短尾猴理毛行为交换策略与群体稳定性研究
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摘要
大多数灵长类动物因外部生态环境因素的作用而形成群体。群体生活有利于个体成员发现食物和抵御天敌,但同时也带来空间、食物、配偶等方面的竞争。群居的动物通过错综复杂的行为进行交往和联系,有些行为(如友好行为和性行为)有利于维持群体关系,有些行为(如攻击行为)则不利于维持群体生活,因此,群居灵长类动物应该存在某种行为机制来平衡和适应某些行为产生的不利影响,否则群体终将面临分裂和瓦解,群体生活的优势也将消失。
关于动物群体形成与维持机制已有许多研究。上世纪七十年代,有学者提出社会行为之间存在交换,且这种交换利于缓解个体间竞争。九十年代中期,Noe和Hammerstein提出了生物市场理论(Biological Market Theory)。该理论认为群体可以看做一个自由市场,社会行为可视为具有特定价值的商品,动物个体可以自由选择伙伴交换行为,并且这种交换随着供求关系的变化而变化。自该理论提出以来,对该学说的验证成为动物研究的热点之一,涉及多个灵长类物种。这些研究验证了行为交换市场的存在,理毛行为是通用交换的“货币”行为,但对行为交换的方式、交换对象选择、交换与群体稳定性的关系等缺乏系统深入和可比较性的研究,难以解释灵长类动物维持群体稳定的行为机制。
本研究以课题组20多年长期跟踪研究的、栖息在安徽黄山的、国家二级保护野生动物短尾猴(Macaca thibetana)为对象,通过对理毛行为和其它社会行为(如攻击、交配等)的研究,探讨野生短尾猴理毛行为策略及其与群体稳定之间的关系,期望回答以下重要科学问题：(1)短尾猴社会群体中存在何种类型行为之间的市场交换?(2)雌性和雄性分别采取何种理毛伙伴选择策略?(3)雌性和雄性对群体稳定的作用如何?本研究不仅有助于阐明理毛行为对维持群体稳定的作用,更有助于揭示行为交换的经济学规律,进而揭示维持群体稳定性和凝聚力的内在行为机制。
本研究采用目标动物取样法(Focal sampling methods)和连续记录方法(Continuously recording methods),分别于2009年5月至2010年8月和2011年9月至2012年5月,实时跟踪猴群并获得野外观测数据,有效观察天数为311天,有效时间为1119.33小时,主要结果如下：
一、发现了短尾猴行为交换的多样性。通过对雌性-雌性之间、雄性-雄性之间、雌性-雄性之间的理毛投资与收益以及社会行为的关系对比分析,发现短尾猴群体内个体之间行为交换类型多样,主要表现为：(1)同种行为之间可以交换(如“理毛”换取“理毛”)。相同或相似等级的同性个体之间,动物个体投资理毛可以交换相同频次或持续时间的理毛行为。这种交换随着投资者理毛行为频次或持续时间的增加而增加,即投资理毛频次越高、持续时间越长,获得的理毛回报也就越高、持续时间也越长；等级距离较大的同性个体之间,理毛行为交换不等价,即低顺位个体为高顺位个体投资理毛不能获取该高顺位个体给予同等价值的理毛回报,表现为低顺位对高顺位的理毛投资显著高于高顺位对低顺位的理毛投资。(2)不同行为之间可以交换(如“理毛”换取“容忍”)。高顺位个体对低顺位个体的容忍度可以作为高顺位个体与低顺位个体之间理毛行为不等价交换的有效补偿,即低顺位为高顺位投资更多的理毛可以换取同等价值的行为,如高顺位对自身的容忍度,降低被攻击的风险。(3)不同类型行为也可以交换(如“理毛”换取“交配”)。虽然异性个体之间理毛投资与收益均呈显著正相关,但异性个体之间的理毛投资与收益之间显著不平等,即理毛行为交换不等价,表现为雌性对雄性投资的理毛行为显著高于雄性对雌性的理毛。进一步的分析发现,交配机会可以作为异性个体之间理毛行为不等价交换的行为补偿,即雌性个体为雄性个体投资更多的理毛可以有效地换取同等价值的不同类型行为,如与相应雄性个体的交配机会。
二、阐明了短尾猴的理毛伙伴选择策略。在短尾猴社会群体中,雄性个体在性成熟后迁出出生群,而雌性个体则永久生活在出生群。因此,个体社会等级变化、雄性迁入、雄性迁出以及分群是野生短尾猴社会群体动态变化的主要因素。依据这些特征,将本研究时期分为五个阶段,采用雌性／雄性优先选择指数(Female/male-preferred index)和伙伴稳定性指数(Partner stability index)分析短尾猴理毛伙伴选择策略及其动态适应性。结果表明,雄性个体和雌性个体均优先选择雌性个体作为理毛投资对象,即雄性-雌性之间、雌性-雌性之间理毛行为显著高于雄性-雄性之间和雌性-雄性之间；不仅如此,通过对不同研究阶段、不同性别组之间理毛伙伴稳定性的比较分析,发现雌性-雌性之间的理毛伙伴稳定性显著高于雄性-雄性之间和雌性-雄性之间,雌性个体之间的理毛伙伴稳定性最强,雌性-雄性之间次之,雄性个体之间理毛伙伴稳定性最差。由此看出,优先选择雌性伙伴并与之保持稳定的伙伴关系与短尾猴的群体结构特征相一致。
三、揭示了性别差异对群体稳定性的不同作用。采用多层次分析社会关系的新方法---社会网络结构分析(Social network analysis),对比分析野生短尾猴理毛行为网络结构中个体间行为交流强度(Strength)、特征向量中心度(Eigenvector centrality)、聚类系数(Clustering coefficient)和个体间亲密度(Affinity)等代表动物个体之间社会关系的特征值,探讨雌性和雄性对群体稳定性的作用。虽然雌性和雄性个体共同维持群体稳定,但是,雌性短尾猴的特征向量中心度水平显著高于雄性个体,即雌性短尾猴对群体稳定的作用显著高于雄性个体；同型相关性聚类分析(Hierarchical cluster analysis)结果进一步表明,短尾猴社会群体内部,存在明显的小聚群现象,聚群的主要方式是亲缘个体之间的频繁交流,因此,亲缘个体间频繁的理毛交往是母系社会群体稳定的行为基础。另一方面,雄性参与群体生活虽然降低了雌性个体的聚类系数,即削弱了雌性个体之间的“派系”或“小聚群”,但显著提高“派系”内部成员个体之间和“派系”之间行为交流的频次,提升了雌性-雌性之间、雄性-雄性之间以及群体水平的凝聚力,从而定量证明了雄性成员对母系社会群体稳定性的作用。
本研究运用了生物学、生态学、行为学、经济学以及社会学的知识,通过实时跟踪猴群、录音实时记录动物行为,探讨了野生短尾猴理毛行为与其他社会行为之间的交换关系,揭示了理毛行为对维持群体稳定的重要意义和行为策略。本课题的主要成果创新在于：(1)发现了动物行为交换的多样性：不仅相同行为可以交换(如“理毛”换取“理毛”),而且不同行为可以交换(如“理毛”换取“容忍”),甚至不同性质的行为也可以交换(如“理毛”换取“交配”),这就从社会关系的角度揭示了动物群体稳定的行为机制；(2)定量证明了雄性成员对群体的重要作用：理毛网络结构分析表明,虽然雌性短尾猴的特征向量中心度水平显著高于雄性个体,即雌性短尾猴对群体稳定的作用显著高于雄性个体,但雄性参与群体生活显著提高了群体成员个体之间行为交流的频次,从而促进了雄性和雌性凝聚力的提升。
维持群体的稳定性和凝聚力,不仅是动物社会面临的问题,也是人类社会面临的挑战。人类和谐社会建设需要照顾各方关系、平衡各方利益。“动物是人类的自然老师”,本研究成果不仅揭示了短尾猴维持社群稳定和增强凝聚力的内在行为机制,促进了社会行为进化中的经济学研究,对短尾猴的保护和管理具有重要意义,同时,对理解人类社会的复杂关系、构建和谐社会也具有较高的科学参考价值。
Recent studies of group living in many species of non-human primates indicate that, similar with human society, the coordinated or cooperative action of several individuals can provide multiple benefits to group members, including earlier predator detection, higher foraging efficiency and improved resource defense. Individuals form social relationships by complex behavioral interactions. Some behaviors (e.g., friendly and sexual) improve to maintain social stability, however, some behaviors (e.g., aggressive) destroy social cohesiveness. Thus, social animals have adaptation strategies to balance these two aspects, otherwise the group will be broken.
There are abundant of researches about group living and maintaining group cohesiveness. In1970s, some authors hypothesized that exchange among behaviors can buffer competition. In1990s, biological market theory was proposed. Within this framework, grooming can be perceived as a commodity of hygienic value, a commodity of social value, or both, and exchanged for some equivalent commodity in the marketplace that is the social group. To date, there are lots of studies focused on testing the biological market theory, especially in nonhuman primates. Previous studies found some evidence for behavioral exchange/interchange and grooming would be currency for exchange/interchange. However, the type of exchange, partner choice, the relationship between exchange/interchange and group stability are less known. Therefore, the behavioral mechanisam of maintaining group stability can not be demonstrated within the systematic study.
The present study, taking Tibetan macaques (Macaca thibetana) as subject, focused on the grooming and other social behaviors (e.g., aggression, copulation), and discussed on the partner choice and its adaptation strategy. The following questions can be answered:(1) what kinds of exchanges in Tibetan macaques?(2) how to select grooming partner for both males and females?(3) what's the function of male/female for group stability? This study can provide insight into the function of grooming on group stability, promote the study for laws of behavioral economics,.and reveal the -behavioral mechanisim of group stability and cohesiveness.
In the present study, using focal sampling methods and continuously recording methods, I obtained behavioral data (311days,1119.33hours) during the study period (May.2009-Aug.2010and Sep.2011). The present study analyzed the dyadic grooming interaction and its adaptation strategy based on analysis of grooming investment among intra-and inter-sexual dyads partner choice, adaptation characteristic and grooming network analysis. The following important results were found as follow:
1The present study found the diversity of behavioral exchange/interchange. This study focus on the grooming investment and the relationships between grooming and other social behaviors (i.e., aggression, copulation) among female-female, male-male, and female-male dyads. The results showed that there was variety of behavioral exchange/interchange among Tibetan macaques.(1) Exchange between the same behaviors, such as grooming exchange for itself. Among intra-sexual dyads (female-female and male-male dyads) with the similar dominance hierarchy, grooming investment could exchange for itself with equal values, such as the same frequency or duration. That is to say, grooming reciprocity also occurred principally between individuals of adjacent rank. Correlation tests implied that individuals increased grooming investment with increasing grooming received, and vice versa. Among intra-sexual dyads, when individuals of different rank groomed, individuals tended to groom up the hierarchy, lower-ranking individuals groomed higher-ranking individuals more than vice versa was to increase the tolerance from higher-ranking individuals by aggression reduced.(2) Interchange between different behavior but the same type, such as grooming for tolerance. In intra-sexual dyads with different ranks, tolerance could be as compensation for the disparity of grooming initiated and received.(3) Interchange between different property behaviors, for example, grooming for copulations. In inter-sexual dyads, compared to grooming strategies in intra-sexual dyads, animal individuals pay more attention to the relationship between grooming investment and copulation opportunity. Grooming given was significantly positive correlated with grooming received, however, female individuals groom males at a greater rate and/or for a longer duration during both the mating season and non-mating seasons. High-ranking, middle-ranking, and low-ranking females preferred to groom and/or copulate with high-ranking, middle-ranking, and low-ranking males, respectively. Female-to-male grooming was correlative with copulation indicated that mating opportunity could be as compensation to offset the grooming disparity among female-male dyads.
2This study found the partner choice strategy based on grooming interaction in Tibetan macaques. Female philopatry and male dispersal is the important characteristics of Tibetan macaques. Variation of dominance hierarchy, immigrating, emigrating, and fission is the factors of group dynamic. Animals have their own adaptation strategies. The current study analyzed the female-preferred index and partner stability index during the five subperiods with group dynamic. Grooming investment was at a greater and/or longer duration in female-female dyads than male-male and female-male dyads. Partner stability in female-female dyads was higher than both male-male and female-male dyads. These results indicate that Tibetan macaques preferred to groom females and formed stability female partner, which is consistent with group structure of Tibetan macaques.
3This study provided the male/female role for group stability. Using social network analysis, the present study analyzed the individual's role for group stability using strength, eigenvector centrality, clustering coefficient and affinity. Both males and females contributed for the group stability and cohesiveness, however, the eigenvector centrality of females were higher than males', which represented that female Tibetan macaques contributed more than males. Further, for grooming frequency, hierarchical cluster analysis identified five cliquishnesses clusteed according to matrilineal kin with a cophenetic correlation coefficient of0.816, it suggested matrilineal kinship play an important role in the group of Tibetan macaques. However, for grooming duration, there was no cliquishness, it suggested that in Tibetan macaques, frequently grooming interactions play more important roles for maintaining group cohesiveness than long duration but nonfrequently grooming. On other hand, although females' clustering coefficient were decreased when male participated, male can be as a adhesive to bridge the behavioral interactions between males and females. Male participation can improve highly the interactions within/among cliquishness, and the cohesiveness among females, males. Therefore, this study give the new evidence of the function for group stability, male participation can promote the group stability and cohesiveness.
This study used the combinded idea within the biology, ecology, behavior, economics and sociology. Focal animal sampling and continuous recording (using a digital voice recorder, model News my RV50) were used to score the activity of the focal monkey when following. The aim of the present study was to focus on the relationship between grooming and other social behaviors, to discuss the strategy for group stability based on dyadic grooming interactions. The innovational results were found as followed:(1) This study found the diversity of behavioral exchange/interchange. Behaviors can be exchanged for itself (i.e., grooming for grooming), for other behavior with the same property (i.e., grooming for tolerance), and can also for behavior with different property (i.e., grooming for copulation). This result provides insight into the behavioral mechanism for group stability.(2) Quantitative proved the male's function for group-living animals. Male participation can improve highly the interactions within/among cliquishness, and the cohesiveness among females, males. Therefore, this study give the new evidence of male's function for group stability, male participation can promote the group stability and cohesiveness.
Maintaining group stability and cohesiveness is the challenge for animals, and it is also a mutual point for zoologists, ecologists, anthropologists and sociologists. Construction of harmonious society need balance the benefit among the whole society. We can get some significant hints from the nature, animals. In view of this point, the present study significantly explained the behavioral mechanism to maintain social bonds and cohesiveness in non-human primates, and also to promote the economic research for evolution of social behaviors. It may open up a new area to concentrate behavior interactions and strategies on both individual and group levels. Meanwhile, this study will help forward conservation and management of Tibetan macaques, and shed light on building harmonious society in human society.
关于动物群体形成与维持机制已有许多研究。上世纪七十年代,有学者提出社会行为之间存在交换,且这种交换利于缓解个体间竞争。九十年代中期,Noe和Hammerstein提出了生物市场理论(Biological Market Theory)。该理论认为群体可以看做一个自由市场,社会行为可视为具有特定价值的商品,动物个体可以自由选择伙伴交换行为,并且这种交换随着供求关系的变化而变化。自该理论提出以来,对该学说的验证成为动物研究的热点之一,涉及多个灵长类物种。这些研究验证了行为交换市场的存在,理毛行为是通用交换的“货币”行为,但对行为交换的方式、交换对象选择、交换与群体稳定性的关系等缺乏系统深入和可比较性的研究,难以解释灵长类动物维持群体稳定的行为机制。
本研究以课题组20多年长期跟踪研究的、栖息在安徽黄山的、国家二级保护野生动物短尾猴(Macaca thibetana)为对象,通过对理毛行为和其它社会行为(如攻击、交配等)的研究,探讨野生短尾猴理毛行为策略及其与群体稳定之间的关系,期望回答以下重要科学问题：(1)短尾猴社会群体中存在何种类型行为之间的市场交换?(2)雌性和雄性分别采取何种理毛伙伴选择策略?(3)雌性和雄性对群体稳定的作用如何?本研究不仅有助于阐明理毛行为对维持群体稳定的作用,更有助于揭示行为交换的经济学规律,进而揭示维持群体稳定性和凝聚力的内在行为机制。
本研究采用目标动物取样法(Focal sampling methods)和连续记录方法(Continuously recording methods),分别于2009年5月至2010年8月和2011年9月至2012年5月,实时跟踪猴群并获得野外观测数据,有效观察天数为311天,有效时间为1119.33小时,主要结果如下：
一、发现了短尾猴行为交换的多样性。通过对雌性-雌性之间、雄性-雄性之间、雌性-雄性之间的理毛投资与收益以及社会行为的关系对比分析,发现短尾猴群体内个体之间行为交换类型多样,主要表现为：(1)同种行为之间可以交换(如“理毛”换取“理毛”)。相同或相似等级的同性个体之间,动物个体投资理毛可以交换相同频次或持续时间的理毛行为。这种交换随着投资者理毛行为频次或持续时间的增加而增加,即投资理毛频次越高、持续时间越长,获得的理毛回报也就越高、持续时间也越长；等级距离较大的同性个体之间,理毛行为交换不等价,即低顺位个体为高顺位个体投资理毛不能获取该高顺位个体给予同等价值的理毛回报,表现为低顺位对高顺位的理毛投资显著高于高顺位对低顺位的理毛投资。(2)不同行为之间可以交换(如“理毛”换取“容忍”)。高顺位个体对低顺位个体的容忍度可以作为高顺位个体与低顺位个体之间理毛行为不等价交换的有效补偿,即低顺位为高顺位投资更多的理毛可以换取同等价值的行为,如高顺位对自身的容忍度,降低被攻击的风险。(3)不同类型行为也可以交换(如“理毛”换取“交配”)。虽然异性个体之间理毛投资与收益均呈显著正相关,但异性个体之间的理毛投资与收益之间显著不平等,即理毛行为交换不等价,表现为雌性对雄性投资的理毛行为显著高于雄性对雌性的理毛。进一步的分析发现,交配机会可以作为异性个体之间理毛行为不等价交换的行为补偿,即雌性个体为雄性个体投资更多的理毛可以有效地换取同等价值的不同类型行为,如与相应雄性个体的交配机会。
二、阐明了短尾猴的理毛伙伴选择策略。在短尾猴社会群体中,雄性个体在性成熟后迁出出生群,而雌性个体则永久生活在出生群。因此,个体社会等级变化、雄性迁入、雄性迁出以及分群是野生短尾猴社会群体动态变化的主要因素。依据这些特征,将本研究时期分为五个阶段,采用雌性／雄性优先选择指数(Female/male-preferred index)和伙伴稳定性指数(Partner stability index)分析短尾猴理毛伙伴选择策略及其动态适应性。结果表明,雄性个体和雌性个体均优先选择雌性个体作为理毛投资对象,即雄性-雌性之间、雌性-雌性之间理毛行为显著高于雄性-雄性之间和雌性-雄性之间；不仅如此,通过对不同研究阶段、不同性别组之间理毛伙伴稳定性的比较分析,发现雌性-雌性之间的理毛伙伴稳定性显著高于雄性-雄性之间和雌性-雄性之间,雌性个体之间的理毛伙伴稳定性最强,雌性-雄性之间次之,雄性个体之间理毛伙伴稳定性最差。由此看出,优先选择雌性伙伴并与之保持稳定的伙伴关系与短尾猴的群体结构特征相一致。
三、揭示了性别差异对群体稳定性的不同作用。采用多层次分析社会关系的新方法---社会网络结构分析(Social network analysis),对比分析野生短尾猴理毛行为网络结构中个体间行为交流强度(Strength)、特征向量中心度(Eigenvector centrality)、聚类系数(Clustering coefficient)和个体间亲密度(Affinity)等代表动物个体之间社会关系的特征值,探讨雌性和雄性对群体稳定性的作用。虽然雌性和雄性个体共同维持群体稳定,但是,雌性短尾猴的特征向量中心度水平显著高于雄性个体,即雌性短尾猴对群体稳定的作用显著高于雄性个体；同型相关性聚类分析(Hierarchical cluster analysis)结果进一步表明,短尾猴社会群体内部,存在明显的小聚群现象,聚群的主要方式是亲缘个体之间的频繁交流,因此,亲缘个体间频繁的理毛交往是母系社会群体稳定的行为基础。另一方面,雄性参与群体生活虽然降低了雌性个体的聚类系数,即削弱了雌性个体之间的“派系”或“小聚群”,但显著提高“派系”内部成员个体之间和“派系”之间行为交流的频次,提升了雌性-雌性之间、雄性-雄性之间以及群体水平的凝聚力,从而定量证明了雄性成员对母系社会群体稳定性的作用。
本研究运用了生物学、生态学、行为学、经济学以及社会学的知识,通过实时跟踪猴群、录音实时记录动物行为,探讨了野生短尾猴理毛行为与其他社会行为之间的交换关系,揭示了理毛行为对维持群体稳定的重要意义和行为策略。本课题的主要成果创新在于：(1)发现了动物行为交换的多样性：不仅相同行为可以交换(如“理毛”换取“理毛”),而且不同行为可以交换(如“理毛”换取“容忍”),甚至不同性质的行为也可以交换(如“理毛”换取“交配”),这就从社会关系的角度揭示了动物群体稳定的行为机制；(2)定量证明了雄性成员对群体的重要作用：理毛网络结构分析表明,虽然雌性短尾猴的特征向量中心度水平显著高于雄性个体,即雌性短尾猴对群体稳定的作用显著高于雄性个体,但雄性参与群体生活显著提高了群体成员个体之间行为交流的频次,从而促进了雄性和雌性凝聚力的提升。
维持群体的稳定性和凝聚力,不仅是动物社会面临的问题,也是人类社会面临的挑战。人类和谐社会建设需要照顾各方关系、平衡各方利益。“动物是人类的自然老师”,本研究成果不仅揭示了短尾猴维持社群稳定和增强凝聚力的内在行为机制,促进了社会行为进化中的经济学研究,对短尾猴的保护和管理具有重要意义,同时,对理解人类社会的复杂关系、构建和谐社会也具有较高的科学参考价值。
Recent studies of group living in many species of non-human primates indicate that, similar with human society, the coordinated or cooperative action of several individuals can provide multiple benefits to group members, including earlier predator detection, higher foraging efficiency and improved resource defense. Individuals form social relationships by complex behavioral interactions. Some behaviors (e.g., friendly and sexual) improve to maintain social stability, however, some behaviors (e.g., aggressive) destroy social cohesiveness. Thus, social animals have adaptation strategies to balance these two aspects, otherwise the group will be broken.
There are abundant of researches about group living and maintaining group cohesiveness. In1970s, some authors hypothesized that exchange among behaviors can buffer competition. In1990s, biological market theory was proposed. Within this framework, grooming can be perceived as a commodity of hygienic value, a commodity of social value, or both, and exchanged for some equivalent commodity in the marketplace that is the social group. To date, there are lots of studies focused on testing the biological market theory, especially in nonhuman primates. Previous studies found some evidence for behavioral exchange/interchange and grooming would be currency for exchange/interchange. However, the type of exchange, partner choice, the relationship between exchange/interchange and group stability are less known. Therefore, the behavioral mechanisam of maintaining group stability can not be demonstrated within the systematic study.
The present study, taking Tibetan macaques (Macaca thibetana) as subject, focused on the grooming and other social behaviors (e.g., aggression, copulation), and discussed on the partner choice and its adaptation strategy. The following questions can be answered:(1) what kinds of exchanges in Tibetan macaques?(2) how to select grooming partner for both males and females?(3) what's the function of male/female for group stability? This study can provide insight into the function of grooming on group stability, promote the study for laws of behavioral economics,.and reveal the -behavioral mechanisim of group stability and cohesiveness.
In the present study, using focal sampling methods and continuously recording methods, I obtained behavioral data (311days,1119.33hours) during the study period (May.2009-Aug.2010and Sep.2011). The present study analyzed the dyadic grooming interaction and its adaptation strategy based on analysis of grooming investment among intra-and inter-sexual dyads partner choice, adaptation characteristic and grooming network analysis. The following important results were found as follow:
1The present study found the diversity of behavioral exchange/interchange. This study focus on the grooming investment and the relationships between grooming and other social behaviors (i.e., aggression, copulation) among female-female, male-male, and female-male dyads. The results showed that there was variety of behavioral exchange/interchange among Tibetan macaques.(1) Exchange between the same behaviors, such as grooming exchange for itself. Among intra-sexual dyads (female-female and male-male dyads) with the similar dominance hierarchy, grooming investment could exchange for itself with equal values, such as the same frequency or duration. That is to say, grooming reciprocity also occurred principally between individuals of adjacent rank. Correlation tests implied that individuals increased grooming investment with increasing grooming received, and vice versa. Among intra-sexual dyads, when individuals of different rank groomed, individuals tended to groom up the hierarchy, lower-ranking individuals groomed higher-ranking individuals more than vice versa was to increase the tolerance from higher-ranking individuals by aggression reduced.(2) Interchange between different behavior but the same type, such as grooming for tolerance. In intra-sexual dyads with different ranks, tolerance could be as compensation for the disparity of grooming initiated and received.(3) Interchange between different property behaviors, for example, grooming for copulations. In inter-sexual dyads, compared to grooming strategies in intra-sexual dyads, animal individuals pay more attention to the relationship between grooming investment and copulation opportunity. Grooming given was significantly positive correlated with grooming received, however, female individuals groom males at a greater rate and/or for a longer duration during both the mating season and non-mating seasons. High-ranking, middle-ranking, and low-ranking females preferred to groom and/or copulate with high-ranking, middle-ranking, and low-ranking males, respectively. Female-to-male grooming was correlative with copulation indicated that mating opportunity could be as compensation to offset the grooming disparity among female-male dyads.
2This study found the partner choice strategy based on grooming interaction in Tibetan macaques. Female philopatry and male dispersal is the important characteristics of Tibetan macaques. Variation of dominance hierarchy, immigrating, emigrating, and fission is the factors of group dynamic. Animals have their own adaptation strategies. The current study analyzed the female-preferred index and partner stability index during the five subperiods with group dynamic. Grooming investment was at a greater and/or longer duration in female-female dyads than male-male and female-male dyads. Partner stability in female-female dyads was higher than both male-male and female-male dyads. These results indicate that Tibetan macaques preferred to groom females and formed stability female partner, which is consistent with group structure of Tibetan macaques.
3This study provided the male/female role for group stability. Using social network analysis, the present study analyzed the individual's role for group stability using strength, eigenvector centrality, clustering coefficient and affinity. Both males and females contributed for the group stability and cohesiveness, however, the eigenvector centrality of females were higher than males', which represented that female Tibetan macaques contributed more than males. Further, for grooming frequency, hierarchical cluster analysis identified five cliquishnesses clusteed according to matrilineal kin with a cophenetic correlation coefficient of0.816, it suggested matrilineal kinship play an important role in the group of Tibetan macaques. However, for grooming duration, there was no cliquishness, it suggested that in Tibetan macaques, frequently grooming interactions play more important roles for maintaining group cohesiveness than long duration but nonfrequently grooming. On other hand, although females' clustering coefficient were decreased when male participated, male can be as a adhesive to bridge the behavioral interactions between males and females. Male participation can improve highly the interactions within/among cliquishness, and the cohesiveness among females, males. Therefore, this study give the new evidence of the function for group stability, male participation can promote the group stability and cohesiveness.
This study used the combinded idea within the biology, ecology, behavior, economics and sociology. Focal animal sampling and continuous recording (using a digital voice recorder, model News my RV50) were used to score the activity of the focal monkey when following. The aim of the present study was to focus on the relationship between grooming and other social behaviors, to discuss the strategy for group stability based on dyadic grooming interactions. The innovational results were found as followed:(1) This study found the diversity of behavioral exchange/interchange. Behaviors can be exchanged for itself (i.e., grooming for grooming), for other behavior with the same property (i.e., grooming for tolerance), and can also for behavior with different property (i.e., grooming for copulation). This result provides insight into the behavioral mechanism for group stability.(2) Quantitative proved the male's function for group-living animals. Male participation can improve highly the interactions within/among cliquishness, and the cohesiveness among females, males. Therefore, this study give the new evidence of male's function for group stability, male participation can promote the group stability and cohesiveness.
Maintaining group stability and cohesiveness is the challenge for animals, and it is also a mutual point for zoologists, ecologists, anthropologists and sociologists. Construction of harmonious society need balance the benefit among the whole society. We can get some significant hints from the nature, animals. In view of this point, the present study significantly explained the behavioral mechanism to maintain social bonds and cohesiveness in non-human primates, and also to promote the economic research for evolution of social behaviors. It may open up a new area to concentrate behavior interactions and strategies on both individual and group levels. Meanwhile, this study will help forward conservation and management of Tibetan macaques, and shed light on building harmonious society in human society.
引文
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