Determinants of Mating Frequency in the Spiny Orbweaving Spider,Micrathena gracilis (Araneae: Araneidae)

Male Micrathena gracilis require two copulations, separated by a dismount, in order to inseminate both reproductive tracts of the female. We examined several factors that might influence a male's copulatory success. Web structure influenced male courtship and dismount tactics, but not copulatory frequency. The presence of another male reduced the likelihood of a given male copulating with both tracts, a limitation mediated by sexual responsiveness of the female. Mating status of the female did influence copulatory frequency; males were less likely to copulate a second time with nonvirgin females. In summary, males modify mating activities to reduce predation by females, to reduce intermale competition, and to avoid expending gametes when there is little chance of fertilization. Females influence males by predatory activities, mediated through web structure, and enhancing sperm competition among males.     

Rhesus macaque copulation calls: Re-evaluating the “honest signal” hypothesis

Male rhesus macaques sometimes give loud calls while thrusting or dismounting during multi-mount copulations.Hauser (1993) has proposed that these calls (1) impose a cost (increased risk of aggression) on calling males, and (2) increase callers' copulation frequencies, supporting the hypothesis that calls function as honest signals (handicaps) that females use to evaluate male quality during mate choice. This hypothesis was re-examined using data collected at Cayo Santiago, Puerto Rico on 40 focal females and their 56 observed copulatory partners. Although attacks by males against copulating pairs were frequent, they were usually directed only against the female of the pair. Males that called were no more likely than silent males to suffer male escalated attack during or immediately following mount series. Male-female dyads in which the male called during copulation were significantly more likely than non-calling dyads to complete the most copulations observed for any given female. Males that called at least once were significantly more likely than non-calling males to complete at least one copulation with a peri-ovulatory female. A log-linear model revealed that male rank and calling were both associated with likelihood of experiencing at least one peri-ovulatory copulation. However, calling was not associated with reception of demonstrated female mate choice behaviors. Controlling for dominance rank, callers did not experience more female proximity maintenance than non-callers. Nor were callers' hip-grasps refused less frequently than non-callers' hip-grasps. These results cast doubt on the hypothesis that rhesus macaque copulation calls are costly, honest indicators of intrinsic male quality. A contrasting alternative hypothesis, that a male's copulatory calls advertise relative immunity from attacks against his copulatory partners, was not supported either. Thus, the function of rhesus macaque copulatory calls remains unknown. The unusually high rate of copulations amongHauser's (1993) subjects may explain the discrepancy in results, but it is unclear how high copulation rates would increase the cost of copulatory calling to males.     

Seasonal changes in reproductive behavior in two ovariectomized female rhesus monkeys treated year-round with estradiol

Two ovariectomized female rhesus monkeys treated year-round with estradiol-filled capsules were used in hour-long behavioral tests with male rhesus monkeys both in and out of the normal breeding season. The study was designed primarily to test male hormonal responses to copulatory behavior and it was expected that behavior of the females would be essentially the same under both seasonal conditions. Several behaviors of the female, however, were found to fluctuate on a seasonal basis, namely (1) proximity to the male, (2) grooming of the male, (3) sexual presentations, and (4) threatening away (a form of sexual invitation). All of these behaviors, except proximity, were found to be positively correlated with the male partner's testosterone levels before and after the behavior test in the only test condition in which most males ejaculated. The preliminary suggestion is made that these females responded to some cue, either from the environment or from the males, that caused a change in their behavior between breeding and nonbreeding seasons despite the constant hormonal environment provided by the estradiol capsules. Since the same behaviors which were sensitive to seasonal effects were positively correlated with male testosterone levels, it is possible that the male's hormonal status affects affects female behavior.     

Spiders,microbes and sex: Bacterial exposure on copulatory organs alters mating behaviour in funnel‐web spiders

Environmental microbes have the potential to be involved in nearly all behavioural processes. For example, mating systems where males use intromittent organs to transfer sperm to females represent a means by which environmental microbes collected by males can breach entry into females' body cavities during mating. However, the degree to which the acquisition of environmental microbes onto important sex structures alters courtship behaviours remains unknown. Here, we collected bacteria from the copulatory organs of Agelenopsis pennsylvanica funnel‐weaving spiders in situ to test whether exposure to bacteria on copulatory organs can alter hosts' courtship behaviour, reproductive success and survival. We used a standardized assay to repeatedly measure each spider's aggressiveness, a behavioural component of both male courtship and female sexual receptivity. Then, we experimentally altered the bacteria present on male and female spiders' copulatory organs with an application of either (a) a mixture of bacteria collected from conspecifics to increase bacterial presence, (b) an antibiotic to reduce bacterial presence or (c) a procedural control. Each spider was paired with a size‐matched spider of the opposite sex whose copulatory organs were unaltered, and we measured the latency until the onset and the duration of courtship. Spiders were then isolated, and we measured each individual's time until death and female fecundity over the next 40 days. We found that female exposure to bacteria had multiple effects on mating dynamics. Males took over four times longer to begin courting females that had been exposed to bacteria compared to unexposed and antibiotic‐treated females. Only when courting these bacteria‐exposed females, males began courtship sooner when females were more aggressive. Lastly, females whose mate had been exposed to bacteria experienced reduced survival. These data suggest that bacteria present on animals' copulatory organs can alter courtship behaviours, female survivorship, and may potentially play a role in mating dynamics.     

Psychobiology of sexual behavior in a whiptail lizard, Cnemidophorus inornatus

This study investigated the effects of social environment on gonadal recrudescence and sexual behavior in male and female Little Striped Whiptail lizards (Cnemidophorus inornatus). The presence of sexually active males facilitates ovarian recrudescence in conspecific females. Similarly, the presence of reproductively active females facilitates testicular recrudescence in conspecific males. Males housed with females, however, had lower average circulating concentrations of testosterone and dihydrotestosterone, and higher average concentrations of corticosterone compared to intact males housed in isolation. In other studies, the presence of reproductively active females partially restored courtship behavior in castrated males compared to castrated males housed in isolation. Despite the stimulatory effects of females on castrates, exogenous androgens are required for complete restoration of all components of sexual behavior in male C. inornatus. Females are receptive to male courtship and copulatory behavior only during the vitellogenic stages; females in previtellogenic or postovulatory ovarian stages aggressively reject male courtship advances. These findings demonstrate reciprocal effects of sexual behaviors of males and females upon each other's reproductive behavior and physiology.     

Decline of sexual behavior in castrated male rats: Effects of female precopulatory behavior

From the population of 89 adult sexually inexperienced Wistar male rats 20 animals that initiated copulatory behavior with females exhibiting low intensity of precopulatory behavior (presenting females) were preselected. Prior to castration all 20 males had the same sexual experience: three ejaculatory series in four weekly sessions with females exhibiting high intensity of precopulatory behavior (darting females). Following castration, the decline of copulatory behavior was much slower for the nine males tested with darting females as compared to the 11 males tested with presenting females. Male precopulatory behaviors (anogenital sniffing, touching flanks, etc) outlasted the loss of copulatory behavior and seem to be less dependent on both external and internal determinants. It is concluded that intensive external sexual stimuli can function to compensate, and therefore mask, the subnormal operation of androgen-dependent mechanisms in initiating the copulatory behavior.     

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.     

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.     

Natural or hormone-induced sexual and social behaviors in the female brown lemming (Lemmus trimucronatus)

The behaviors of intact or ovariectomized, estradiol benzoate-treated or estradiol benzoate followed by progesterone-treated female brown lemmings were compared. Intact, diestrous females engaged in more social interactions with a male than did ovariectomized females (Experiment 1). In the first 5 min of a 1-hr mating exposure (Experiment 2, Test A) intact females in natural estrus engaged in more social and sexual behaviors than did ovariectomized females in estrogen-induced estrus. However, during the last 5 min of the 1-hr exposure (Test B) ovariectomized females receiving estrogen alone continued to show high levels of sexual activity with a male partner, while intact estrous females or females receiving estrogen followed by progesterone showed an apparent drop in sexual receptivity and an increase in aggressivity. Aggressive behaviors, as indexed by threat-leap behaviors on the part of the female may increase in the presence of progesterone. Declines in sexual activity, occurring within 1 hr of progesterone injection, were apparently dependent on the interaction of progesterone and copulatory events which may affect both the male and female.     

Female crickets assess relatedness during mate guarding and bias storage of sperm towards unrelated males

Recent evidence shows that females exert a post‐copulatory fertilization bias in favour of unrelated males to avoid the genetic incompatibilities derived from inbreeding. One of the mechanisms suggested for fertilization biases in insects is female control over transport of sperm to the sperm‐storage organs. We investigated post‐copulatory inbreeding‐avoidance mechanisms in females of the cricket Teleogryllus oceanicus. We assessed the relative contribution of related and unrelated males to the sperm stores of double‐mated females. To demonstrate unequivocally that biased sperm storage results from female control rather than cryptic male choice, we manipulated the relatedness of mated males and of males performing post‐copulatory mate guarding. Our results show that when guarded by a related male, females store less sperm from their actual mate, irrespective of the relatedness of the mating male. Our data support the notion that inhibition of sperm storage by female crickets can act as a form of cryptic female choice to avoid the severe negative effects of inbreeding.     

Rival male relatedness does not affect ejaculate allocation as predicted by sperm competition theory

When females are sexually promiscuous, the intensity of sperm competition for males depends on how many partners females mate with. To maximize fitness, males should adjust their copulatory investment in relation to this intensity. However, fitness costs associated with sperm competition may not only depend on how many males a female has mated with, but also how related rival males are. According to theoretical predictions, males should adjust their copulatory investment in response to the relatedness of their male rival, and transfer more sperm to females that have first mated with a non-sibling male than females that have mated to a related male. Here, for the first time, we empirically test this theory using the Australian field cricket Teleogryllus oceanicus. We expose male crickets to sperm competition from either a full sibling or non-sibling male, by using both the presence of a rival male and the rival male's actual competing ejaculate as cues. Contrary to predictions, we find that males do not adjust ejaculates in response to the relatedness of their male rival. Instead, males with both full-sibling and non-sibling rivals allocate sperm of similar quality to females. This lack of kin biased behaviour is independent of any potentially confounding effect of strong competition between close relatives; kin biased behaviour was absent irrespective of whether males were raised in full sibling or mixed relatedness groups.     

Heterosexual interactions in laboratory-housed stumptail macaques (Macaca arctoides): observations during the menstrual cycle and after ovariectomy.

Heterosexual interactions of pairs of stumptail macaques (Macaca arctoides) were studied in relation to the female menstrual cycle and after ovariectomy. Five intact male and 10 tubal-ligated female macaques were observed in laboratory pair tests of 20-mins duration, and data were obtained on various male and female behaviors. Each male was tested with the same two females during four 40-day observation periods. Males were tested daily and females were tested every other day. After two 40-day testing periods, one female partner of each male was ovariectomized and the other was sham-operated. Blood was collected regularly from the females during the course of observation and serum levels of estradiol and progesterone were determined by radioimmunoassay. The midcycle peak of estradiol was observed to occur approximately 18 days prior to menses. A distinct secondary peak in estradiol was observed to occur during the luteal phase of all cycles examined. Of 28 different male and female behaviors studied only female presentation to male sexual contact showed a significant midcycle peak related to the endogenous estradiol surge. After ovariectomy a significant decrease in the frequency of several male copulatory behaviors was observed, but most males continued to copulate regularly with their spayed partners throughout the period of this study. Thus, the pattern of copulatory behavior observed in stumptail macaques over the cycle of the female partner and subsequent to ovariectomy differs from that observed in other macaque species studied in the laboratory. It is concluded that cyclical fluctuations in the level of ovarian hormones are not significantly related to measures of sexual interactions in laboratory tests of this species, although the maintenance of copulation and associated behaviors at high levels depends to some degree upon the ovary.     

根田鼠的熟悉性及其自然动情下的配偶选择

报道了在室内雌鼠自然动情条件下根田鼠两性个体配偶选择的3种熟悉性效应的5组实验结果,包括雌性对熟悉和陌生雄鼠、配偶和陌生雄鼠、配偶和熟悉雄鼠的择偶行为实验 , 雄性对熟悉和陌生雌鼠、配偶和陌生雌鼠的择偶行为实验。在实验前, 将1对性成熟的雌雄鼠 (非亲缘关系) 每天关养8 h 共8 d 建立熟悉性; 配对雌雄鼠至少生育1胎则为配偶关系。在 Y 形迷宫内的30 min 实验中, 两个被选择鼠分别以项链拴在横跨选择箱顶端中央的一根铁丝上, 限制它们在各自箱内活动; 而允许异性选择鼠从中立箱自由进入两个选择箱。记录选择鼠对异性被选择鼠的访问、社会探究、攻击、交配和友好行为的频次和时间。经Wilcoxon 关联样本 T 检验发现, 除交配行为不显著外, 雌性根田鼠均选择熟悉性较高的雄鼠; 而雄鼠不具这种选择性。该结果提示雌雄根田鼠的不同择偶行为也许就是反映该种婚配制度特征的直接行为过程。因此, 在择偶行为中, 雌性根田鼠表现的单配性倾向和雄鼠的多配性倾向, 只能以其婚配制度为一雄多雌制的假设来解释。我们的实验还说明在多配制田鼠中, 一雄多雌制与混交制在雌鼠择偶行为上是不同的, 而与单配制的相似或相近, 故简单地比较多配制与单配制则不能反映田鼠亚科动物婚配制度的多样性。     

Sexual behaviour and reproductive competition in male wild and domestic Norway rats

The copulatory behaviour of male wild Norway rats (Rattus norvegicus) exposed individually to females in hormone-induced oestrus is characterized by fewer mounts and intromissions per ejaculatory series, shorter ejaculation latencies, and longer ejaculatory clasps than the copulatory behaviour of domestic males. Competition between wild and domestic males tested simultaneously in pairs produced only minor changes in copulatory behaviour. The reproductive success of wild and domestic male pairs simultaneously exposed to naturally cycling females was nearly equal in both the laboratory and an outdoor enclosure. Genotype of the female had little effect on male copulatory behaviour. The copulatory behaviour of sexually naive and experienced domestic rats did not differ. However, most sexually naive wild males failed to exhibit copulatory behaviour under the test conditions employed.     

Sexual and social behavior of adult saddle-back tamarins (Saguinus fuscicollis), castrated as neonates

The behavior of four adult males castrated as neonates was compared with that of three neonatally sham-castrated males and two untreated males serving as controls. Two males castrated as sexually experienced adults served as additional controls. All males lived in permanent male-female pairs. The interactions of neonatally castrated, sham-castrated, and intact control males with strange conspecifics were tested in a situation analogous to territorial defense. In addition, the interactions of all males with their female pair mates were tested. Neonatally castrated males showed less injurious aggression against strangers and interacted with them less frequently than control males. There were significant differences between both groups of males in some of the agonistic behaviors recorded, but not in all. In pair tests, neonatally castrated males failed to show any coordinated male copulatory behavior but control males did. Males castrated as adults showed rates of mounting and thrusting similar to those of intact controls. Neonatally castrated males tended to interact with their females on a lower level than neonatally sham-castrated and untreated control males. How ever, most differences between both groups in the levels of pair mate interactions were not statistically significant. It is concluded that castration during neonatal life prevents the display of male copulatory behavior in adulthood and selectively affects the display of a number of other behaviors in adulthood. It is not possible at this point to decide whether these results are due to hormonal deficiencies on an organizational or activational level or both. However, data from studies currently in progress suggest that organizational processes might be responsible for part of the effect.     

Copulatory courtship and cryptic female choice in red flour beetles Tribolium castaneum

Males of many animal species engage in courtship behaviours during and after copulation that appear to be solely aimed at stimulating the female. It has been suggested that these behaviours have evolved by cryptic female choice, whereby females are thought to impose biases on male postmating paternity success. Males of the red flour beetle Tribolium castaneum rub the lateral edges of the females' elytra with their tarsi during copulation. We manipulated female perception of this behaviour by tarsal ablation in males, thus preventing males from reaching the edge of the female elytra with their manipulated legs, and by subsequently performing a series of double-mating experiments where the copulatory behaviour was quantified. We found a positive relationship between the intensity of the copulatory courtship behaviour and relative fertilization success among unmanipulated males. This pattern, however, was absent in manipulated males, where female perception of male behaviour differed from that actually performed. Thus, female perception of male copulatory courtship behaviour, rather than male behaviour per se, apparently governs the fate of sperm competing over fertilizations within the female, showing that copulatory courtship is under selection by cryptic female choice.     

Paternity analyses in wild‐caught and laboratory‐reared Caribbean cricket females reveal the influence of mating environment on post‐copulatory sexual selection

Polyandry is ubiquitous in insects and provides the conditions necessary for male‐ and female‐driven forms of post‐copulatory sexual selection to arise. Populations of Amphiacusta sanctaecrucis exhibit significant divergence in portions of the male genitalia that are inserted directly into the female reproductive tract, suggesting that males may exercise some post‐copulatory control over fertilization success. We examine the potential for male–male and male–female post‐copulatory interactions to influence paternity in wild‐caught females of A. sanctaecrucis and contrast our findings with those obtained from females reared in a high‐density laboratory environment. We find that female A. sanctaecrucis exercise control by mating multiple times (females mount males), but that male–male post‐copulatory interactions may influence paternity success. Moreover, post‐copulatory interactions that affect reproductive success of males are not independent of mating environment: clutches of wild‐caught females exhibit higher sire diversity and lower paternity skew than clutches of laboratory‐reared females. There was no strong evidence for last male precedence in either case. Most attempts at disentangling the contributions of male–male and male–female interactions towards post‐copulatory sexual selection have been undertaken in a laboratory setting and may not capture the full context in which they take place – such as the relationship between premating and post‐mating interactions. Our results reinforce the importance of designing studies that can capture the multifaceted nature of sexual selection for elucidating the role of post‐copulatory sexual selection in driving the evolution of male and female reproductive traits, especially when different components (e.g. precopulatory and post‐copulatory interactions) do not exert independent effects on reproductive outcomes.     

Copulatory behaviors and body condition predict post-mating female hormone concentrations, fertilization success, and primary sex ratios in Japanese quail

Environmental cues and social interactions are known to influence reproductive physiology and behavior in vertebrates. In female birds, male courtship displays can result in the growth of ovarian follicles, the production of reproductive hormones, and stimulation of oviduct development, all of which have the potential to influence maternal investment. Male Japanese quail follow a typical sequence of copulatory behaviors during a mating interaction and often force copulations with unreceptive females. We hypothesized that female Japanese quail could adjust maternal investment in response to male copulatory behaviors during a single mating interaction. We investigated the relationships between 1) male copulatory behaviors and post-mating concentrations of steroids in the female, 2) female steroid concentrations and fertilization success of inseminations and 3) female steroid concentrations and the offspring sex ratio. We found that male condition and copulatory behaviors predicted female steroid concentrations and maternal investment in eggs laid after a mating trial. The body condition of one or both mates was a significant predictor of the changes in female corticosterone and testosterone concentrations after mating, whereas specific male copulatory behaviors significantly predicted changes in female progesterone concentrations. Male and female body condition, male neck grabs and post-mating concentrations of female corticosterone, progesterone, and testosterone were all significant predictors of egg fertilization rates. Female body condition, male copulation efficiency, and female testosterone concentrations were significant predictors of offspring sex ratios. Our results show that phenotypic and behavioral characteristics of male Japanese quail modulate female steroid concentrations and result in changes in maternal investment.     

Conflict between sexes in the water strider, Gerris lacustris: a test of two hypotheses for male guarding behavior

We studied the effect of operational sex ratio on female reluctanceand male persistence to mate as well as on the length of copulationand postcopulatory guarding in Gerris lacustris by adding fivesurplus males or females to the basin with a pair in tandem.In the control treatment, a pair alone was tested. Accordingto the copulatory guarding hypothesis (CGH), males should prolongmating and guard females in the presence of surplus males. Accordingto the convenience polyandry hypothesis (CPH), females shouldshow lower levels of resistance to prolonged mating in the presenceof surplus males because the mating male protects the femaleagainst harassment from other males. As expected on the basisof both the CGH and CPH, mating (copulation + guarding) averagedlonger in the male-biased treatment. The behavior of males andfemales during mating suggested that both hypotheses hold true:females showed less resistance to prolonged mating (as predictedfrom CPH), and male behavior suggested stronger efforts to stayon the female when surplus males were present (as predictedfrom CGH). Comparisons of the treatment with surplus femaleswith the results from the mating pair without surplus individualssuggested that the capabilities of water striders in tandemto assess the sex of nearby nonmating striders are limited.