秦岭金丝猴一个投食猴群等级关系的研究
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摘要
2002年7月至2003年6月，采用随即取样法(Ad lib．Sampling)和目标取样法(Focul animal sampling)对秦岭金丝猴(Rhinopithecus roxellana)西梁投食猴群进行了观察，发现该群是由混合猴群和全雄猴群组成，混合猴群由8个家庭单元、89只个体组成，每个家庭单元几乎是由5个年龄性别组组成，每个家庭平均由一个成年雄性、3.75个成年雌性、1.13个亚成年雌性和3.75青少年猴组成，在生殖季节常有婴猴。婚配制度属于混交的一雄多雌制。通过年龄性别组和性比的分析，说明雄性个体在成长中有逐渐离群的现象，在3龄时离群最多，家庭单元中未见4龄雄性个体，表明川金丝猴混合群是典型雌性为主的家族性社会，成年雄性多是从群外迁入的，从而避免了猴群中近亲繁殖的现象。西梁猴群8个家庭单元的繁殖率明显不同，可能与成年雌性的年龄有关。通过连续两年对混合猴群生育的观察比较，推测川金丝猴在野生条件下多为隔年生育。
采用随即取样法，对秦岭金丝猴西梁投食猴群家庭单元之间的攻击和屈服行为的指向和次数进行记录，采用计算优势指数的方法分析，得出家庭单元之间的等级排列是线性的。研究时间分为交配和产仔两个阶段，在交配季节等级关系的排列为：黑头家庭＞长毛家庭＞井字头家庭＞秃头家庭＞断指家庭＞八字头家庭＞刀疤家庭＞红腿家庭；在产仔季节等级关系为：黑头家庭＞长毛家庭＞秃头家庭＞八字头家庭＞井字头家庭＞断指家庭＞刀疤家庭＞红腿家庭；在全年内等级关系为：黑头家庭＞长毛家庭＞井字头家庭＞秃头家庭＞八字头家庭＞断指家庭＞刀疤家庭＞红腿家庭。由此可见，8个家庭单元在两个阶段及全年内的等级结构排列是不同的。等级较高的家庭单元能够在很长时间内维持自己的地位，而低等级家庭，很难通过争斗提高自己的等级地位。家庭单元常以家庭成员的通力合作提高本家庭的地位，变化多发生在产仔时期。其原因主要是等级较高的家庭成员一般比低等级家庭成员更加团结，低等级的家庭成员之间明显松散，且在产仔季节有些家庭成员团结变得松散，与家庭单元数量及其内的成年雄性强弱无关。家庭单元之间常有冲突，多以劣位家庭单元的回避而结束，从而建立等级关系而达到整个猴群的和平和稳定。猴群中无“猴王”的存在，个体的优势是以单元的优势来体现的，黑头家庭单元虽然在群中占有绝对的优势，但不能充当“猴王”的角色。
对秦岭金丝猴西梁投食猴群家庭单元内部攻击和屈服行为的指向和次数进行
记录，采用计算优势指数的方法，对家庭单元内部的等级进行排列。时间也分为
交配和产仔两个阶段。结果显示，在两个阶段中，8个家庭单元内部等级结构不同。
在交配季节等级关系的排列为:l)黑头>圆脸>小坑>白点>一撮毛>大棒褪;
2)长毛>白线>黑尾>铁锚>浅浅>黑风>双旋>白头>小白;3)八字头>金
箍>背头>香肠>二旋>红头帕>奶嘴>右缺;4)井字头>瘤瘤>宽板猴>窄板
猴>鸡冠>断尾>小黑尾>蛋塔;5)秃头>红冠>半黑线>黄冠>双行>黑线>
黄顶>白顶;6)断指>黑拓>扎扎红>鬼头>红拓>扎扎黑;7)刀疤>白背>
发卡>凹陷>平头>无名>奶酪;8)红腿>双瘤>弯毛>黑背>四孔。在产仔季
节等级关系为:l)黑头>小坑>圆脸>白点>一撮毛>大棒糙;2)长毛>铁锚
>白线>黑尾>浅浅>黑风>双旋>白头>小白;3)八字头>金箍>背头>香肠
>红头帕>二旋>奶嘴>右缺;4)井字头>宽板猴>断尾>鸡冠>窄板猴>瘤瘤
>小黑尾>蛋塔;5)秃头>黄冠>半黑线>红冠>双行>黑线>黄顶>白顶;6)
断指>黑拓>鬼头>扎扎红>红拓>扎扎黑;7)刀疤>发卡>白背>凹陷>平头
>无名>奶酪;8)红腿>双瘤>弯毛>黑背>四孔。在全年平均的等级关系为:
l)黑头>圆脸>小坑>白点>一撮毛>大棒糙;2)长毛>黑尾>白线>铁锚>
浅浅>黑风>双旋>白头>小白;3)八字头>金箍>背头>香肠>红头帕>二旋
>奶嘴>右缺;4)井字头>瘤瘤>宽板猴>窄板猴>鸡冠>断尾>小黑尾>蛋塔;
5)秃头>半黑线>红冠>黄冠>双行>黑线>黄顶>白顶;6)断指>黑拓>鬼
头>红拓>扎扎红>扎扎黑;7)刀疤>白背>发卡>凹陷>平头>无名>奶酪;
8)红腿>双瘤>弯毛>黑背>四孔。结果显示，在每一个家庭内部“家长”居最
高地位，成年雌猴占有第二阶层，亚成体处于第三阶层，青年猴处于第四阶层。
年龄性别组之间等级序列在交配季节和生殖季节没有变化。对交配和产仔阶段每
个家庭内成年雌性等级进行比较，在交配季节成年雌性的等级结构为:圆脸>小
坑;白线>黑尾>铁锚;金箍>背头>香肠;瘤瘤>宽板猴>窄板猴>鸡冠>断
尾;红冠>半黑线>黄冠>双行>黑线;黑拓>扎扎红>鬼头>红拓>扎扎黑;白
背>发卡>凹陷>平头;双瘤>弯毛>黑背。在产仔季节成年雌性的等级结构为:
小坑*>圆脸;铁锚*>白线>黑尾;金箍>背头>香肠;宽板猴*>断尾*>
鸡冠*>窄板猴>瘤瘤;黄冠*>半黑线>红冠>双行>黑线;黑拓*>鬼头>
扎扎红>红拓>扎扎黑;发卡*>白背*>凹陷>平头:双瘤*>弯毛*>黑背
*。由此可见，成年雌性的等级顺位不是固定不变的。对每个家庭成员等级的比
较，发现家庭中没有产仔成员的等级结构几乎不变。家庭中产仔成员的等级结构
发生变化，一般为产仔雌性等级地位高于未产仔雌性。这是成年产仔雌性等级顺
身
位提高的原因之一。
采用目标取样法，即对相互理毛的个体进行锁定，每次观察20mi
Data on social structure and sex ratio of a herd of Sichuan snub-nosed monkey were collected in Zhouzhi National Reserve, which lies in the north slope of the Qinling Mountains from July 2002 to June 2003. By the use of Ad libitum Sampling and focal sampling methods the group in the west Ridge was studied. The results revealed that the whole group was composed of both lots of one-male-units and all-male-unit. One-male-units shows there are 8 one-male-units, each of which on average is made up of one adult male, 3.75 adult female, 1.13 sub-adult female and 3.75 Juvenile, and additional 1.5 infants in breeding season. Marriage should belong to harem above promiscuity. Through analyzing the sex-aged and sex ratio respectively, results showed that male individuals would leave from their one-male-units gradually during the courses of growth, especially in 3 years old and there are no male individuals at the age of about 4 years. This revealed that one-male-units were matrilineal societies, that is to say
, female individuals have been in birth-group all ones life. In order not to inbreed, each male of one-male-units often come out of all-males-unit or other one-male-units. Adult females in wild field incline to reproduce every other year by comparison of the west herd's reproduction in the successive two years. Because of Adult females'age-difference, adult females in each of one-male-units breed differently.
If the group was consisted of same kind of animals, dominance hierarchy was reported among individuals. By the use of Ad libitum Sampling data on the direction and number of aggressive behavior and submissive behavior in the west group of Sichuan snob-nosed monkey was collected from July to December 2002 and April to June 2003. The phase of September to December is called the mating season and that of April to June is named the birth season.and dominance index analyzing data of aggressive behavior and submissive behavior showed that dominance hierarchy among eight units is a linear dominance ranking order. As to mating season the ranking order was: Heitou>Changmao >Jingzitou>Toutu>Duanzhi>Bazitou>Daoba>Hongtui; In birth season, Heitou > Changmao > Toutu > Bazitou > Jingzitou > Duanzhi > Daoba > Hongtui and in the whole year, Heitou > Changmao> Jingzitou>Toutu> Bazitou > Duanzhi> Daoba > Hongtui, which shown that the rank order between the highest
dominant families such as Heitou and Changmao, and the lowest subordinate families such as Daoba and Hongtui had not changed, and changed among Toutu , Bazitou, Jingzitou and Duanzhi mainly in birth season. The unchanging reasons are that the higher dominant families's individuals hold together tighter than those of the lower subordinate families and in birth season the variational cause was that the extent of solidarity among families's individuals changed relaxation and the dominant had nothing to do with families's quantity and their adult male. Eight families in the west group often collide with each other and conflict usually ends by subordinate avoiding. There is no monkey-king in the west group. Although Heitou unit's dominance index is the highest, its unit does not play a king role, and individual's dominance is revealed by unit's dominance.
Subsequently, Ad libitum Sampling was used to record aggressive behavior and submissive behavior in each one-male-unit. Dominance index analysis showed that a linear dominance ranking order exists in each one-male-unit. In mating season a linear dominance ranking order was: Heitou>Yuanlian >Xiaoke>Yizuomao>Dabangchui; Changmao > Baixian > Heiwei > Tiemao > Qianqian > Heifeng > Shuangxuan > Baitou
> Xiaobai; Bazitou > Jingu > Beitou > Xiangchang > Erxuan > Hongtoupa > Naizui > Youque; Jingzitou > Liuliu > Kuanbanhou > Zaibanhou > Jiguan > Duanwei > Xiaoheiwei > Jidan; Tutou > Hongguan > Banheixian > Huangguan > Shuanghang > Heixian > Huangding > Balding; Duanzhi > Heituo > Zhazhahong > Guitou > Hongtuo >Zhazhahei; Daoba>Baibei>Faqia>Aoxian>Pingtou>Wuming>Nailao; Hongtui >Shuangliu>Wanmao>Heibei>Sikong. In the birth season,
采用随即取样法，对秦岭金丝猴西梁投食猴群家庭单元之间的攻击和屈服行为的指向和次数进行记录，采用计算优势指数的方法分析，得出家庭单元之间的等级排列是线性的。研究时间分为交配和产仔两个阶段，在交配季节等级关系的排列为：黑头家庭＞长毛家庭＞井字头家庭＞秃头家庭＞断指家庭＞八字头家庭＞刀疤家庭＞红腿家庭；在产仔季节等级关系为：黑头家庭＞长毛家庭＞秃头家庭＞八字头家庭＞井字头家庭＞断指家庭＞刀疤家庭＞红腿家庭；在全年内等级关系为：黑头家庭＞长毛家庭＞井字头家庭＞秃头家庭＞八字头家庭＞断指家庭＞刀疤家庭＞红腿家庭。由此可见，8个家庭单元在两个阶段及全年内的等级结构排列是不同的。等级较高的家庭单元能够在很长时间内维持自己的地位，而低等级家庭，很难通过争斗提高自己的等级地位。家庭单元常以家庭成员的通力合作提高本家庭的地位，变化多发生在产仔时期。其原因主要是等级较高的家庭成员一般比低等级家庭成员更加团结，低等级的家庭成员之间明显松散，且在产仔季节有些家庭成员团结变得松散，与家庭单元数量及其内的成年雄性强弱无关。家庭单元之间常有冲突，多以劣位家庭单元的回避而结束，从而建立等级关系而达到整个猴群的和平和稳定。猴群中无“猴王”的存在，个体的优势是以单元的优势来体现的，黑头家庭单元虽然在群中占有绝对的优势，但不能充当“猴王”的角色。
对秦岭金丝猴西梁投食猴群家庭单元内部攻击和屈服行为的指向和次数进行
记录，采用计算优势指数的方法，对家庭单元内部的等级进行排列。时间也分为
交配和产仔两个阶段。结果显示，在两个阶段中，8个家庭单元内部等级结构不同。
在交配季节等级关系的排列为:l)黑头>圆脸>小坑>白点>一撮毛>大棒褪;
2)长毛>白线>黑尾>铁锚>浅浅>黑风>双旋>白头>小白;3)八字头>金
箍>背头>香肠>二旋>红头帕>奶嘴>右缺;4)井字头>瘤瘤>宽板猴>窄板
猴>鸡冠>断尾>小黑尾>蛋塔;5)秃头>红冠>半黑线>黄冠>双行>黑线>
黄顶>白顶;6)断指>黑拓>扎扎红>鬼头>红拓>扎扎黑;7)刀疤>白背>
发卡>凹陷>平头>无名>奶酪;8)红腿>双瘤>弯毛>黑背>四孔。在产仔季
节等级关系为:l)黑头>小坑>圆脸>白点>一撮毛>大棒糙;2)长毛>铁锚
>白线>黑尾>浅浅>黑风>双旋>白头>小白;3)八字头>金箍>背头>香肠
>红头帕>二旋>奶嘴>右缺;4)井字头>宽板猴>断尾>鸡冠>窄板猴>瘤瘤
>小黑尾>蛋塔;5)秃头>黄冠>半黑线>红冠>双行>黑线>黄顶>白顶;6)
断指>黑拓>鬼头>扎扎红>红拓>扎扎黑;7)刀疤>发卡>白背>凹陷>平头
>无名>奶酪;8)红腿>双瘤>弯毛>黑背>四孔。在全年平均的等级关系为:
l)黑头>圆脸>小坑>白点>一撮毛>大棒糙;2)长毛>黑尾>白线>铁锚>
浅浅>黑风>双旋>白头>小白;3)八字头>金箍>背头>香肠>红头帕>二旋
>奶嘴>右缺;4)井字头>瘤瘤>宽板猴>窄板猴>鸡冠>断尾>小黑尾>蛋塔;
5)秃头>半黑线>红冠>黄冠>双行>黑线>黄顶>白顶;6)断指>黑拓>鬼
头>红拓>扎扎红>扎扎黑;7)刀疤>白背>发卡>凹陷>平头>无名>奶酪;
8)红腿>双瘤>弯毛>黑背>四孔。结果显示，在每一个家庭内部“家长”居最
高地位，成年雌猴占有第二阶层，亚成体处于第三阶层，青年猴处于第四阶层。
年龄性别组之间等级序列在交配季节和生殖季节没有变化。对交配和产仔阶段每
个家庭内成年雌性等级进行比较，在交配季节成年雌性的等级结构为:圆脸>小
坑;白线>黑尾>铁锚;金箍>背头>香肠;瘤瘤>宽板猴>窄板猴>鸡冠>断
尾;红冠>半黑线>黄冠>双行>黑线;黑拓>扎扎红>鬼头>红拓>扎扎黑;白
背>发卡>凹陷>平头;双瘤>弯毛>黑背。在产仔季节成年雌性的等级结构为:
小坑*>圆脸;铁锚*>白线>黑尾;金箍>背头>香肠;宽板猴*>断尾*>
鸡冠*>窄板猴>瘤瘤;黄冠*>半黑线>红冠>双行>黑线;黑拓*>鬼头>
扎扎红>红拓>扎扎黑;发卡*>白背*>凹陷>平头:双瘤*>弯毛*>黑背
*。由此可见，成年雌性的等级顺位不是固定不变的。对每个家庭成员等级的比
较，发现家庭中没有产仔成员的等级结构几乎不变。家庭中产仔成员的等级结构
发生变化，一般为产仔雌性等级地位高于未产仔雌性。这是成年产仔雌性等级顺
身
位提高的原因之一。
采用目标取样法，即对相互理毛的个体进行锁定，每次观察20mi
Data on social structure and sex ratio of a herd of Sichuan snub-nosed monkey were collected in Zhouzhi National Reserve, which lies in the north slope of the Qinling Mountains from July 2002 to June 2003. By the use of Ad libitum Sampling and focal sampling methods the group in the west Ridge was studied. The results revealed that the whole group was composed of both lots of one-male-units and all-male-unit. One-male-units shows there are 8 one-male-units, each of which on average is made up of one adult male, 3.75 adult female, 1.13 sub-adult female and 3.75 Juvenile, and additional 1.5 infants in breeding season. Marriage should belong to harem above promiscuity. Through analyzing the sex-aged and sex ratio respectively, results showed that male individuals would leave from their one-male-units gradually during the courses of growth, especially in 3 years old and there are no male individuals at the age of about 4 years. This revealed that one-male-units were matrilineal societies, that is to say
, female individuals have been in birth-group all ones life. In order not to inbreed, each male of one-male-units often come out of all-males-unit or other one-male-units. Adult females in wild field incline to reproduce every other year by comparison of the west herd's reproduction in the successive two years. Because of Adult females'age-difference, adult females in each of one-male-units breed differently.
If the group was consisted of same kind of animals, dominance hierarchy was reported among individuals. By the use of Ad libitum Sampling data on the direction and number of aggressive behavior and submissive behavior in the west group of Sichuan snob-nosed monkey was collected from July to December 2002 and April to June 2003. The phase of September to December is called the mating season and that of April to June is named the birth season.and dominance index analyzing data of aggressive behavior and submissive behavior showed that dominance hierarchy among eight units is a linear dominance ranking order. As to mating season the ranking order was: Heitou>Changmao >Jingzitou>Toutu>Duanzhi>Bazitou>Daoba>Hongtui; In birth season, Heitou > Changmao > Toutu > Bazitou > Jingzitou > Duanzhi > Daoba > Hongtui and in the whole year, Heitou > Changmao> Jingzitou>Toutu> Bazitou > Duanzhi> Daoba > Hongtui, which shown that the rank order between the highest
dominant families such as Heitou and Changmao, and the lowest subordinate families such as Daoba and Hongtui had not changed, and changed among Toutu , Bazitou, Jingzitou and Duanzhi mainly in birth season. The unchanging reasons are that the higher dominant families's individuals hold together tighter than those of the lower subordinate families and in birth season the variational cause was that the extent of solidarity among families's individuals changed relaxation and the dominant had nothing to do with families's quantity and their adult male. Eight families in the west group often collide with each other and conflict usually ends by subordinate avoiding. There is no monkey-king in the west group. Although Heitou unit's dominance index is the highest, its unit does not play a king role, and individual's dominance is revealed by unit's dominance.
Subsequently, Ad libitum Sampling was used to record aggressive behavior and submissive behavior in each one-male-unit. Dominance index analysis showed that a linear dominance ranking order exists in each one-male-unit. In mating season a linear dominance ranking order was: Heitou>Yuanlian >Xiaoke>Yizuomao>Dabangchui; Changmao > Baixian > Heiwei > Tiemao > Qianqian > Heifeng > Shuangxuan > Baitou
> Xiaobai; Bazitou > Jingu > Beitou > Xiangchang > Erxuan > Hongtoupa > Naizui > Youque; Jingzitou > Liuliu > Kuanbanhou > Zaibanhou > Jiguan > Duanwei > Xiaoheiwei > Jidan; Tutou > Hongguan > Banheixian > Huangguan > Shuanghang > Heixian > Huangding > Balding; Duanzhi > Heituo > Zhazhahong > Guitou > Hongtuo >Zhazhahei; Daoba>Baibei>Faqia>Aoxian>Pingtou>Wuming>Nailao; Hongtui >Shuangliu>Wanmao>Heibei>Sikong. In the birth season,
引文
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