Sex differences in scent marking in Propithecus edwardsi of Ranomafana National Park, Madagascar

In this study we compared the scent-marking rates of females with those of males. Specifically, we examined the ability of season, dominance status, and natal status to explain the frequency of scent marking in female sifakas living wild in Ranomafana National Park, Madagascar, and compared the results with those published for males [Pochron et al., American Journal of Primatology, in press]. We also sought to determine whether vulnerability to infanticide affects marking frequency in adults of either sex, and whether female reproductive status affects female marking behavior. We found that males marked at twice the rate of females, and like males, females in single-female groups marked at the highest rates. Dominant females and non-natal females marked at higher rates than did subordinate females and natal females, a pattern also seen in males. This suggests that scent marks may convey important information about status. Neither females nor males varied their marking frequency with the presence of vulnerable infants. Females did not alter marking frequency with reproductive state, and like males, they marked at higher rates in the period prior to the mating season than they did in the mating season itself. This implies that females may use scent marks more for intrasexual aggression or territoriality than for mate attraction.     

Female pied flycatchers trade between male quality and mating status in mate choice

According to mate choice theory, females should consider both male quality and mating status when choosing a mate. In birds, strong experimental evidence indicates that females prefer males with elaborate traits. No comparable evidence exists to determine whether females take male mating status into account or how they may trade between male quality and male mating status. We studied mate choice of female pied flycatchers (Ficedula hypoleuca) in outdoor aviaries where the effect of territory quality could be eliminated and where we could control which males were mated and which were unmated. We used male plumage colour as our measure of male quality. In the aviaries, focal females could easily compare males and assess their plumage colour and mating status, and resident females were prevented from attacking prospecting females because of separation in different compartments. The study provided evidence for a trade-off in mate choice. Females may compromise by choosing an already mated male if he is more brightly coloured and, presumably, of higher quality than available unmated males. The study did not support the idea that polygyny is based on male deception of females, but the results were consistent with the female aggression hypothesis.     

Mating patterns in late-maturing female Mediterranean tarantulas may reflect the costs and benefits of sexual cannibalism

During courtship and mating, males of some invertebrate predators risk being killed and consumed by females, who in turn can obtain a foraging benefit from feeding on males. In these invertebrates, the sex ratio at the end of the mating season is usually female biased, probably due to sexual cannibalism and other sources of male mortality. Thus, at the end of the mating season males can be a limited resource to females as both mates and prey. Because of the high risk incurred when approaching females, males should show mate choice. To date there are little data on the costs and benefits of sexual cannibalism in natural populations. For one month we followed the mating patterns of 60 late-maturing Mediterranean tarantula, Lycosa tarentula L., females in a desert grassland population. The later a female matured, the shorter was her cohabitation time with males and the lower her probability of cohabiting with a male at all, suggesting that late-maturing females may be limited in their access to males as mates. At the end of the mating season, nonsexually cannibalistic late-maturing females also had poorer body conditions than did both sexually cannibalistic late-maturing females and early-maturing females, suggesting that late-maturing females may be also limited in their access to males as food. Females had higher mating success if they were smaller or in better condition (better fed). This pattern may reflect either male choice, or the possibility that small, well-fed females have higher mating success because they are less aggressive towards males. Copyright 2003 Published by Elsevier Ltd on behalf of The Association for the Study of Animal Behaviour.      

Non-reproductive copulation behavior among Tibetan macaques (<Emphasis Type="Italic">Macaca thibetana</Emphasis>) at Huangshan,China

Non-reproductive copulation, which takes place outside of the mating season and does not result in conception and birth, was studied in a free-ranging group of Tibetan monkeys (Macaca thibetana) at Mt. Huangshan, China, in the birth seasons of 1992 and 1997. We employed all occurrence and focal animal samplings to record sexual and related behaviors and affiliation interactions, respectively. Compared with sexual behavior in mating season, non-reproductive copulation occurred at a lower frequency, with less frequent ejaculation, less harassment, shorter mount duration, and an absence of pause with vocalization. It often took place in a situation in which non-lactating females were involved in social conflict or approached males for mating. Neither pregnant nor lactating females were observed to mate in the birth seasons. Copulation during the birth season did not increase a sexually receptive female’s delivery the next year, nor was it associated with increased proximity, grooming, or agonistic aid for the mating pair. However, copulated pairs spent more time co-feeding, presumably reflecting an increased tolerance on the part of the male. Adolescent males, who rarely copulated in the mating season, engaged in mating activity in birth seasons as well. Therefore, though birth-season copulation had no reproductive functions, it may have fulfilled social functions for females, such as post-aggression appeasing by males or gaining access to resources. This also offered good opportunities for adolescent males and females to develop their sexual skills for later competition.     

Sexual behavior of adult male chimpanzees of the Mahale Mountains national park,Tanzania

This study, based on 687 hr of focal observations, aims to describe overall patterns of the sexual behavior of the adult male chimpanzees of the Mahale Mountains, to compare the results with previous reports, and to explain the variations between studies. Genital inspection of cycling females by adult males was eight times as frequent as that of lactating females, and twice as frequent as that of pregnant females. Inspection of the genitals of cycling females increased dramatically 7–10 days before the onset of maximal swelling and gradually decreased as the day of ovulation approached. Adult males likely obtained information on the attractivity of females by inspecting their genitals. Mating was usually initiated by male courtship and followed by pelvic thrusts in a dorsoventral posture, performed on, rather than above, the ground, which continued for 7 s. on average, and was typically followed by female squeaking and darting from the male, or by the male grooming the female. Higher-ranking males mated with females in the peri-ovulatory period more frequently than did lower-ranking males. In particular, two alpha males mated with such females more often than did any other adult males. A male who interfered with a mating pair was dominant over the mating male in other agonistic contexts. The duration of intromission was correlated with neither dominance rank nor age. However, when an adult male declined in rank from alpha in 1991 to third in 1992, he showed a significantly shorter duration of intromission. This indicates that for a particular male, the alpha rank guaranteed longer duration of intromission. Allies of alpha males tended to mate with peri-ovulatory females more frequently than expected from their low dominance ranks. The number of mating partners was not correlated with male dominance rank, but was sometimes negatively correlated with male age. Females were significantly more likely to emit a copulatory squeak when mating with younger, rather than older, adult males. Male dominance rank and the rate of female copulatory squeaking were not correlated. Weaning infants regularly interfered with their mothers' mating. Occasionally, unrelated adolescent males and rarely females pushed themselves in between copulating adults. Female choice was indicated when they performed a “penis erection check” or took the initiative in courtship, or on the other hand showed strong reluctance to mate with particular males. Young adult males more often received erection checks than did prime males, while none of the three old adult males did. Courtship initiated by estrous females was not directed to two of the oldest males, the exception of which was the alpha male. The oldest males, except for the alpha, were consistently avoided by many estrous females, both young and old. In response to female reluctance, males behaved violently, however, this was not effective, because other more dominant males came to rescue the female. Neither courtship nor mating was seen between mature sons and their mothers, nor between brothers and sisters.     

Time, gender, and seasonality in vervet activity: A chronobiological approach

A chronobiological approach has been used to highlight the issue of how climatic factors and breeding seasonality may affect male and female diurnal activity budget. We investigated gender differences for vervet monkeys in the diurnal distribution of feeding, locomotion, inactivity, and social grooming within and between birth, pre-mating, and mating seasons. The main climatic traits were that days were shorter, drier, and cooler in the mating season. Non-parametric statistics showed that female activities changed with time of day to a greater extent than did male activities. When the constraints imposed by climatic factors and mate competition increased, from the birth to the mating season, male maintenance activities were more independent of the time of day whereas females continued to vary. Gender differences in timing were therefore greater in the mating season, except for social grooming; males and females were more coordinated in diurnal timing of their grooming in the mating season. It is argued that these changes may result in reduced levels of male-female contest competition, and may be related to higher levels of male-female socialization in the mating season. Finally, it is inferred that an out-of-phase synchronization between inactivity and feeding for males in the birth season, and for females in the mating season, may result from each sex investing more time feeding at the expense of resting in those periods.     

云南拉沙山黑白仰鼻猴交配行为和出生季节

灵长类的交配模式对于了解和掌握雄性的交配策略和社群的稳定机制非常重要,但是目前有关亚洲灵长类交配模式的数据较少;因此,该研究于2011年1—12月,分别采用全事件取样法和焦点动物瞬时扫描取样法收集了拉沙山黑白仰鼻猴群的交配行为和出生数据。猴群全年交配,有2个峰期,一个是繁殖交配高峰期(8一10月);另一个在出生季节,但其非繁殖交配的生物学意义尚不清楚。雌性通过俯卧/注视雄性或跳落邀配。爬跨射精比为8.8,射精交配稀少(11.4%),这说明雄性并非每次交配都射精,因而支持黑白仰鼻猴交配模式的主体为多次爬跨射精或处于从单次爬跨射精到多次爬跨射精连续谱上段的观点。雄性邀配的射精爬跨多于雌性,说明多次爬跨射精不仅是雄性的一个交配策略,而且可以决定交配模式在连续谱的位置。交配时间后延6~7个月,交配频次与婴猴出生率相关。拉沙山猴群出生模式为严格的季节性,这进一步证实了前人的观点。婴猴出生具有一定的同步性,且不同猴群婴猴出生的同步模式不同。     

Seasonal variation in genital and body size, sperm displacement ability, female mating rate, and male harassment in two calopterygid damselflies (Odonata: Calopterygidae)

Sperm competition is a pervasive force. One adaptation is the male ability to displace the rivals' sperm that females have stored from previous copulations. In the damselfly, Calopteryx haemorrhoidalis asturica , males with wider aedeagi displace more spermathecal sperm. The present study documents that the same mechanism operates in another damselfly, Hetaerina americana . However, this genital width in both species decreases along the season, but late-emerging females have more sperm displaced than early-emerging females. Because territorial males mated more and were larger in body and genital size than nonterritorial males, late-season females mated with considerably larger males with respect to female size and this produced higher sperm displacement. Assuming female benefits from storing sperm but that such benefit does not prevail if males displace sperm, it is predicted that, along the season, females will mate less and male harassment (in terms of male mating attempts and oviposition duration) will increase. These predictions were corroborated. In H. americana , it was also tested whether spermathecal sperm became less viable along the season. The results obtained did not corroborate this. This is the first evidence indicating that season affects sperm displacement ability and female mating frequency due to changes in male body and genital size.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 815–829.     

黄山短尾猴繁殖的季节性和雌性性活动方式

1987年开始,我们对中国黄山的短尾猴(Macaca thibetana)的性和繁殖行为进行了研究。本文根据1992年全年野外观察,对黄山短尾猴的繁殖季节性和雌性性活动方式进行深入研究,以揭示短尾猴繁殖方式和雌性性行为如何影响雄性性行为。研究采用全事件取样和目标动物取样法。结果表明,短尾猴黄山种群的交配和产仔是季节性的：交配季节发生在7月至12月,其特征是交配频率高并伴有射精;产仔季节为1月至4月。交配季节之外的交配虽有少量发生．但交配频率很低且几乎不射精。成年雌性的会阴部有轻微肿胀的性皮,但性皮的大小和颜色既不随猴群繁殖季节变化,也不随雌性个体生理周期变化。同时,雌性缺乏典型的性要求的行为——发情。因此,我们认为雌性短尾猴倾向于隐蔽自己的排卵。对46个连续观察日中成年雄性对雌性的性行为分析还表明：雄性能够区别潜在繁殖和不繁殖的雌性但不能对同一雌性的生理状况做出准确判断。隐蔽排卵是雌性在性接受期间接受大量交配的主要原因。本研究否定了短尾猴的交配是非季节性的假设(Wadaand Xicmg,1996)[动物学报51(3)：365—375,2005]。     

Scent marking and mate choice in the prairie vole, Microtus ochrogaster

Scent marking is common among male and female rodents and might be used in male-male competition and as a mechanism for mate attraction. I tested the hypotheses that females would choose males based on their frequency and placement of scent marks, and that a female would advertise interest in a particular male by placing her scent marks on or near those of a preferred mating partner. In a series of experiments conducted with prairie voles, Microtus ochrogaster, females did not choose mates based on the frequency or placement of scent marks by males nor did they advertise their interest in a particular male through the frequency or placement of scent marks. The number of males chosen that scent-marked more than their opponents did not differ significantly between females exposed (11 of 15) and not exposed (10 of 15) to scents of males. Females exposed and not exposed to scents of males preferred seven of the same males that had scent-marked more than their opponents. When a third group of females was exposed to four times more scent of the less preferred than preferred males, they still chose the preferred males. Thus, the frequency and placement of scent marks by males were not used to assess males for mate choice nor did female prairie voles use scent to advertise their preference for a mating partner. In that scent marking is common in male and female mammals, scent quality might be more important than quantity in male-male competition and mate attraction.Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved .     

Ejaculation by male rhesus in the absence of female partners

The number of ejaculatory plugs found beneath the cages of male rhesus over a 14-month period indicated a high incidence of ejaculation in the home cage in the absence of a female partner. Frequency of ejaculation in the home cage was not related to concurrent tests of sexual behavior with receptive females. Moreover, ejaculation in the home cage within 22 hr or less of sex tests did not affect the frequency of ejaculation in tests with stimulus females. Long- and short-term vasectomized males ejaculated as frequently in their home cages as nonvasectomized males. No ejaculatory plugs, of course, were found beneath the cages of males castrated 2 years earlier.     

Male age, mating status and nuptial gift quality in a bushcricket

Female mate choice occurs in many animals, and in some species females prefer older males. Because older males have demonstrated their survival ability, they may be of higher genetic quality, providing genetic benefits to the offspring of their mates. However, in species where females receive direct benefits of matings, younger males may be more likely to provide more fertile or more nutritious ejaculates, so females may discriminate against older males. Males of the bushcricket Ephippiger ephippiger (Orthoptera: Tettigoniidae) produce large spermatophores at mating (>30% of body weight, circa 10% protein content). Female E. ephippiger discriminate against the song of older males. We examined the effects of male age and mating history on male reproductive investment (spermatophore size, sperm number, nitrogen content). Males produced spermatophores with significantly fewer sperm and of lower nitrogen content on their fourth mating, despite free access to food and a 1-week interval between matings, indicating that there is a cost of mating to males. There was no indication that older virgin males produced lower-quality spermatophores. Rather, older males produced bigger spermatophores of higher nutritional value and containing more sperm. Male age and mating history seem likely to be strongly correlated in the field. We conclude that female E. ephippiger probably prefer the songs of younger males, because in the field, this preference correlates with male mating history and therefore resources provided at mating. Thus, female preference for younger males could reflect discrimination against low-quality nuptial gifts.     

Sexual behaviour of male and female tammar wallabies (Macropus eugenii) at post-partum oestrus

This study describes the sexual behaviour of the adult male and female tammar wallaby from the time a female gives birth to the end of post-partum oestrus. Eleven females were continuously observed from around the time of birth until up to 3 h after the last observed mating attempt. Males began to associate with females from around the time of birth, thus females appear to be attractive to males at, but not prior to parturition. Females subsequently allowed males to mate with them from 1.3±0.8h (mean±S.D.) post-partum, although the first ejaculation was delayed until 1.8±0.8h after birth owing to intense intermale aggression directed towards any male which attempted to mate with an oestrous female. Mating chases, in which a post-partum female was pursued by several males, were observed from 1.3±0.8h post-partum. All of this mating activity usually occurred within the first 6h after birth.     

Mating Tactics in Response to Costs Incurred by Mating with Multiple Males in Wild Female Japanese Macaques

We investigated the costs of mating with multiple males in terms of feeding time, traveling distances, sexual proceptivity, and male aggression, for wild female (Macaca fuscata yakui) on Yakushima Island, Japan. We analyzed all-day focal sampling data from 7 females during the mating season (Sept.-Nov. 1996). On days when estrous females copulated with multiple males, they decreased their feeding time to half that of anestrous days, traveled longer distances, showed more proceptive sexual behaviors and received more aggression from subordinate males than on days when they copulated with only the 1st-ranking male. On days when females copulated with only the 1st-ranking male, they showed no difference in feeding time with that of anestrous days, and expended less effort than the above mating pattern because of short traveling distances, diminished sexual proceptivity and a lower frequency of aggression received. The results suggest that the costs of estrous vary according to female sexual proceptivity and the number and social status of mating partners. Female Japanese macaques exhibit a mixed mating strategy over prolonged estrous periods, which may provide females with opportunities to maximize the benefits of copulating with multiple males and to minimize the costs of estrus by mating with only the 1st-ranking male. During an estrous cycle, females may be adjusting efforts for reproduction and survival; i.e., mating vs. feeding.     

Seasonal rank changes for adolescent and subadult natal males in a free-ranging group of rhesus monkeys

Changes in dominance rank for adolescent and subadult natal males in a semi-free-ranging rhesus macaque group were seasonal. Three 4-year-old natal males of the highest-ranking matriline occupied high ranks in the adult male dominance hierarchy during the premating period. In the mating season they dropped in rank, and this decrement was related to a concomitant drop in their alliances. After the mating season, these males rose in rank along with their two 3-year-old kin to occupy ranks 2–5 and 7–8 in the adult male dominance hierarchy. Three-year-old natal males of matrilines lower ranking than first did not become integrated into the hierarchy at this time.     

Male crickets adjust the viability of their sperm in response to female mating status

Sperm competition theory predicts that males should allocate sperm according to the number of competing ejaculates. Prudent allocation of sperm in response to different levels of sperm competition has been found across a number of taxa; however, some studies suggest that males may not always allocate sperm as expected. Here we examine sperm allocation in the Australian field cricket Teleogryllus oceanicus, using female mating status (virgin, singly mated, or multiply mated) to manipulate male perception of sperm competition risk and intensity. Consistent with theory, we found that male crickets adjust their ejaculates in response to female mating status. However, rather than altering the absolute numbers of sperm transferred to a female, males altered the quality of their sperm. Males ejaculated sperm of low viability (proportion of live vs. dead sperm) when mating with virgins, increased sperm viability when mating with singly mated females, but reduced sperm viability when mating with multiply mated females. Our results show that variation in ejaculate quality can be an important aspect of strategic ejaculation by males and suggest caution in the interpretation of studies in which males do not appear to allocate sperm according to theory.     

The Effect of Female Quality on Male Ejaculatory Expenditure and Reproductive Success in a Praying Mantid

Strategic ejaculation is a behavioural strategy shown by many animals as a response to sperm competition and/or as a potential mechanism of cryptic male choice. Males invest more mating resources when the risk of sperm competition increases or they invest more in high quality females to maximize their reproductive output. We tested this hypothesis in the false garden mantid Pseudomantis albofimbriata, where females are capable of multiply mating and body condition is an indicator of potential reproductive fitness. We predicted male mantids would ejaculate strategically by allocating more sperm to high quality females. To determine if and how males alter their ejaculate in response to mate quality, we manipulated female food quantity so that females were either in good condition with many eggs (i.e. high quality) or poor condition with few eggs (i.e. low quality). Half of the females from each treatment were used in mating trials in which transferred sperm was counted before fertilisation occurred and the other half of females were used in mating trials where fertilisation occurred and ootheca mass and total eggs in the ootheca were recorded. Opposed to our predictions, the total number of sperm and the proportion of viable sperm transferred did not vary significantly between female treatments. Male reproductive success was entirely dependent on female quality/fecundity, rather than on the number of sperm transferred. These results suggest that female quality is not a major factor influencing postcopulatory male mating strategies in P. albofimbriata, and that sperm number has little effect on male reproductive success in a single mating scenario.     

Effect of multiple mating on the reproductive performance of the rice leaffolder moth,Cnaphalocrocis medinalis (Lepidoptera: Crambidae)

To clarify whether multiple mating of females and males affects the reproductive performance of the rice leaffolder moth, Cnaphalocrocis medinalis (Guenée), we examined the effect of the number of matings (once, twice, or three times) for females (female treatment) and males (male treatment) on the incidence of moth mating, number of eggs laid, egg hatchability, and adult longevity. We also compared the effect of multiple mating imposed on males or females separately with the effect of that imposed on both sexes simultaneously (both sexes treatment). The incidence of mating of females and males that mated three times (3-mated females and males) was significantly lower than for females and males that mated twice or once (2-mated or 1-mated females and males). The incidence of mating of 1-mated moths (both sexes) was significantly higher than for 2-mated or 3-mated moths (both sexes). Two-mated or 3-mated females laid significantly more eggs with significantly higher hatchability than 1-mated females. Females that mated with 1-mated males (second male mating) or 2-mated males (third male mating) laid significantly fewer eggs than those that mated with virgin males (first male mating). Females laid significantly more eggs after the second and third matings for moths of both sexes than after the first mating for moths of both sexes. The mechanisms of improvement and decline of female reproductive performance when multiple mating was imposed on males or females are also discussed in relation to the reproductive biology of C. medinalis.     

Differences in potential reproductive rates of male and female seahorses related to courtship roles

Dwarf seahorses, Hippocampus zosterae (Syngnathidae), are distinguished by extreme morphological specialization for paternal care, the formation of monogamous pair bonds and mating repeatedly over the course of a breeding season. To determine the potential reproductive rates of male and female dwarf seahorses, we measured (1) the maximum number of offspring produced per breeding cycle when sexually receptive mates were unlimited, and (2) the relative time each sex was unavailable for mating ('time out'). We paired sexually isolated males and females with sexually receptive partners and observed them from the day of introduction through to copulation, to determine the length of time it takes each sex to prepare to mate. We conducted additional experiments to determine the length of gestation, which when added to the time needed to prepare to mate and copulate gives an estimate of total reproductive cycle duration, T. We estimated potential reproductive rate by dividing the mean number of offspring produced per breeding cycle by the duration of the breeding cycle (T). We estimated reproductive 'time out' by identifying the period of time males and females were physiologically capable of mating ('time in', S) and subtracting time S from time T. When provided with sexually receptive partners, females took 2 days longer than males to complete courtship and copulation, but neither males nor females remated during gestation. Therefore, males could potentially produce 17% more offspring than females over the course of one breeding season. Females had reproductive 'times out' 1.2 times longer than did males, as they were only capable of mating during the 4 h directly preceding copulation. Thus, H. zosterae males have higher potential reproductive rates and shorter reproductive 'times out' compared with H. zosterae females. These results and previous work indicating that seahorses display traditional courtship roles support the prediction that the sex having the higher potential reproductive rate, or equivalently, the shorter 'time out', will compete more intensely for access to the opposite sex. Copyright 2000 The Association for the Study of Animal Behaviour.     

Puberty in male Japanese macaques (Macaca fuscata): Social and sexual behavior in a confined troop

The social and sexual behaviors of four groups of males (4, 5, 9, and 10 years old) were compared throughout one complete mating season in a confined troop of Japanese macaques (Macaca fuscata). The aim of this research was to document social behavior changes that occurred during adolescence and to relate them to changes in sexual behavior that occurred at puberty. The 4-year-old males all mounted females and ejaculated but they had significantly fewer ejaculations and female partners than did the older males. The 4-year-old males also showed a pattern of delayed development in tail carriage and courtship, and they played more frequently than the older males. There were no significant differences in the frequency of aggression among the groups, but both 4-year-old and 5-year-old males were displaced by adult males and females more frequently than were the older males. We concluded that the development of adult patterns of social behavior in this genus is not climactic, but occurs slowly one to two years after physiological puberty has been reached.