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]。     

非优势顺位雄性黄山短尾猴的交配策略

在多雄多雌、顺位决定交配机会、雄性偏斜繁殖的非人灵长类社会中,低顺位雄性为提高自己的繁殖成功率,通常采取多样的交配策略以获得较多的交配机会。对非优势顺位雄性交配策略的研究可以增加对灵长类社会性行为复杂性的理解,对于深入探讨动物的婚配制度及繁殖策略具有十分重要的科学意义。本研究采用目 标动物取样法和全事件记录法,在２０１２ 年９ －１２ 月（交配期）记录了安徽黄山短尾猴鱼鳞坑ＹＡ１ 群中４ 只成年雄性个体的交配及有关的行为。研究发现：（１）非优势顺位雄性个体在远离优势顺位雄性视野范围的交配频次和交配时间显著高于优势顺位视野范围内;（２）与优势顺位雄性个体相比,非优势顺位雄性的强行交配（forced copulation）和隐秘交配（clandestine copulation）比例较高;（３） 就交配对象而言,对成功生育的雌性个体,优势顺位雄性在选择交配对象时具有明显的倾向性,非优势顺位雄性在选择时倾向性不显著;对未生育的雌性,优势顺位雄性更倾向于选择没有生殖经历的亚成年雌性个体,非优势顺位雄性则倾向于选择处于哺乳后期的成年雌性个体;（４）在具体交配策略上,优势顺位雄性选择跟随（follow） 雌性,非优势顺位雄性则通过做鬼脸 （grimacing）和性追求（sexual chasing）直接获取交配机会。本研究结果表明黄山短尾猴中非优势雄性个体形成了多变的交配策略,更多采取强制性方式,以获得更多的交配机会,多数的交配都是机会性的。     

藏酋猴交配活动中的性打搅

1986—1987年期间,对藏酋猴(Macaca thihetana)黄山鱼鳞坑群交配活动中的性打搅行为进行了为期9个月的观察和研究。在766次交配中,交配个体受到了266次性打搅。性打搅行为大部分发生在交配雄性开始爬跨雌性时(89.2％),较少发生在交配雄性爬跨雌性之前(0.7％)和射精时或射精后(10.1％)。在266次有性打搅的交配中,参加性打搅的个体达473只次,其中,成年个体参于性打搅的频率高于亚成年和幼年个体,而雌雄个体之间各自参于性打搅的频率无明显差异。各个体因在群中的顺位不同,其参加交配活动和受到的性打搅的频率也不同。高顺位雄性和低顺位雌性比低顺位雄性和高顺位雌性较多地参于交配活动,同时它们受到的性打搅的频率也较高。因此,受到性打搅频率的高低与参于交配的频率有关,而与个体顺位的高低无关。在性打搅的行为类型中,63.8％是性打搅者直接朝着交配的雄性,6.6％朝着交配的雌性,29.6％无明确朝向。低顺位雄性和雌性的性打搅行为很少能中断高顺位雄性的交配活动,而高顺位雄性的性打搅却能成功地中断低顺位雄性或雌性的交配活动。然而,在266次有性打搅行为发生的交配中,仅8.6％的交配被性打搅中断。为此,本文就不同性别和顺位的个体所进行的性打搅行为的动机进行了探讨。     

雄性黄山短尾猴攻击行为和粪便睾酮水平季节性变化

睾酮对启动和维持雄性动物的性行为及保持雄性第二性征有重要作用,高浓度的睾酮还可促进雄性的攻击性。但是长时期维持高的睾酮水平还会使雄性付出很高的能量代价,所以雄性不会长时期地维持高的睾酮水平。本研究对黄山短尾猴鱼鳞坑YA2群雄性个体交配期（2006年9月-12月）与产仔期（2007年1月-4月）攻击行为进行研究,同时收集研究个体的新鲜粪便,并用放射免疫分析法测定粪便睾酮水平,以探讨雄性黄山短尾猴不同时期睾酮水平差异及睾酮水平与攻击行为的关系。结果表明,成年雄性黄山短尾猴攻击行为发生频次在交配期显著高于非交配期;成年雄性粪便睾酮水平在交配期显著高于非交配期;交配期成年雄性的粪便睾酮水平与攻击行为发生频次有较显著的正相关关系,而在非交配期无显著相关;低顺位个体在交配期和非交配期都没有观察到攻击行为。本研究的结果支持了“繁殖竞争假说”。     

不同社群序位布氏田鼠的繁殖行为

在实验室条件下观察了捕自内蒙古草原的不同序位布氏田鼠的繁殖行为特征。结果表明, 雌鼠对雄性配偶存在着激烈的竞争, 高序位雌鼠获得被选择、交尾及产仔的机会最多, 较低序位雌鼠若被选择并且产仔, 则其序位将随之上升; 高序位雄鼠对各个序位雌鼠均可选择, 且在繁殖期间对同性个体表现很强的攻击性, 以获得更多的交配机会。在社群繁殖中高序位雌、雄个体的作用明显高于低序位个体, 这是布氏田鼠种群长期选择进化的生存策略。实验支持了布氏田鼠为混交制、一雄多雌的婚配制度推论。     

雄性黄山短尾猴(Macaca thibetana)攻击支持雌性以换取交配回报

生物市场理论认为动物个体之间通过某种协定交换有价值的商品,使双方均受益。该研究采用目标动物法、行为取样法和连续记录法,对浮溪黄山野生猴谷鱼鳞坑短尾猴(Macaca thibetana)A1群(YA1群)和A2群(YA2群)成年个体在非繁殖季节(2011年8月—12月)和繁殖季节(2012年2月—5月)的雄性攻击支持雌性行为和交配行为进行研究,探讨雄性攻击支持雌性与交配之间的关系。两猴群在繁殖季节和非繁殖季节雄性攻击支持雌性与交配行为均呈显著正相关;YA2群繁殖季节与非繁殖季节攻击支持后交配频次均显著高于随机交配;YA1群在繁殖季节攻击支持后交配频次与随机交配频次差异不显著,但在非繁殖季节攻击支持后交配频次显著高于随机交配,说明短尾猴成年雄性攻击支持雌性可以换取与该雌性个体的交配回报。本研究验证了生物市场理论中社会行为存在交换,首次证明了雄性攻击支持可以换取雌性的交配回报,为进一步研究雄性性竞争与雌性选择提供了实例。     

秦岭野生雌性川金丝猴粪便睾酮水平与邀配频次的季节性变化

为研究野生雌性川金丝猴(Rhinopithecus roxellana)不同季节体内睾酮水平与性行为频次的变化及二者的关系,我们以秦岭一野生川金丝猴种群为研究对象,通过长时间连续跟踪观察,完成了该种群中所有87只金丝猴的个体识别。用无损伤取样法采集了三只成年雌性川金丝猴不同季节的粪样,对粪样中的睾酮进行抽提测定,同时对采样个体的性行为进行记录。结果显示:1)成年雌猴在交配盛期与非交配盛期,粪样中睾酮水平均呈明显的周期性波动,交配盛期时睾酮水平的变化周期为24.56 d±4.07 d,非交配盛期时为43.67 d±2.89 d,显著长于交配盛期;2)成年雌猴在交配盛期的粪样睾酮水平为2.85 ng±1.12 ng/g,邀配频次为0.98±1.04次/d,非交配盛期时睾酮水平为1.71 ng±0.77 ng/g,邀配频次为0.28±0.53次/d,与交配盛期相比,非交配盛期的睾酮水平和邀配频次均明显降低;3)交配盛期各成年雌猴的粪样睾酮水平与邀配频次均有较显著的正相关关系(相关系数分别为0.631、0.683和0.659),而在非交配盛期这种相关关系并不存在(相关系数分别为0.091、0.493和0.205)。本研究表明,成年雌性川金丝猴体内睾酮水平、睾酮水平变化周期及邀配行为频次均具有明显的季节性差异,这种季节性差异可能是川金丝猴行季节性繁殖的内在原因和外在表现之一。本研究亦表明交配盛期雌猴的性行为频次在一定程度上可能受体内睾酮水平的控制,而在非交配盛期这种控制作用较弱。     

短尾猴和日本猴雄性性行为的比较研究

本文比较研究了短尾猴和日本猴的雄性性行为。短尾猴属单次爬跨射精型种类,每次交配平均持续23.2秒,日本猴属多次爬跨射精型种类,每次交配平均持续8.2分,交配期间雄性平均爬跨雌性lO.6次才能达到射精。短尾猴第1顺位雄性是群中的主要交配者,它占有交配总数的70.9 %;而日本猴第5顺位以下的雄性占交配总数的64%。交配中,两种猴雄性间相互打搅行为的发生频率大致相当。短尾猴高顺位雄性的打搅行为能使低顺位雄性的交配中断,但日本猴高顺位的打搅行为只能使低顺位雄性交配的54.3%中断。     

野生黄山短尾猴雄性睾酮的季节性变化

本文研究了黄山短尾猴野生群体雄性睾酮的变化规律.采用放射性免疫分析方法,对黄山短尾猴鱼鳞坑YA2群成年雄性个体在交配期(2005年10月至2006年1月;2006年7月至9月)与非交配期(2006年2月至6月)的粪便睾酮水平进行了测定,共采集到5只雄性个体的新鲜粪便样品426个.结果表明:在群体水平,黄山短尾猴成年雄性睾酮浓度交配季节(12.283±5.745 ng/ml)显著高于非交配季节(9.424±4.987 ng/ml),季节性变化显著(P<0.01);个体水平上,不同个体雄性睾酮均呈现显著的季节性变化(P<0.01);成年雄性睾酮分泌水平与生境温度变化呈显著正相关关系(P<0.05),且在交配季节初期雄性睾酮浓度最高,交配季节末期降到最低.研究结果支持短尾猴季节性繁殖的结论,且环境温度是影响短尾猴雄性睾酮浓度的可能因素.     

雄性不育对布氏田鼠交配行为和繁殖的影响

利用雄性结扎不育方法研究雄性结扎不育对布氏田鼠交配行为和繁殖的影响。结果表明：雄性结扎不育后不影响布氏田鼠雌雄两性的交配行为;不育雄性的存在对雌性的正常交配产生了干扰,降低了雌性的有效交配次数,并导致雌性怀孕率和产仔数的下降。结果一方面说明结扎不育雄性方法可用于布氏田鼠的不育控制研究,另一方面为不育后的竞争性繁殖干扰假设提供了实验证据。     

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.     

Quantitative observations of grooming behavior in free-rangingMacaca mulatta

Quantitative methods of observation and analysis were used in a 12-month study of grooming behavior of free-rangingMacaca mulatta on La Cueva Island at La Parguera, Puerto Rico. Observations lasting 30–120 minutes were made from eight positions on the island at a standard time of day when monkeys were either feeding or resting. Two dependent variables were obtained: (1) the number of monkeys present in the observation area were noted by age and sex class at five-minute intervals throughout each observation, and (2) the frequency of grooming encounters was tabulated by the age and sex class(es) of groomer and recipient. These data were computed as grooms/hour/possible interacting combination of monkeys. Grooming frequencies were higher in non-feeding situations than when monkeys were feeding. The largest social group had the lowest mean grooming rates, while the smallest group had the highest grooming frequencies. More grooming occurred during the November-to-February mating season than at other periods of the year. Adult females were involved in over 60% of all grooming behavior, juveniles participated in 25% of the grooming, while adult males groomed females, primarily during the mating season, and rarely groomed other males or juveniles. Genealogical relationships, levels of group aggression and the feeding or resting context all influenced the frequency of grooming. This study provides support for the hypothesis that the basic social unit for rhesus macaques consists of a core of adult females with their juvenile and infant progeny.     

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。     

Male Demography,Female Mating Behavior,and Infanticide in Wild Patas Monkeys (Erythrocebus patas)

Infanticide by males has been hypothesized to be a naturally selected behavioral strategy that increases the infanticidal male's reproductive success. The sexual selection hypothesis has been challenged via alternative, nonadaptive hypotheses that dispute its empirical and theoretical bases. Two of the most widely recognized alternatives are the social pathology hypothesis, in which infanticide results from overcrowding or recent human disturbance, and the generalized aggression hypothesis, in which infanticide is an epiphenomenon of increased male aggression. We report the first case of infanticide in wild, seasonally breeding patas monkeys (Erythrocebus patas) living at a low population density in a stable habitat, conditions which do not support the social pathology hypothesis. Its exceptional occurrence is consistent with the sexual selection hypothesis: over a 7-year period the infanticidal male was the only one of 13 resident males that was not present during the actual conception season but was present during the following birth season. Also consistent with this hypothesis, mothers were differentially targeted for male aggression, which increased sevenfold during the days surrounding the infanticide and then decreased to baseline levels after the infanticide. Aggression targeted at mothers does not support the generalized aggression hypothesis. As predicted by the sexual selection hypothesis, females began soliciting mating immediately after the infanticide, despite its occurrence in the nonconceptive season.     

Inbreeding avoidance in rhesus macaques: Whose choice?

Whether nonhuman primates avoid copulating with close kin living in their social group is controversial. If sexual aversion to relatives occurs, it should be stronger in females than in males because of females' greater investment in each offspring and hence greater costs resulting from less viable offspring. Data presented here show that adult male rhesus macaques breeding in their natal groups at Cayo Santiago experienced high copulatory success, but copulated less with females of their own matrilineages than with females of other matrilineages. Adult females were never observed to copulate with males of their own matrilineage during their fertile periods. Although natal males sometimes courted their relatives, examination of two measures of female mate choice showed that females chose unrelated natal males over male kin. Female aversion to male kin was specific to the sexual context; during the birth season, females did not discriminate against their male relatives in distributing grooming. Evolved inbreeding avoidance mechanisms probably produce different outcomes at Cayo Santiago than in wild rhesus macaque populations. Gender differences in sexual aversion to relatives may be partly responsible for differences between studies in reported frequency of copulations by related pairs. © 1993 Wiley-Liss, Inc.     

Social control of reproduction in banded mongooses

Recent theoretical work suggests that the distribution of reproduction, or degree of reproductive skew, in animal societies depends crucially on (1) whether dominant individuals can fully control subordinate reproduction, and (2) how subordinate reproduction affects the fitness of dominants. I investigated these two factors in cooperatively breeding banded mongooses, Mungos mungo. Female packmates entered oestrus together and were closely guarded by dominant males. These males were aggressive to subordinate males who attempted to mate, but females still managed to mate with males other than their mate guard. Older females were guarded and mated a few days before their younger packmates, yet all females usually gave birth on the same day, suggesting that older females may have a longer gestation period. Moreover, older females carried more fetuses. Overall, ca. 83% of adult females conceived in each breeding attempt and 71% carried to term. These results indicate that, among males, dominant individuals did not have full control over the mating attempts of subordinates (since they could not fully control the mating behaviour of the females they guarded), while among females there was little or no attempt to prevent subordinates from breeding (at least, prior to parturition). Two within-group infanticides by males suggested that some control over reproduction may be exercised postpartum. Per capita survivorship of young in the den increased with the number of females who gave birth. Thus, dominant females may benefit from subordinate reproduction, providing a possible explanation for the lack of reproductive suppression among females in this species. Copyright 2000 The Association for the Study of Animal Behaviour.     

ANNUAL REPRODUCTIVE CYCLE OF THE FEMALE HAWAIIAN MONK SEAL (MONACHUS SCHAUINSLANDI)

Abstract: The annual reproductive cycle is described for the adult female Hawaiian monk seal ( Monachus schauinslandi ) from data collected at Laysan Island (1982–1991) and Lisianski Island (1982–1983) in the Northwestern Hawaiian Islands. Pupping, lactation, weaning, and molting were directly observed, while mating was rarely observed and was, therefore, inferred from the occurrence of mounting injuries and from adult male and female association patterns. Pooled birth rates during the study period were 0.544 for all adult-sized females and 0.675 for females parous in earlier years. For parturient females, pupping peaked in late March and early April, weaning in May, mounting injuries in May and June, and molting in July. For non-parturient females, the median mounting injury and molting dates occurred 17 and 28 days earlier, respectively. Pupping date set the timing of subsequent events in the annual cycle, but the timing of those events was adjusted by loss of the pup or poor physical condition of the female. Individual pupping patterns varied widely. The mean interval for births in consecutive years was 381 days; females that pupped in consecutive years gave birth later each season. Conversely, females who skipped a year or more gave birth earlier their next pupping season.     

Seasonal breeding in rhesus monkeys: Influence of the behavioral environment

The accepted model of breeding seasonality in rhesus monkeys states that females become reproductively active in response to an environmental cue and that males become sexually active in response to ovulating females. This model must be modified to include direct responses of the male to the physical environment, endocrine responses of males to sexual activity, and responses of the female to the sexual activation of fellow group members. The complex set of social stimuli that influences the breeding readiness of both sexes may serve to delimit more precisely the annual periods of conception and birth than would be the case if each individual responded only to the changing physical environment.     

The development of affiliative and coercive reproductive tactics in male chimpanzees

Like many animals, adult male chimpanzees often compete for a limited number of mates. They fight other males as they strive for status that confers reproductive benefits and use aggression to coerce females to mate with them. Nevertheless, small-bodied, socially immature adolescent male chimpanzees, who cannot compete with older males for status nor intimidate females, father offspring. We investigated how they do so through a study of adolescent and young adult males at Ngogo in Kibale National Park, Uganda. Adolescent males mated with nulliparous females and reproduced primarily with these first-time mothers, who are not preferred as mating partners by older males. Two other factors, affiliation and aggression, also influenced mating success. Specifically, the strength of affiliative bonds that males formed with females and the amount of aggression males directed toward females predicted male mating success. The effect of male aggression toward females on mating success increased as males aged, especially when they directed it toward females with whom they shared affiliative bonds. These results mirror sexual coercion in humans, which occurs most often between males and females involved in close, affiliative relationships.