秦岭川金丝猴（Rhinopithecus roxellana）繁殖竞争与肢体偏好的研究
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摘要
第一部分秦岭川金丝猴繁殖竞争的研究
第一章秦岭川金丝猴主雄替换后单元内雌雄两性繁殖策略的研究
2003年3月至2004年5月,利用焦点动物取样法和行为取样法对分布在秦岭北坡周至国家自然保护区玉皇庙地区川金丝猴西梁群关于主雄替换对单元内两性繁殖关系的影响进行研究。研究结果发现:主雄替换后,单元新主雄会倾向于自主选择非处于哺乳期成年雌性作为主要繁殖对象。新主雄通过单元外交配行为,不仅可以增加扩散自身基因的可能性,而且可以刺激单元内交配行为的发生进而增加自身繁殖成功机会。哺乳期雌性个体在其婴猴消失后的首个生育间隔显著短于研究群内未失婴猴雌性的平均生育间隔,这意味着野生成年雌性川金丝猴面对自身婴猴消失会采取积极的适应性繁殖策略。此外,亚成年雌性川金丝猴通过积极与新主雄建立良好繁殖关系进而获得其在成年后的繁殖成功。总之,野生川金丝猴单元新主雄、成年雌性和亚成年雌性个体面对主雄替换及之后的婴猴消失各自采取了不同的繁殖策略,这在一定程度上体现了该物种对于主雄替换这一避免近亲繁殖机制的行为适应性。
第二章秦岭川金丝猴雌性扩散背后的交配竞争及相关繁殖行为研究
2001年10月至2005年5月,对分布在秦岭北坡周至国家自然保护区玉皇庙地区川金丝猴西梁群关于雌性个体扩散与繁殖成功关系进行研究。研究结果发现:成年雌性个体和业成年雌性个体均发生扩散,成年雌性个体的迁出率和迁入率均高于亚成年雌性个体。交配竞争是成年雌性个体扩散的主要推动力,而亚成年雌性个体扩散可能是为了避免近亲繁殖。成年雌性个体在扩散过程中通过采用多样化的繁殖策略进而提高自身繁殖成功率。
第三章秦岭川金丝猴单元大小及组成对雌性繁殖成功影响的研究
动物的社会性在其群居利益大于群居代价时得以进化。群组大小和群组构成是评价动物社会性的两个基础衡量尺度,它们对雌性繁殖成功发挥重要影响作用。2001年10月至2008年5月,对分布在秦岭北坡周至国家自然保护区玉皇庙地区川金丝猴西梁群关于一雄多雌单元大小和组成对川金丝猴雌性繁殖成功的影响进行长期研究。研究结果发现:川金丝猴雌性繁殖成功与不同单元大小组别的平均值和与单元内成年雌性数目均呈显著负线性相关。川金丝猴的这一研究结果支持van Schaik(1983)的掠夺-组内取食竞争假说。组内扰乱竞争可能存在于川金丝猴社群中,并影响着雌性繁殖成功。
第二部分秦岭川金丝猴肢体偏好的研究
第四章秦岭川金丝猴单手和双手触达任务中手偏好行为的研究
2003年7月至2004年1月,利用焦点动物取样法和行为取样法对分布在秦岭北坡周至国家自然保护区玉皇庙地区的川金丝猴西梁群成年个体的手偏好行为进行观察研究。在单手取食触达和双手爬背触达两个行为任务中,焦点动物个体的左手偏好和右手偏好均在不同程度上分别有所体现,且无性别因素影响。64.29%的焦点动物个体在单手取食触达任务中表现出明显的手偏好,100%的焦点动物个体在双手爬背触达任务中表现出明显的手偏好。根据手偏好指数,71.43%的焦点动物个体在单手取食触达任务中表现出左手偏好,这在一定程度上支持姿势起源理论。川金丝猴在双手爬背触达任务中的手偏好强度明显高于其在单手取食触达任务中的手偏好强度,这说明身体姿势是川金丝猴手偏好表达的重要影响因素之一。
第五章秦岭川金丝猴四肢前行和双足前移任务中脚偏好行为的研究
2007年3月至2008年4月,利用焦点动物行为取样法对分布在秦岭北坡周至国家自然保护区玉皇庙地区的川金丝猴西梁群成年个体在移动中的脚偏好行为进行观察研究。本研究选用了四肢前行和双足前移两个行为任务,研究中选择21个成年个体作为焦点动物,其中包括9个成年雄性和12个成年雌性。在两个行为任务中,焦点动物个体的左脚偏好和右脚偏好均在不同程度上分别有所体现,且无性别因素影响。57.14%的焦点动物个体在四肢前行任务中表现出明显的脚偏好,100%的焦点动物个体在双足前移任务中表现出明显的脚偏好。根据脚偏好指数,61.90%的焦点动物个体在四肢前行任务中表现出右脚偏好,这在一定程度上支持姿势起源理论。川金丝猴在双足前移任务中的脚偏好强度明显高于其在四肢前行任务中的脚偏好强度,这说明身体姿势是川金丝猴脚偏好表达的重要影响因素之一。
第六章秦岭川金丝猴新生婴猴吸乳偏好和母猴携抱偏好行为的研究
2007年3月至4月,利用焦点动物行为取样法对分布在秦岭北坡周至国家自然保护区玉皇庙地区的野生川金丝猴西梁群个体的早期母婴互动行为不对称性(即婴猴吸乳偏好和母猴携抱偏好)进行观察研究。本研究选择10个焦点母婴对,结果发现,母猴携抱偏好和婴猴吸乳偏好在个体水平上表达显著,在群组水平上表达不显著;母猴携抱偏好与婴猴吸乳偏好之间相关不显著:野生川金丝猴在出生后一周内即表现出明显的头部转向偏好。
Part One
Reproductive Competition of Sichuan Snub-nosed Monkeys (Rhinopithecus roxellana) in the Qinling Mountains,China
Chapter 1:Impact of male takeover on intra-unit sexual interactions and subsequent inter-birth interval in Rhinopithecus roxellana
Data were collected on sexual interactions before and after a male takeover of a one male unit(OMU) of Sichuan snub-nosed monkeys(Rhinopithecus roxellana) at the Zhouzhi National Nature Reserve in the Qinling Mountains of China.The original unit consisted of an adult male,two adult and two subadult females,two female juveniles and a single infant.Following the takeover,the new resident male copulated with one adult female, which was not lactating.Subsequent to the disappearance of her infant,the second(lactating) female entered breeding condition and began to solicit copulation with the new resident male. Subadult females also engaged in matings with the new male.The new resident male was observed mating,on three occasions,with females in two other OMUs.These are the first observations of sexual behavior in free-ranging Sichuan snub-nosed monkeys after an OMU takeover.Sexual interactions play an important role in establishing relationships between the new male and resident females in the OMU.
Chapter 2:Mate competition and reproductive correlates of female dispersal in Rhinopithecus roxellana
Different mating systems in group-living animals have characteristic behavioral correlates that are primarily related to mate competition.Mate competition may push individuals to selectively make dispersal decisions for the purpose of maximizing of opportunities for reproduction.The Sichuan snub-nosed monkey(Rhinopithecus roxellana) is a polygynous primate species endemic to China.We provide the first data on female dispersal in a free-ranging group of R.roxellana at the Zhouzhi National Nature Reserve in the Qinling Mountains of China.Both adult and subadult female dispersal occurred. Immigration/emigration rates of adult females are higher than those of subadult females.Mate competition is one apparent driving force behind adult female dispersal,and inbreeding avoidance is the possible proximate factor influencing subadult female dispersal.Adult female R.roxellana employ various reproductive strategies related to dispersal,which may increase their reproductive success.
Chapter 3:Effect of group size and composition on female reproductive success in Rhinopithecus roxellana
Animal sociality evolves when gregarious benefits exceed those costs.Group size and group composition,as two basic dimensions of animal sociality,has various effects on the morphological and behavioral variables of group-living animals,such as individual reproductive success.This is the first study on the effect of group size and group composition on female reproductive success in the Rhinopithecus spp.From 2001 to 2008,our data was collected for a flee-ranging group of R.roxellana at the Zhouzhi National Nature Reserve in the Qinling Mountains of China.Female reproductive success of R.roxellana has a significantly linear correlation with both the mean size of each size group and the number of adult females.It suggests that the R.roxellana accords with the basic assumptions of the van Schaik(1983)'s predation-intragroup feeding competition hypothesis.This result raises the possibility that the within-group scramble competition works on female reproductive success in R.roxellana.
Part Two
Limb Laterality of Sichuan Snub-nosed Monkeys (Rhinopithecus roxellana) in the Qinling Mountains,China
Chapter 4:Hand preference during unimanual and bimanual reaching actions in Rhinopithecus roxellana
Hand preferences were investigated during one unimanual action(food-reaching) and one bimanual action(mount-reaching) in a free-ranging group of Sichuan snub-nosed monkeys(Rhinopithecus roxellana) at the Zhouzhi National Nature Reserve in the Qinling Mountains of China.Nine of fourteen individuals tested on the unimanual food-reaching action and all six individuals tested on the bimanual mount-reaching action exhibited a manual preference.Both significant right- and left-handed preferences were observed in the two actions.Sex did not affect either direction or strength of hand preference in the unimanual action.Hand preference for the bimanual action was stable over time,and the strength of hand preference was significantly stronger in the bimanual action than in the unimanual action.
Chapter 5:Foot Preference during quadrupedal and bipedal actions in Rhinopithecus roxellana
Foot preferences were investigated during the quadrupedal walking action and the bipedal shifting action for a free-ranging group of Sichuan snub-nosed monkeys (Rhinopithecus roxellana) at the Zhouzhi National Nature Reserve in the Qinling Mountains of China.Twelve of 21 individuals tested on the quadrupedal action and all 21 individuals tested on the bipedal action exhibited a significant foot preference.Both significant right- and left-footed preferences were observed and sex did not affect either direction or strength of foot preference in both actions.The finding that 61.90%of focal R.roxellana showed the right-foot preference both in quadrupedal action based on the footed index and in bipedal action based on the z-score are in partial agreement with the postural origin hypothesis on footedness.The strength of foot preference was significantly stronger in the bipedal action than in the quadrupedal action,supporting the view that posture could be a crucial factor influencing foot preference as well as hand preference in this species.
Chapter 6:Neonate nipple preference and maternal cradling laterality in Rhinopithecus roxellana
This study of Sichuan snub-nosed monkeys(Rhinopithecus roxellana) constitutes the first report on asymmetric patterns of early mother-infant interactions among Old World monkeys in the wild.It was found that neonate nipple preference and maternal cradling laterality are both evident on the individual level.Although there is no significant group-level preference direction,the group preference strength on both measures is evident.Moreover, neonate nipple preference is not significantly correlated with maternal cradling laterality on either LBI scores or the direction.Taken together,it is possible to suggest that wild R. roxellana showy independent orientation laterality from the first week of life.
第一章秦岭川金丝猴主雄替换后单元内雌雄两性繁殖策略的研究
2003年3月至2004年5月,利用焦点动物取样法和行为取样法对分布在秦岭北坡周至国家自然保护区玉皇庙地区川金丝猴西梁群关于主雄替换对单元内两性繁殖关系的影响进行研究。研究结果发现:主雄替换后,单元新主雄会倾向于自主选择非处于哺乳期成年雌性作为主要繁殖对象。新主雄通过单元外交配行为,不仅可以增加扩散自身基因的可能性,而且可以刺激单元内交配行为的发生进而增加自身繁殖成功机会。哺乳期雌性个体在其婴猴消失后的首个生育间隔显著短于研究群内未失婴猴雌性的平均生育间隔,这意味着野生成年雌性川金丝猴面对自身婴猴消失会采取积极的适应性繁殖策略。此外,亚成年雌性川金丝猴通过积极与新主雄建立良好繁殖关系进而获得其在成年后的繁殖成功。总之,野生川金丝猴单元新主雄、成年雌性和亚成年雌性个体面对主雄替换及之后的婴猴消失各自采取了不同的繁殖策略,这在一定程度上体现了该物种对于主雄替换这一避免近亲繁殖机制的行为适应性。
第二章秦岭川金丝猴雌性扩散背后的交配竞争及相关繁殖行为研究
2001年10月至2005年5月,对分布在秦岭北坡周至国家自然保护区玉皇庙地区川金丝猴西梁群关于雌性个体扩散与繁殖成功关系进行研究。研究结果发现:成年雌性个体和业成年雌性个体均发生扩散,成年雌性个体的迁出率和迁入率均高于亚成年雌性个体。交配竞争是成年雌性个体扩散的主要推动力,而亚成年雌性个体扩散可能是为了避免近亲繁殖。成年雌性个体在扩散过程中通过采用多样化的繁殖策略进而提高自身繁殖成功率。
第三章秦岭川金丝猴单元大小及组成对雌性繁殖成功影响的研究
动物的社会性在其群居利益大于群居代价时得以进化。群组大小和群组构成是评价动物社会性的两个基础衡量尺度,它们对雌性繁殖成功发挥重要影响作用。2001年10月至2008年5月,对分布在秦岭北坡周至国家自然保护区玉皇庙地区川金丝猴西梁群关于一雄多雌单元大小和组成对川金丝猴雌性繁殖成功的影响进行长期研究。研究结果发现:川金丝猴雌性繁殖成功与不同单元大小组别的平均值和与单元内成年雌性数目均呈显著负线性相关。川金丝猴的这一研究结果支持van Schaik(1983)的掠夺-组内取食竞争假说。组内扰乱竞争可能存在于川金丝猴社群中,并影响着雌性繁殖成功。
第二部分秦岭川金丝猴肢体偏好的研究
第四章秦岭川金丝猴单手和双手触达任务中手偏好行为的研究
2003年7月至2004年1月,利用焦点动物取样法和行为取样法对分布在秦岭北坡周至国家自然保护区玉皇庙地区的川金丝猴西梁群成年个体的手偏好行为进行观察研究。在单手取食触达和双手爬背触达两个行为任务中,焦点动物个体的左手偏好和右手偏好均在不同程度上分别有所体现,且无性别因素影响。64.29%的焦点动物个体在单手取食触达任务中表现出明显的手偏好,100%的焦点动物个体在双手爬背触达任务中表现出明显的手偏好。根据手偏好指数,71.43%的焦点动物个体在单手取食触达任务中表现出左手偏好,这在一定程度上支持姿势起源理论。川金丝猴在双手爬背触达任务中的手偏好强度明显高于其在单手取食触达任务中的手偏好强度,这说明身体姿势是川金丝猴手偏好表达的重要影响因素之一。
第五章秦岭川金丝猴四肢前行和双足前移任务中脚偏好行为的研究
2007年3月至2008年4月,利用焦点动物行为取样法对分布在秦岭北坡周至国家自然保护区玉皇庙地区的川金丝猴西梁群成年个体在移动中的脚偏好行为进行观察研究。本研究选用了四肢前行和双足前移两个行为任务,研究中选择21个成年个体作为焦点动物,其中包括9个成年雄性和12个成年雌性。在两个行为任务中,焦点动物个体的左脚偏好和右脚偏好均在不同程度上分别有所体现,且无性别因素影响。57.14%的焦点动物个体在四肢前行任务中表现出明显的脚偏好,100%的焦点动物个体在双足前移任务中表现出明显的脚偏好。根据脚偏好指数,61.90%的焦点动物个体在四肢前行任务中表现出右脚偏好,这在一定程度上支持姿势起源理论。川金丝猴在双足前移任务中的脚偏好强度明显高于其在四肢前行任务中的脚偏好强度,这说明身体姿势是川金丝猴脚偏好表达的重要影响因素之一。
第六章秦岭川金丝猴新生婴猴吸乳偏好和母猴携抱偏好行为的研究
2007年3月至4月,利用焦点动物行为取样法对分布在秦岭北坡周至国家自然保护区玉皇庙地区的野生川金丝猴西梁群个体的早期母婴互动行为不对称性(即婴猴吸乳偏好和母猴携抱偏好)进行观察研究。本研究选择10个焦点母婴对,结果发现,母猴携抱偏好和婴猴吸乳偏好在个体水平上表达显著,在群组水平上表达不显著;母猴携抱偏好与婴猴吸乳偏好之间相关不显著:野生川金丝猴在出生后一周内即表现出明显的头部转向偏好。
Part One
Reproductive Competition of Sichuan Snub-nosed Monkeys (Rhinopithecus roxellana) in the Qinling Mountains,China
Chapter 1:Impact of male takeover on intra-unit sexual interactions and subsequent inter-birth interval in Rhinopithecus roxellana
Data were collected on sexual interactions before and after a male takeover of a one male unit(OMU) of Sichuan snub-nosed monkeys(Rhinopithecus roxellana) at the Zhouzhi National Nature Reserve in the Qinling Mountains of China.The original unit consisted of an adult male,two adult and two subadult females,two female juveniles and a single infant.Following the takeover,the new resident male copulated with one adult female, which was not lactating.Subsequent to the disappearance of her infant,the second(lactating) female entered breeding condition and began to solicit copulation with the new resident male. Subadult females also engaged in matings with the new male.The new resident male was observed mating,on three occasions,with females in two other OMUs.These are the first observations of sexual behavior in free-ranging Sichuan snub-nosed monkeys after an OMU takeover.Sexual interactions play an important role in establishing relationships between the new male and resident females in the OMU.
Chapter 2:Mate competition and reproductive correlates of female dispersal in Rhinopithecus roxellana
Different mating systems in group-living animals have characteristic behavioral correlates that are primarily related to mate competition.Mate competition may push individuals to selectively make dispersal decisions for the purpose of maximizing of opportunities for reproduction.The Sichuan snub-nosed monkey(Rhinopithecus roxellana) is a polygynous primate species endemic to China.We provide the first data on female dispersal in a free-ranging group of R.roxellana at the Zhouzhi National Nature Reserve in the Qinling Mountains of China.Both adult and subadult female dispersal occurred. Immigration/emigration rates of adult females are higher than those of subadult females.Mate competition is one apparent driving force behind adult female dispersal,and inbreeding avoidance is the possible proximate factor influencing subadult female dispersal.Adult female R.roxellana employ various reproductive strategies related to dispersal,which may increase their reproductive success.
Chapter 3:Effect of group size and composition on female reproductive success in Rhinopithecus roxellana
Animal sociality evolves when gregarious benefits exceed those costs.Group size and group composition,as two basic dimensions of animal sociality,has various effects on the morphological and behavioral variables of group-living animals,such as individual reproductive success.This is the first study on the effect of group size and group composition on female reproductive success in the Rhinopithecus spp.From 2001 to 2008,our data was collected for a flee-ranging group of R.roxellana at the Zhouzhi National Nature Reserve in the Qinling Mountains of China.Female reproductive success of R.roxellana has a significantly linear correlation with both the mean size of each size group and the number of adult females.It suggests that the R.roxellana accords with the basic assumptions of the van Schaik(1983)'s predation-intragroup feeding competition hypothesis.This result raises the possibility that the within-group scramble competition works on female reproductive success in R.roxellana.
Part Two
Limb Laterality of Sichuan Snub-nosed Monkeys (Rhinopithecus roxellana) in the Qinling Mountains,China
Chapter 4:Hand preference during unimanual and bimanual reaching actions in Rhinopithecus roxellana
Hand preferences were investigated during one unimanual action(food-reaching) and one bimanual action(mount-reaching) in a free-ranging group of Sichuan snub-nosed monkeys(Rhinopithecus roxellana) at the Zhouzhi National Nature Reserve in the Qinling Mountains of China.Nine of fourteen individuals tested on the unimanual food-reaching action and all six individuals tested on the bimanual mount-reaching action exhibited a manual preference.Both significant right- and left-handed preferences were observed in the two actions.Sex did not affect either direction or strength of hand preference in the unimanual action.Hand preference for the bimanual action was stable over time,and the strength of hand preference was significantly stronger in the bimanual action than in the unimanual action.
Chapter 5:Foot Preference during quadrupedal and bipedal actions in Rhinopithecus roxellana
Foot preferences were investigated during the quadrupedal walking action and the bipedal shifting action for a free-ranging group of Sichuan snub-nosed monkeys (Rhinopithecus roxellana) at the Zhouzhi National Nature Reserve in the Qinling Mountains of China.Twelve of 21 individuals tested on the quadrupedal action and all 21 individuals tested on the bipedal action exhibited a significant foot preference.Both significant right- and left-footed preferences were observed and sex did not affect either direction or strength of foot preference in both actions.The finding that 61.90%of focal R.roxellana showed the right-foot preference both in quadrupedal action based on the footed index and in bipedal action based on the z-score are in partial agreement with the postural origin hypothesis on footedness.The strength of foot preference was significantly stronger in the bipedal action than in the quadrupedal action,supporting the view that posture could be a crucial factor influencing foot preference as well as hand preference in this species.
Chapter 6:Neonate nipple preference and maternal cradling laterality in Rhinopithecus roxellana
This study of Sichuan snub-nosed monkeys(Rhinopithecus roxellana) constitutes the first report on asymmetric patterns of early mother-infant interactions among Old World monkeys in the wild.It was found that neonate nipple preference and maternal cradling laterality are both evident on the individual level.Although there is no significant group-level preference direction,the group preference strength on both measures is evident.Moreover, neonate nipple preference is not significantly correlated with maternal cradling laterality on either LBI scores or the direction.Taken together,it is possible to suggest that wild R. roxellana showy independent orientation laterality from the first week of life.
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