using two heterogeneric cosmopolitan species as case study: Planococcus citri (Risso) and Pseudococcus calceolariae (Maskell). Male mating behavior of the studied species differed in respect to mate selection, and frequency and duration of copulation. Females played an active role in mate selection, by either facilitating or resisting copulation. This is a first evidence of female sexual active behavior in scale insects. In both species, male predisposition to mate was affected by previous exposition to light. The frequency of courtship and copulation decreased with the age of females for Pl. citri but not for Ps. calceolariae. Mating frequency increased with female colony density in both species.     

Information content of female copulation calls in yellow baboons.

In a wide variety of animal species, females produce vocalizations just before, during, or immediately after copulation. Observational and experimental evidence indicates that these copulation calls are sexually selected traits, functioning to promote competition between males for access to the calling female. In this paper, we present an acoustic analysis of variation in the form of copulation calls of female yellow baboons, Papio cynocephalus cynocephalus. In particular, we examine whether information about three factors-the calling female's reproductive state, the occurrence or absence of ejaculation, and the dominance rank of the mating male-is encoded in call structure and hence is potentially available to male receivers attending to the signal. Although several features of copulation calls were correlated with each of these factors, when all three were included in multiple regressions only reproductive state and rank of the mating male had independent effects on call form. These findings indicate that female copulation calls in this species signal information about the proximity to ovulation of the calling female and also the relative competitive strength of her mating partner.     

Dual Signaling during Mating in Brown-headed Cowbirds (Molothrus ater,Family Emberizidae/Icterinae)

Male brown-headed cowbirds (Molothrus ater) vocalize to females during pair formation, a period usually lasting several days. Males also vocalize to females in the seconds immediately prior to females' adopting copulatory postures. The two major classes of male vocalizations occurring during courtship and copulation are songs and flight whistles. Observations across the species' North American range suggest that the function of these two courtship vocalizations may differ geographically. Aviary observations of eastern and midwestern populations suggested, furthermore, that the precise timing of song and whistle used during copulation sequences differs, with flight whistles occurring most often after a copulatory posture but before the male mounts and the pair copulates. Such timing of the two signals suggested different proximate functions. Here, we report three experiments that addressed the communicative properties of the two signals in two midwestern populations. First, we tested females of the two populations in two playback experiments to determine copulatory responsiveness and discrimination of the two signals. We asked whether females of the two populations gave more copulatory responses to the playbacks of songs and flight whistles of males of their own population than to those of males of the other population, and whether females responded differently to songs than to whistles. In the third experiment, we observed courtship interactions among males and females from one population in a large aviary to assess the use of flight whistles in relation to courtship success. Females of both populations responded more frequently to playbacks of songs than to playbacks of flight whistles and showed reliably more responsiveness to local song variants. Thus, information in male song can be used by females to discriminate the local population. The aviary data revealed that the rate of flight whistling correlated strongly with male courtship success. Thus, the vocal antecedents to mating in midwestern cowbirds include close-range signaling to females followed by longer range signaling, perhaps to other males and to females other than the mate. Acoustic and behavioral differences between these two signals in diverse parts of the cowbirds' range suggest that the function of ‘speciestypical’ signals such as songs or whistles may not be fixed, a conclusion in keeping with the growing evidence of vocal and social mallcability in these brood parasitic birds.     

Reproductive Biology and Behavior of Rhabditis pellio, (Schneider) (Rhabditida: Rhabditidae)

Laboratory studies were conducted on the mating behavior of Rhabditis pellio males and females, which were maintained on a culture of Flavobacterium sp. bacteria isolated from earthworms. The mean time that elapsed between first contact of the sexes and their ultimate separation was 23.2 min. However, only 5.0 min were required for copulation (the interval during which male spicules were inserted into the female vagina). Three-day-old females that were permitted to mate once on their first day of adult life produced only one-third as many larvae as did females that were permitted unlimited mating. However, the longevity of females was found to decrease with an increase in the number of matings. Both males and females that were permitted to mate daily produced the greatest number of offspring when they were 4 days old. When the initiation of mating was delayed beyond their third day of life, the number of larvae produced by females decreased. In approximately one-half of the copulations, males failed to inseminate their female partners.     

The role of mating history and male size in determining mating behaviours and sexual conflict in a water strider

We conducted three experiments to test the effects of mating history of both sexes and of male body size on mating behaviours in the water strider, Gerris buenoi. Our manipulations influenced the interests of both sexes and, thus, the degree of conflict over mating behaviours. Mating history was a dichotomous variable (deprived/mated), depending on holding conditions in the laboratory. Experiment 1 considered and found independent effects of male and female mating history on latency to copulation and copulation duration. In experiment 2, we manipulated only female mating history, using unsuccessful struggle rates as evidence for female reluctance and conflict over mating. Finally, we investigated the relation between male body size and mating history on copulation duration. We predicted that intersexual conflict over mating would be lowest when females were deprived, because female interests under these conditions should more closely match those of males. Deprived females began mating in half the time of mated females and were twice as likely to mate because of reduced reluctance. Furthermore, copulation duration for deprived males was about one and a half times longer than that for mated males. Although previous studies examining nonrandom mating patterns by size predicted longer copulations for small males, we found that small males prolonged copulation when deprived more than large males. We conclude that females primarily influence copulation frequency, but males primarily influence copulation duration. Our results favour the hypothesis that reduced mating opportunity for small males accounts for their extended copulation duration. Finally, our findings provide evidence for strong effects of male body size on selection mechanisms in water striders, and support the hypothesis of conflicting pre- and postcopulatory selection mechanisms in this group. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.     

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ABSTRACT

Research on African elephant (Loxodonta africana) vocal communication has increased in recent years, yet there has been very little data collected on the vocal production of infant African elephants. Vocalizations were recorded from a group of five adult female African elephants and 3 dependent offspring (1 male and 2 female) at Disney's Animal Kingdom, Florida, U.S.A., using custom-designed audio-recording collars worn by the adult females. We measured both source and filter features of infant ‘rumble’ vocalizations made during affiliative social interactions and after cessation of nursing from adult females. Rumble vocalizations produced in the ‘nurse cessation’ context exhibited an upward shift in formant frequency locations, compared to rumbles produced during the ‘affiliation’ context. Additionally, call duration increased and fundamental frequencies decreased after nurse cessations for the male, but both females showed the opposite acoustic response. When infant rumbles accompanied nurse cessations, nursing was more likely to resume within 30 seconds compared to nurse cessations without vocalizations. These results suggest that infant rumbles associated with cessation of nursing reflect the motivational state of infants and may influence maternal responsiveness.     

Avoidance of maladaptive,precocious copulation in the cabbage white butterfly,Pieris rapae crucivora

In the cabbage white butterfly, Pieris rapae crucivora, copulation hinders normal expansion and hardening of the wings of newly emerged females. The resulting permanent wing deformation makes it impossible for females to fly and therefore, to find an oviposition site and nectar sources. An attempt was made to clarify whether the newly emerged female butterfly avoids copulation. Observation of wing expansion and hardening reveals that the wings are fully expanded and hard by 20–30 min after emergence. In the field, presentation of females with soft wings to males shows that males will attempt to copulate with these females. However, newly emerged females prevent successful completion of copulation by assuming the mate refusal posture, and thereby avoid a potentially maladaptive copulation. The discussion focusses on the question as to why females and not males avoid early copulation.