人工辅助投食滇金丝猴一雄多雌单元之间的等级序列

群居灵长类动物等级序列是长期进化的结果,也是动物行为生态学研究的热点。灵长类动物个体间通过等级序列,合理的利用自然资源、避免过多的伤亡、保护群体内的幼弱者,从而更好地适应复杂的生态环境。2013年6月至2014年1月,采用焦点动物取样法对白马雪山国家级自然保护区人工辅助投食滇金丝猴群的6个一雄多雌单位之间等级序列进行了研究。通过对一雄多雌单元之间攻击-屈服行为数据分析发现：一雄多雌单位之间存在明显的等级序列（大个子单元＞单疤单元＞红脸单元＞联合国单元＞偏冠单元＞花唇单元）;单元之间成年雄性的攻击行为多于成年雌性;单位之间的等级序列与取食次数、第一序位取食总次数呈显著正相关,与各单元取食总时间无明显相关性;同时,一雄多雌单元之间的等级序列与单元内成年雌性数量呈显著正相关。研究结果符合群居灵长类动物攻击-屈服假说和资源优先占据-雌性高序列偏好假说。     

岷山山系川金丝猴群的社会结构——以唐家河国家级自然保护区川金丝猴为例

为进一步深入理解不同地理区域川金丝猴群的社会结构特点,于2014年3—7月,采用焦点动物取样法对岷山山系唐家河国家级自然保护区辅助投食成功的一个川金丝猴群进行了观察。研究表明:该猴群共有138只个体,包括16只成年雄性,48只成年雌性,11只亚成年雌性,3只亚成年雄性,36只青少年猴以及24只婴幼猴;猴群中成年个体与未成年个体的比例为0.86∶1。唐家河川金丝猴群具有典型的重层社会结构:即由11个一雄多雌单元(one-male units,OMUs)组成2个繁殖分队(breeding band),并分别与1个密切联系的全雄单元(all-male unit,AMU)共同构成。其中,每个OMU分别由6~15只个体组成,平均为10.64只±2.77只;AMU共21只个体,以青少年个体最多,占61.9%。方差分析表明:2个繁殖分队中的OMU大小差异无统计学意义(P>0.05)。     

中国云南塔城滇金丝猴的社会结构

在云南塔城附近的山地森林中,我们利用摄象机记录了一群滇金丝猴通过山谷和在水源地饮水的过程。我们利用这些记录分析这群滇金丝猴的种群大小、组成成员以及个体的空间分布,并利用连续通过个体之间的时间间隔来揭示社会结构。塔城滇金丝猴群由366个个体组成,是目前所知最大的滇金丝猴群体。猴群内雌性个体较多。猴群可以分为26个单位,其中有19个单雄多雌单位、5个多雄多雌单位和2个全雄单位。统计结果表明单位内个体间的时间间隔显著比单位间的短。2/3以上的亚成年雄性个体和全雄单位一起活动。猴群在地面活动时全雄单位位于猴群的外围,充当前卫和后卫。塔城猴群的生命表组成结构与吾牙普亚猴群的大致一样,同样是多层社会结构,单雄多雌和全雄单位构成猴群的主体。亚成年雄性多数与全雄单位一起活动,表明雄性可能会从其出生的单雄多雌单位中迁移出来。     

扩散或不扩散：雄性太行山猕猴繁殖成功比较研究

典型的猕猴（Macaca multta）社群为多雄多雌型,雌性留群并形成母系单元,雄性多在亚成年或成年期离开出生群,而群内成年雄性多为外部迁入个体。雄性的迁出被认为可降低近亲繁殖或提升繁殖成功。然而,诸多野外数据显示,少数本群出生雄性个体会居留于出生群一年或数年。尽管驱动雄性离群的因素较复杂（如社会关系、近亲回避、繁殖成功）,但繁殖成功的差异可能是驱动雄性离群的主导因素。为探讨居留于出生群是否影响雄性的繁殖成功,于2010年3月至2014年1月,在太行山猕猴国家级自然保护区王屋山地区,以一群野生太行山猕猴为研究对象,采用非损伤取样法并结合分子生物学方法,分析了群内出生和迁入成年雄性个体的繁殖成功（以子代数量评估）。研究发现：(1)群内51只子代个体中有36只个体可以匹配到其遗传学父亲;(2)4个迁入雄性繁衍了34个子代,仅1只群内出生雄性ZM繁衍了2只子代,但群内出生的雄性BB未匹配到子代。本研究提示,迁入雄性较本群出生雄性的繁殖成功较高,即迁移有助于繁殖成功的提升。     

野生太行山猕猴的社会结构

2009年2月至2010年7月,在河南太行山猕猴国家级自然保护区济源管理局所辖的天坛山管护区,基于个体识别和野外跟踪,观察了一个野生太行山猕猴群(王屋1群, WW-1)的社会结构,采用随机取样法(Ad libitum sampling)记录了该群内成年个体之间、母系单元之间的竞争行为过程,进而依据"David得分法"构建了WW-1群内成年个体及母系单元间的优势等级, 并采用瞬时扫描取样法(Instantaneous andscan sampling)记录了成年个体到达投食区的移动顺序, 分析了平均移动序位与社会顺位的相关性。结果表明: 1)WW-1群体大小为41只个体, 由7个成年雄性和13个成年雌性以及21个未成年猕猴组成, 群内的成年雌性个体分别隶属于3个母系单元; 2)群中成年雄性、成年雌性、亚成年雄性、亚成年雌性、青少年雄性、青少年雌性、婴幼猴雄性、婴幼猴雌性的比例为1∶1.86∶0.29∶0.43∶0.86∶1.29∶0.14∶0, 且未成年个体占全群的51.2%; 3)WW-1群表现出严格的优势等级结构, 成年个体优势顺位由高到低依次是: 豁鼻>次红>白鼻>痞子>红脸>尖脸>小白脸>皱眉>光鼻>长毛>黑颊>小不点>斑点>斑眼>灰头, 母系单元间的优势顺位由高到低依次是红脸单元>长毛单元>斑点单元。研究结果提示, 太行山猕猴的社会结构为多雄多雌型; 成年雄性个体之间优势等级较成年雌性严格, 而成年雌性的社会顺位受所在母系单元社会顺位的影响; WW-1群的α位为成年雄性, 其在获取资源上具有优先性。     

短尾猴的种群生物学

短尾猴是高等灵长类动物,中国特有物种。是多雄多雌群体,猴群中性成熟雌雄性比接近于1：1,雌性略多于雄性。虽然雌性定居、雄性移民,但大多数雄性的迁出发生在成年以后,而不是成年之前。社会和生态因素在调节猴群的数量中都有重要作用。雄性个体之间具有复杂的行为,雄猴之间频繁交往,以减少群体的紧张。猴群内具有严格的社会等级关系,年轻雄猴是群体的最高统治者。     

中国怒江片马地区怒江金丝猴种群动态及社会组织初探

2011 年10 月至2013 年10 月在云南高黎贡山自然保护区泸水县片马镇地区对一群怒江金丝猴的社会组织进行研究,基于野外实地记录与获得的视频资料统计猴群的个体数量,研究群数量初步估计为100 只。根据体型、毛色及是否携带婴猴等特征来划分所有个体的性别年龄组,依据单元内个体通过林窗时移动的时间间隔比单元间短的原则来划分不同繁殖单元及全雄群。所有视频中共发现22 个一雄多雌单元(OMU) 和1 个全雄群(AMU),因此怒江金丝猴的社会组织与其他金丝猴物种相似,由多个一雄多雌及其后代组成繁殖单元(OMUs)与雄性个体组成全雄群(AMU)组成重层社(Modular society);另外基于GPS 位点初步确定猴群活动的海拔范围为1 900 - 3 700 m,猴群生境面积约为12 km² ,初步获得猴群的密度为8 只/ km² ,比缅甸的种群密度(1 只/ km² )大得多。基于种群性别年龄计算种群动态参数,雄雌比为1∶ 2.1, 婴雌比为1∶ 4. 7,成幼比为2.5∶ 1。与滇金丝猴种群相比,该种群有很低的婴雌比和很高的成幼比。     

繁殖盛期雄性普通东方小蛙的个体大小不影响交配成功率

通过比较抱对和非抱对普通东方小蛙(Crinia signifera) 的吻尾干骨长(体长) 和检测个体较大或状态较好的雄体是否为成功繁殖个体, 检测雌蛙体长与交配成功的雄蛙的体长和相对状态(体重/体长) 之间是否存在线形关系, 评估交配与个体大小之间的关系。在实验室内, 令未交配雌体选择随机选出的雄体, 观测雌体是否与最大的雄体交配; 将抱对的蛙暴露于其它雄蛙, 观测其它较大或较小的雄蛙是否取代已抱对的雄蛙。结果表明: 抱对雄蛙和雌蛙的个体大小无显著的关系, 成功交配的雄蛙并不比未交配的雄蛙大, 其它雄蛙不能取代已抱对的雄蛙。在这一C. signifera种群中, 繁殖盛期雄性个体的大小似乎不影响交配成功率, 在此期间即便存在配偶选择, 亦决定于与雄性个体大小无关的其它因素。在更长的繁殖期内, 雄性个体大小则与交配成功率有关, 这是该种的典型特征。     

非人灵长类重层社会中一雄多雌体系的分化

重层社会是指社群内个体通过两个或多层面的纽带关系将彼此维系在一起形成的社会模式,是非人灵长类进化中最为复杂的一种社会结构,但仅见于狮尾狒、埃及狒狒和我国的金丝猴中。这些物种虽与其它叶猴类一样以一雄多雌社会单元作为社群的基本单位,但单元间不再具有领域性排斥对方而是相互聚集结合在一起形成大群;这样复杂的社群内个体如何稳定有序地组织在一起,长期以来受到广泛的关注。从现有对埃及狒狒和狮尾狒社会系统的研究看来,虽然这些物种具有相似的社会结构,但社群内个体间的组织模式与社会动力却截然不同。研究重层社会一雄多雌制体系的多样性,有助于我们了解物种利用社会组织适应环境变化的具体过程。但对于我国金丝猴重层社会组织机制的了解仍不充分,而成为当前研究的热点之一。本文从行为生态学的角度,阐述了现生灵长类重层社会物种在个体扩散机制、雄性替换模式、个体间联盟与繁殖策略等方面的差异,介绍了母系与非母系一雄多雌社会系统的经纬,综述了各物种维持社会稳定的动力与形成原因。旨在为研究者对我国仰鼻猴属物种社会组织与结构方面的研究启发思路,促进我国灵长类行为生态学理论体系的发展,并为保护金丝猴这一濒危物种提供科学依据。     

Adult male replacement in a group of silvered leaf-monkeys (Presbytis cristata) at Kuala Selangor,Malaysia

During observations of silvered leaf-monkeys at Kuala Selangor, Malaysia, the resident adult male in one group was replaced by an immigrant male. Three months after the replacement, all of the dependent infants in the group disappeared. The similarity of these observations to observations of male-takeovers and subsequent infanticide among Indian and Ceylonese langurs suggests a similar pattern of social change among all langurs which live in one-male groups.     

Female sexual swellings in the Asian colobine Simias concolor

During a preliminary study of pig-tailed langurs (Simias concolor) in the Pagai Islands, Indonesia, it was discovered that females exhibited conspicuous swelling of the urogenital triangle. The pig-tailed langur is the first Asiatic colobine found to have prominent sexual swellings and the only colobine with sexual swellings that lives in one-male groups. Because all anthropoids with conspicuous sex skin typically live in groups having female-biased adult sex ratios, it is possible that females might compete amongst themselves for the male in one-male groups, or the best males in multimale groups. Sexual swellings may therefore have resulted from sexual selection for signals attractive to males in female-female competition, as suggested earlier by Bercovitch (California Anthropologist 8:9–12, 1978). Prolonged observation of recognizable individuals will be required to test this hypothesis in the pig-tailed langur and other species.     

Take-over and infanticide in South Indian Hanuman langurs (Presbytis entellus)

A take-over and infanticide were observed in a South Indian population of Hanuman langurs (Presbytis entellus). A one-male, bisexual troop of langurs (Troop B1) was attacked by a band of two adult males and one subadult male. During the take-over, the resident male and one elderly female were driven from the troop. The youngest infant, about two months old, was fatally wounded by an attack from the dominant incoming male (S1). However, other young infants in Troop B1 were left unharmed. All three incoming males remained in Troop B1 but the troop appeared to remain functionally one-male, as S1 did not allow the other adult male to copulate. There was no sign that S1 attempted to eject the other two males from troop B1. © 1993 Wiley-Liss, Inc.     

My primate studies

In this paper, I describe my 62 years in primatology focusing on some of the key findings from fieldwork conducted in Japan, India, and Africa. My first study on nonhuman primates described in detail the division of a troop of Japanese macaques at Takasakiyama. After that, I had an opportunity to work on Hanuman langurs at Dharwar, India. These langurs lived in one-male, multi-female groups. This type of group structure was maintained through takeovers by all-male parties. The adult male and all juvenile males were chased out of the group. By this process, the one-male, multi-female group system was maintained. The incoming adult male bit and killed all infants in the group. Mothers who lost their infants went into estrus and mated with the newly arrived male. For many years, scientists ignored these events or ruled them out as abnormal behavior. My work on Japanese macaques suggested that concentrated resources created by artificial feeding exaggerated dominance rank hierarchies among individuals, whereas it is comparatively relaxed in the natural environment. I also investigated the population dynamics of a troop and the life histories of individuals. From these studies, I documented the frequency of twin births, the carrying of dead infants by mothers, and the occurrence of physical malformations. These observations were made possible through artificial feeding, revealing the merits and demerits of this approach. I pointed out that authors and journal editors must be careful to acknowledge important elements of the environment where studies are conducted, and these should be described when reporting results in scientific articles. My studies of chimpanzees were conducted at Bossou, Guinea. I suggested that there are males who lived outside of bisexual groups. Chimpanzees in this population made and used many kinds of tools. Some of them were observed only at Bossou, and a few were only discovered 20 years after the establishment of Bossou as a research site. After decades of research on tool use in this species, I also suggested that there are cultural zones throughout the geographic distribution of chimpanzees.

     

Determinants of female dispersal in Thomas langurs

Female dispersal occurs in a number of primate species. It may be related to: avoidance of inbreeding, reduction in food competition, reduction of predation risk, or avoidance of infanticide in combination with mate choice. Female dispersal was studied for a 5-year period in a wild population of Thomas langurs (Presbytis thomasi) that lived in one-male multi-female groups. Juvenile and adult individuals of both sexes were seen to disperse. Females appeared to transfer unhindered between groups, mostly from a larger group to a recently formed smaller one. They transferred without their infants and when not pregnant, and seemed to transfer preferentially during periods when extra-group males were harassing their group. During these inter-group encounters extra-group males seemed to try to commit infanticide. Thus, the timing of female transfer was probably closely linked to infanticide avoidance. Moreover, females seemed to transfer when the resident male of their group was no longer a good protector. The observations in the present study suggest that females transferred to reduce the risk of infanticide. Female dispersal may have another ultimate advantage as well, namely inbreeding avoidance. Due to the dispersal of both females and males the social organization of Thomas langurs was rather fluid. New groups were formed when females joined a male; male takeovers were not observed. Bisexual groups had only a limited life span, because all adult females of a bisexual group could emigrate. This pattern of unhindered female dispersal affects male reproductive strategies, and in particular it might lead to infanticidal behavior during inter-group encounters. Am. J. Primatol. 42:179–198, 1997. © 1997 Wiley-Liss, Inc.     

Male competition and coalitions in langurs (Presbytis entellus) at Jodhpur,Rajasthan, India

During a 15 month study on free ranging langurs (Presbytis entellus) at Jodhpur, Rajasthan, India, 5 adult male replacements were observed as a result of nontroop male invasions into the home ranges of 3 neighbouring one-male troops comprising 16–28 members each. Jodhpur langurs have no breeding season. Periods of instability during resident male changes lasted 11–119 days. Linear dominance hierarchies could be detected within the 3 main rival male bands of 2, 5, and 28–35 members. The respective alphas drove their allies away after their bands succeeded cooperatively at occupying a troop. During gradual replacements interim residencies alternated with multi-male stages. A large band's alpha may have had better chances to win the competition, since adult and nonadult allies functioned as “buffers” in agonistic encounters. The role of kin selection in structuring the composition of male bands and male coalitional behaviour cannot yet be quantified. Tactical “deceit” of powerful males to cause unrealistic expectations and in this way agonistic engagement of less strong males can be ruled out. “Sneaking copulations” is a proximate advantage for subordinate supporters, since they participated in 61.9% of all sexual interactions. Female promiscuity might reflect a strategy to induce male-male competition and thus select for a strong resident.     

黑长臂猿（Hylobates concolor）的配偶制及其与行为、生态和进化的关系

黑长臂猿是猿类中野外行为生态研究最少的一种,因而对其社群结构及配偶体制一直有着很大争议．本文基于近两年在无量山和哀牢山１０个月的野外工作,着重阐述了黑长臂猿的配偶体制,我们认为黑长臂猿是以一夫一妻和一夫多妻（至少是一夫两妻）的形式存在,并与其行为、生态及系统发育相关联．黑长臂猿的系统发育地位、成年雌性的领域性及群体的领域性不强和群体的领域较其它长臂猿类大得多可能是形成一夫多委的原因;而黑长臂猿在中国各分。布区的生境均受到不同程度的破坏,这在一定程度上也影响着其社群结构,特别是狩猎有着更直接的影响,活动范围的限制等,可能是形成一夫一妻的原因．     

Paternity analysis of alternative male reproductive routes among the langurs (Semnopithecus entellus) of Ramnagar

Because primate males usually invest very little in offspring, male reproductive success will mainly be determined by access to fertile females with differences in access leading to differential male reproductive success. To determine the outcome of alternative male reproductive routes, we investigated a wild population of Hanuman langurs at Ramnagar, South Nepal, where groups were either one-male or multimale. Paternity was established by DNA analysis from faeces for 42 infants in five groups. In one-male groups all infants were sired by the only resident male in the group. In multimale groups the alpha male fathered significantly more infants (57%) than all other resident males. Nonresident males sired at least 21% of all infants born into multimale groups. Because of the lower mean number of infants sired by alpha males during their whole tenure as alpha compared with males in one-male groups (2.3 versus 6.8) and the higher maximum value (8.9 versus 6.0), the alpha male route is considered the riskier option. Based on demographic data we suggest that the considerable variance in short-term reproductive success might not translate to the same degree into differences in long-term reproductive success. Copyright 2001 The Association for the Study of Animal Behaviour.     

Seasonality of group size,feeding, and breeding in wild red‐shanked douc langurs (Lao PDR)

In Asian colobines, small one‐male groups (OMG) seem to predominate alongside all‐male groups (AMG), while larger multimale groups (MMG) are rare, but are reported for Hanuman langurs and red‐shanked douc langurs. Recently, however, it has been speculated that the genus Pygathrix could have multilevel societies based on (1) a theoretical extension of the multilevel societies found in Rhinopithecus to all odd‐nosed colobines and (2) first data for black‐shanked douc langurs. This assumes bands composed of small OMG with a skewed adult sex ratio. Band size may vary with seasonal food availability resulting in smaller bands when feeding competition is increased. To investigate the social organization of red‐shanked douc langurs and potential seasonal influences, we observed 2 unhabituated groups at Hin Namno National Protected Area, Lao PDR from March 2007 to August 2008 for 803 hr. We recorded births and performed group counts and scan sampling of feeding behavior. Most births (79% of N = 15) occurred from June–September, indicating a 4‐month peak conception season from November to February. Group size averaged 24.5 individuals (range 17–45) with 2.45 adult males (range 1–4). Although the smaller group remained at a stable size (about 18 individuals), the larger group reduced from about 45 to 25 individuals during the 7‐months long lean season, when less than 50% of the feeding time was spent on fruits. This suggests feeding competition as a potential cause of seasonal variation in group size. With 1.9 females per male the skew in adult sex ratio was much lower compared with Rhinopithecus, indicating MMG rather than multilevel societies. However, data on the spacing and interaction patterns between recognized individuals need to be collected and analyzed before the social organization can be determined. Detailed ecological data are furthermore required to investigate the basis for the seasonal changes in group size found. Am. J. Primatol. 73:1134–1144, 2011. © 2011 Wiley Periodicals, Inc.     

Social stability and change among forest hanuman langurs (Presbytis entellus)

The social organization of hanuman langurs (Presbytis entellus, Colobinae) was studied in Kanha Tiger Reserve, Central Indian Highlands between 1980 and 1983, followed by six brief return visits over ten years. Censuses of the 7 km² Kanha meadows demonstrated little change in population density and structure between 1982 and 1990; the population was consistently composed of one-male troops and all-male bands. During the return visits the focal C troop extended its known 74.5 ha range by only 5.6%. By 1993 two identified, habituated, adult females remained within the troop. The adult male was resident in C troop for ≥nine years and eight months.