非人灵长类雌性等级的研究方法

在非人灵长类动物等级研究中, 有关判定与描述优势等级方法的选择至关重要。早期对旧大陆灵长类的等级研究主要关注雄性, 对雌性涉猎较少。目前主要应用攻击-屈服、取食、携婴、相互理毛等行为标准来研究灵长类的雌性等级, 也发现不同的物种往往有不同的雌性等级模式。依据社群结构的完整性将其分为线性等级与非线性等级, 并采用不同的方式进行描述。本文通过查阅文献资料, 对雌性等级的研究内容、判定标准以及描述方式进行论述, 以便抛砖引玉提高非人灵长类雌性等级的研究水平。     

非人灵长类性打搅行为研究进展

性打搅行为是指非人灵长类社群内非交配个体对正在发生性行为个体的干扰行为,该行为在非人灵长类中普遍存在,通过综述目前已知的36种非灵长类的性打搅行为的报道,探讨性打搅行为的功能和意义。性打搅行为具有更高度的种间和种内差异,种间的性打搅的方式不同、同时受到社会结构和婚配制度的影响;在种内受到性别、年龄、等级等因素的影响。结合川金丝猴性打搅的研究结果,从性打搅的类型、被打搅的反应、性打搅的功能以及相关假说等方面入手,对非人灵长类性打搅行为进行系统总结,旨在为我国相关领域的研究提供参考,为生殖行为学研究者启发思路,推动本领域的发展。     

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。     

非人灵长类的杀婴行为及其适应意义

杀婴现象普遍存在于非人灵长类种类中,本文通过综述目前已知的21 种非人灵长类的69 例杀婴行为记
录,探讨了非人灵长类杀婴行为的适应性意义。大多数雄性杀婴行为符合雄性繁殖策略假说,既满足以下3 个
条件:(1)杀婴雄性与受害婴仔没有父子关系;(2)失去婴猴的母亲会提前进入发情状态;(3)受害雌性随后
可能与杀婴雄性交配。不过有些个别的杀婴现象与雄性繁殖利益无关,文中进一步介绍了误伤假说、减少未来
竞争者假说、肉食假说和病态行为假说。杀婴无疑会对受害母亲造成巨大的损失。雌性防范杀婴的策略包括直
接抵抗、回避接触雄性、积极发情、依靠保护雄性和改变繁殖群组成等多种方式。但是雄性杀婴现象不会完全
消失,因为出现该行为的根本原因是雌性防范雄性杀婴的难度较大,而雄性实施杀婴行为的风险较低、利益较
高。     

非人灵长类的亲缘选择

亲缘选择是动物进化的重要研究领域之一,非人灵长类因具有丰富的社会网络,是亲缘选择研究领域的重要类群。动物进行亲缘选择的前提是亲缘识别,并常通过社会行为的亲缘偏向表现。因此,本文从非人灵长类的亲缘识别机制和亲缘关系对其社会行为的影响两方面进行了综述:熟悉性和表现型匹配是目前普遍认同的非人灵长类亲缘识别机制,同时这两种机制并不相互排斥,它们可能共同在灵长类的亲缘识别中起作用;在非人灵长类中,亲缘关系是影响社会行为模式的主导因子,它影响着多种灵长类个体的友好行为、攻击行为和性行为的选择,同时亲缘偏向行为在不同物种中表现不尽相同,说明亲缘选择理论可以部分解释灵长类的行为,但存在一定的局限性。本文分析了两种亲缘识别机制的异同以及在实际研究中利用亲缘选择理论解释非人灵长
类社会行为的局限及可能原因。目前,对非人灵长类社会中的亲缘选择研究正逐步深入,其中分子遗传学技术的应用是重要的推动力量。同时,依然存在诸如汉密尔顿规则参数估计和新大陆猴的亲缘选择研究案例的难点,有待研究者进一步探究。     

色彩信号在非人灵长类动物性选择中的作用

动物的多彩羽毛、毛发和皮肤不仅在种内交流、反捕食防御、体温调节等方面起到重要作用,而且在性选择中具有非常重要的意义。许多非人灵长类种类具有多色毛发和裸露皮肤,同时又拥有复杂多样的交配系统和视觉系统,因此成为研究色彩信号在性选择中作用的代表类群。本文阐述了性选择压力下颜色信号的功能以及非人灵长类的研究优势,综合分析了颜色信号在非人灵长类性选择中的作用：(1)异性间偏好鲜艳颜色,鲜艳颜色可能意味着身体更健康;(2)同性间颜色鲜艳意味着个体等级高低及竞争烈度强弱;(3)所有研究主要以山魈(Mandrillus sphinx)和猕猴(Macaca mulatta)为对象,且多以皮肤颜色为信号,涉及毛色的研究几乎空白;(4)尽管视觉模型比光谱法及数字摄影更精准,但在现有条件下数字摄影更加方便可行;(5)表型颜色与内在生理机制联系及相关基因研究还需要进一步深入挖掘。     

灵长类社会玩耍的行为模式、影响因素及其功能风险

社会玩耍是指两个或两个以上个体共同参与的一种互作性玩耍行为,个体间的行为彼此适应并相互影响。社会玩耍行为在灵长类物种的社会交往过程中普遍发生,作为未成年个体一种重要的发育行为,其对个体的生存技能和成年后的繁殖成功具有重要影响。灵长类物种的社会玩耍不仅仅表现为追逐、摔跤、跳跃等一些常见行为,部分物种还发展出自己特有的行为。一般而言,社会玩耍在婴儿后期和青少年早期的发生频率最高,然后随着年龄增长直到成年时期,这类行为的平均发生频率将逐渐下降。未成年雄性个体要比同年龄段的雌性个体更喜欢玩耍,但常因物种、研究对象年龄等因素表现不同甚至相反;很多物种的个体喜欢与有亲缘关系的个体玩耍,或者与性别相同、年龄相仿、等级相近的个体玩耍。总之,非人灵长类个体社会玩耍的发育不但受环境参量如食物、场地等的影响,而且还与个体的年龄、性别、等级、亲缘关系等社群因素紧密相关。未成年个体在玩耍过程中,获得了身体机能的快速发育、完善了生存技能、建立了个体间的友好关系、增强了认识自身及适应周围环境的能力,从而为顺利过渡到成年期和履行自己的职能打好基础,但有时却需要承担玩耍过程中受伤甚至死亡的风险。玩耍作为灵长类社会的一种行为文化,对其研究有助于对人类自身行为进化的不断认识,相信这方面的理论将会得到后来者的不断创新和丰富,也期望这方面的理念及经验能被及时运用到保护繁育等实践活动中。     

太行山猕猴的性打搅行为

性打搅是指动物在交配期间受到同种其他个体化学信息素、面部表情、声音、攻击等方面干扰的现象。性打搅行为作为非人灵长类繁殖行为生态研究的重要话题,具有高度的种间和种内差异,在种内主要受到性别和年龄等因素的影响。本研究于2021年9月至2022年8月在太行山猕猴国家级自然保护区五龙口地区,以太行山猕猴(Macaca mulatta tcheliensis) WLK-1A群为研究对象,采用焦点动物取样法和连续记录法观察并记录目标个体的交配行为,以期探讨太行山猕猴的性打搅行为及其性别-年龄组差异。结果表明：(1)太行山猕猴的性打搅行为仅发生在交配期,高峰期为11月;(2)从性打搅的发起者来看,雄性(成年和亚成年)个体占29.1%,雌性(成年和亚成年)个体占14.9%,青少年个体占7.5%,婴幼个体占48.5%;(3)与接触式性打搅相比,雄性(P=0.011)和雌性(P=0.033)的性打搅均以非接触式性打搅为主,青少年个体的非接触式与接触式性打搅之间无显著差异(P=0.334),婴幼猴以接触式性打搅为主(P=0.001);(4)雄性倾向于打搅交配中的雌性(P=0.040),雌性对交配中雄性与雌性...     

笼养雌性短尾猴月经周期及性行为

在多雄多雌的灵长类社会中,成年雌雄个体的性行为并不局限于雌性受孕期,但仍认为性行为在受孕期最为活跃。对于雌性隐藏发情的灵长类物种,雌雄个体的性行为与雌性月经周期的相关性尚不清楚。本研究以短尾猴(Macaca thibetana)为研究对象,研究雌性隐藏发情灵长类的性行为与月经周期阶段的关系。在短尾猴繁殖季节,从2019年10月2日至2020年1月14日连续采集了安徽省铜陵市动物园6只成年雌性短尾猴的490份粪便样本,采用放射免疫分析法检测粪便中雌二醇激素的浓度,并用焦点动物取样法和行为取样法采集目标个体(4只雌性、4只雄性)的性行为数据。本研究利用Spearman相关系数检验雌激素与性行为的相关性;独立样本T检验分析雌性短尾猴发情的邀配性指标(雌性呈臀行为)、吸引性指标(雄性性检查行为)和交配行为在月经周期三个阶段(受孕期前:受孕期前的5 d;受孕期:最有可能排卵的2 d加上精子能在雌性生殖道中存活的3 d;受孕期后:受孕期后的5 d)的分布差异性。结果表明,雌性短尾猴的雌二醇浓度在繁殖季节呈现周期性波动,周期长度为(31.1±1.8)d(n=7,范围23～37 d);同时在月经周期内,雌二醇浓度与雌性呈臀行为呈显著负相关(r=﹣0.616,n=15,P=0.015),而与交配行为和雄性性检查行为均无显著相关性;雌性呈臀行为的平均频次主要集中在受孕期前(T=﹣1.215,df=12,P=0.044);交配行为和雄性性检查行为在月经周期的三个阶段无显著性差异。短尾猴的性行为并没有集中在雌性的受孕期,说明雌激素对短尾猴的性行为无严格控制作用,且雄性短尾猴无法准确识别雌性的受孕期。     

非人灵长类和解行为的研究方法与内容

对近年来有关非人灵长类和解行为的研究方法和内容进行了综述。和解行为在非人灵长类中普遍存在,并且有特定的行为模式,要证明它的存在需做匹配观察。分析表明,在潜在合作伙伴中,亲缘关系较近、统治风格缓和、等级距离接近的个体之间,以及一方携带婴儿的雌性个体与其它个体之间,其和解倾向较高,反之和解倾向较低。攻击强度与和解倾向无关,和解行为的发起者因种而异。     

Observations on the evolution of the genitalia and copulatory behaviour in male primates

Comparative studies of genital anatomy and sexual behaviour in male primates show that penile morphology and copulatory patterns tend to be more specialized in species which have a multimale or dispersed (non-gregarious) mating system. The penis may be longer and more complex morphologically in such species. Copulatory patterns involving a series of intromissions or prolonged single intromissions are more common than in species with a monogamous or polygynous mating system. Elongation of the baculum occurs in species which prolong intromission into the post-ejaculatory period. Sexual selection may have favoured the evolution of such features in species where females mate with a number of males rather than with a single partner. Relative testis size is also greatest under such conditions and larger testes occur in some non-gregarious or seasonally breeding prosimians, as well as in anthropoids with multimale mating systems.     

Primate frugivory in Kibale National Park, Uganda, and its implications for human use of forest resources

In an attempt to understand the practical and/or economic implications of primate seed dispersal, it was established which seed species are dispersed by frugivorous primates in Kibale National Park, Uganda, and which of this sort of species were used by Ugandan people. A list of fruit species consumed by Kibale primates was compiled using primary data and by reviewing all known published accounts of their fruit diet. Primates consume the fruit of 87 Kibale forest tree species; the seeds of 11% of these species are destroyed by the primates. The remaining 77 species are dispersed by either one, two, three or all four of the frugivorous Kibale primates. Of these 77 species, 42% have some utility to local Ugandan inhabitants, suggesting that maintaining populations of primates is important not only for natural forest regeneration, but also for human habitat use. This report illustrates the complexity of the seed dispersal process and suggests links not only between plants and their dispersers, but also between sets of plants/dispersers and the human populations that rely on forest resources.     

Big-bodied males help us recognize that females have big pelves

Schultz ([1949] Am. J. Phys. Anthropol. 7:401-424) presented a conundrum: among primates, sexual dimorphism of the pelvis is a developmental adjunct to dimorphism in other aspects of the body, albeit in the converse direction. Among species in which males are larger than females in body size, females are larger than males in some pelvic dimensions; species with little sexual dimorphism in nonpelvic size show little pelvic dimorphism. Obstetrical difficulty does not explain this relationship. The present study addresses this issue, evaluating the relationship between pelvic and femoral sexual dimorphism in 12 anthropoid species. The hypothesis is that species in which males are significantly larger than females in femoral size will have a higher incidence, magnitude, and variability of pelvic sexual dimorphism, with females having relatively larger pelves than males, compared with species monomorphic in femoral size. The results are consistent with the hypothesis. The proposed explanation is that the default pelvic anatomy in adulthood is that of the female; testosterone redirects growth from the default type to that of the male by differentially enhancing and repressing growth among the pelvic dimensions. Testosterone also influences sexual dimorphism of the femur. The magnitude of the pelvic response to testosterone is greater in species that are sexually dimorphic in the femur than in those that are monomorphic.     

Multiple speciation events in an arthropod with divergent evolution in sexual morphology

Sexual selection can facilitate divergent evolution of traits related to mating and consequently promote speciation. Theoretically, independent operation of sexual selection in different populations can lead to divergence of sexual traits among populations and result in allopatric speciation. Here, we show that divergent evolution in sexual morphology affecting mating compatibility (body size and genital morphologies) and speciation have occurred in a lineage of millipedes, the Parafontaria tonominea species complex. In this millipede group, male and female body and genital sizes exhibit marked, correlated divergence among populations, and the diverged morphologies result in mechanical reproductive isolation between sympatric species. The morphological divergence occurred among populations independently and without any correlation with climatic variables, although matching between sexes has been maintained, suggesting that morphological divergence was not a by-product of climatic adaptation. The diverged populations underwent restricted dispersal and secondary contact without hybridization. The extent of morphological difference between sympatric species is variable, as is diversity among allopatric populations; consequently, the species complex appears to contain many species. This millipede case suggests that sexual selection does contribute to species richness via morphological diversification when a lineage of organisms consists of highly divided populations owing to limited dispersal.     

How intraspecific variation in seed‐dispersing animals matters for plants

Seed dispersal by animals is a complex phenomenon, characterized by multiple mechanisms and variable outcomes. Most researchers approach this complexity by analysing context‐dependency in seed dispersal and investigating extrinsic factors that might influence interactions between plants and seed dispersers. Intrinsic traits of seed dispersers provide an alternative way of making sense of the enormous variation in seed fates. I review causes of intraspecific variability in frugivorous and granivorous animals, discuss their effects on seed dispersal, and outline likely consequences for plant populations and communities. Sources of individual variation in seed‐dispersing animals include sexual dimorphism, changes associated with growth and ageing, individual specialization, and animal personalities. Sexual dimorphism of seed‐dispersing animals influences seed fate through diverse mechanisms that range from effects caused by sex‐specific differences in body size, to influences of male versus female cognitive functions. These differences affect the type of seed treatment (e.g. dispersal versus predation), the number of dispersed seeds, distance of seed dispersal, and likelihood that seeds are left in favourable sites for seeds or seedlings. The best‐documented consequences of individual differences associated with growth and ageing involve quantity of dispersed seeds and the quality of seed treatment in the mouth and gut. Individual specialization on different resources affects the number of dispersed plant species, and therefore the connectivity and architecture of seed‐dispersal networks. Animal personalities might play an important role in shaping interactions between plants and dispersers of their seeds, yet their potential in this regard remains overlooked. In general, intraspecific variation in seed‐dispersing animals often influences plants through effects of these individual differences on the movement ecology of the dispersers. Two conditions are necessary for individual variation to exert a strong influence on seed dispersal. First, the individual differences in traits should translate into differences in crucial characteristics of seed dispersal. Second, individual variation is more likely to be important when the proportions of particular types of individuals fluctuate strongly in a population or vary across space; when proportions are static, it is less likely that intraspecific differences will be responsible for changes in the dynamics and outcomes of plant–animal interactions. In conclusion, focusing on variation among foraging animals rather than on species averages might bring new, mechanistic insights to the phenomenon of seed dispersal. While this shift in perspective is unlikely to replace the traditional approach (based on the assumption that all important variation occurs among species), it provides a complementary alternative to decipher the enormous variation observed in animal‐mediated seed dispersal.     

Sexual dimorphism in postcranial skeletal shape suggests male-biased specialization for physical competition in anthropoid primates

Sexual dimorphism often arises as a response to selection on traits that improve a male's ability to physically compete for access to mates. In primates, sexual dimorphism in body mass and canine size is more common in species with intense male–male competition. However, in addition to these traits, other musculoskeletal adaptations may improve male fighting performance. Postcranial traits that increase strength, agility, and maneuverability may also be under selection. To test the hypothesis that males, as compared to females, are more specialized for physical competition in their postcranial anatomy, we compared sex-specific skeletal shape using a set of functional indices predicted to improve fighting performance. Across species, we found significant sexual dimorphism in a subset of these indices, indicating the presence of skeletal shape sexual dimorphism in our sample of anthropoid primates. Mean skeletal shape sexual dimorphism was positively correlated with sexual dimorphism in body size, an indicator of the intensity of male–male competition, even when controlling for both body mass and phylogenetic relatedness. These results suggest that selection on male fighting ability has played a role in the evolution of postcranial sexual dimorphism in primates.     

Sexual Selection in the Loud Calls of Male Primates: Signal Content and Function

Researchers have used sexual selection theory and hypotheses based on intersexual mate choice and intrasexual mate competition to explain the role of spontaneous long-distance vocalizations emitted by adult male primates, relying on the tacit assumption that assessment or identity cues are encoded in the vocalizations. I review the published literature and aim to substantiate a relationship between sexual selection and long-distance vocal communication in primates. First, I review findings from nonprimate taxa to determine the relative importance of inter- and intrasexual selection and to provide a background for examining primates. Next, I describe several hypotheses for signal content and function in adult male loud calls. Then, I examine the available data across Primates for evidence to support or to refute these hypotheses and to determine if they meet proposed criteria for demonstrating sexual selection [Snowdon, C. T. (2004). Sexual Selection in Primates: New and Comparative Perspectives]. Signal content refers to patterns of acoustic features within vocalizations from which listeners might extract cues or information about the signaler. I interpret signal function, in turn, from behavioral responses of receivers and assume it has ultimate effects on the evolution and design of acoustic signals if direct fitness consequences exist. After the general review across primates, I propose orangutans as a candidate species for further evaluation of sexual selection in vocal communication. The available evidence corroborates a demonstrable relationship between sexual selection and adult male loud calls based on individual recognition, but it is necessary to obtain additional data to affirm a direct benefit to reproductive success.     

Seed Dispersal by Primates in Asian Habitats: From Species,to Communities,to Conservation

Primates are among the most important seed dispersers in the habitats they occupy. Understanding the extent of, and gaps in, our knowledge of seed dispersal by Asian primates is essential, because many of these primates are extremely vulnerable to anthropogenic disturbance. In this review, I show how initial studies focused on the role of individual species in seed dispersal have expanded more recently to consider their role in the wider frugivore community. There are five functional groups of primate seed dispersers in Asia; most of our information comes from the (usually) highly frugivorous macaques and gibbons, while our understanding of the roles played by orangutans and, especially, colobines and lorises remains rudimentary. Preliminary community-wide studies suggest a pivotal role for gibbons and macaques in frugivore communities, with higher dispersal overlap with other mammals than with birds. The gaps in our knowledge are plentiful, however, including understanding fruit selection in detail, determining how seed dispersal roles might change across different habitats, evaluating the balance between mutualisms and antagonisms in orangutans and macaques, describing postdispersal processes, and documenting how habitats are impacted by changes in primate abundance and behavior.     

Sexual signalling in female crested macaques and the evolution of primate fertility signals

ABSTRACT: BACKGROUND: Female signals of fertility have evolved in diverse taxa. Among the most interesting study systems are those of multimale multifemale group-living primates, where females signal fertility to males through multiple signals, and in which there is substantial inter-specific variation in the composition and reliability of such signals. Among the macaques, some species display reliable behavioural and/or anogenital signals while others do not. One cause of this variation may be differences in male competitive regimes: some species show marked sexual dimorphism and reproductive skew, with males fighting for dominance, while others show low dimorphism and skew, with males queuing for dominance. As such, there is variation in the extent to which rank is a reliable proxy for male competitiveness, which may affect the extent to which it is in females' interest to signal ovulation reliably. However, data on ovulatory signals are absent from species at one end of the macaque continuum, where selection has led to high sexual dimorphism and male reproductive skew. Here we present data from 31 cycles of 19 wild female crested macaques, a highly sexually dimorphic species with strong mating skew. We collected measures of ovarian hormone data from faeces, sexual swelling size from digital images, and male and female behaviour. RESULTS: We show that both sexual swelling size and female proceptivity are graded-signals, but relatively reliable indicators of ovulation, with swelling size largest and female proceptive behaviours most frequent around ovulation. Sexual swelling size was also larger in conceptive cycles. Male mating behaviour was well timed to female ovulation, suggesting that males had accurate information about this. CONCLUSION: Though probabilistic, crested macaque ovulatory signals are relatively reliable. We argue that in species where males fight over dominance, male dominance rank is surrogate for competitiveness. Under these circumstances it is in the interest of females to increase paternity concentration and assurance in dominants beyond levels seen in species where such competition is less marked. As such, we suggest that it may be variation in male competitive regimes that leads to the evolution of fertility signalling systems of different reliability.