矮岩羊种群生态学研究
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摘要
矮岩羊为国家Ⅱ级保护动物,特产于我国金沙江畔,主要分布在青藏高原东南缘的四川省巴塘县境内。2006年8月、2007年7～12月和2008年3～7月,我们在四川省竹巴笼自然保护区对矮岩羊行为谱、行为PAE编码、日活动节律、活动时间分配、集群特征和生境选择季节性变化进行了研究,主要结果如下:
针对行为的3要素,建立了矮岩羊行为的三位编码系统:依据矮岩羊的姿势、动作和环境建立P码、A码和E码,对一种特定的行为依其发生的环境,涉及的姿势和动作建立PAE编码系统。观察分辨和记录到矮岩羊的11种姿势,75种动作,14种活动环境,92种行为,分列于取食、反刍、休息、运动、排遗、调温、发情、交配、育幼、冲突、通讯、聚群和杂类等13大类行为。标明各种行为在雄性、雌性和幼体之间的相对发生频次以及发生季节,并建立了各行为要素码与行为序位码之间的联系。
按年龄、性别把矮岩羊种群分为成雄、成雌、亚成体和幼体4个组,采用焦点取样-瞬时记录法记录一次岩羊的行为。观察记录矮岩羊的取食、卧息、站立、运动和其它行为等5种行为,采用非参数检验的Mann-Whitney U检验和Kruskal-Wallis H检验对矮岩羊不同性别-年龄组以及季节变化差异进行了比较分析。结果表明:矮岩羊的日活动时间分配,取食(44.56±2.72%)时间比例最高,其次是卧息(25.54±3.19%),再次是站立(15.15±0.60%)和运动(11.39±0.55%),其它行为(3.40±0.31%)最低;一年中,8月取食时间比例最低(40.80%)、卧息最高(29.35%),12月取食最高(49.66%)、卧息最低(18.85%),站立、运动和其它行为变化不大;春秋冬3季均出现2个取食高峰和1个卧息高峰,夏季则出呈现3个取食高峰和2个卧息高峰,冬季取食时间比例(47.01%)最高、卧息(21.95%)最低,夏季则与之相反;成年雄性(4.40%)与成年雌性(2.80%)、成体(3.60%)与亚成体(2.70%)仅在其它行为上前者显著高于后者,而在取食、卧息、站立和运动上均无显著差异(P＞0.05);幼体在3个年龄组中取食(40.70%)和站立(12.10%)时间比例上均最低,卧息最高(32.30%),而在运动时间分配上成体、亚成体和幼体差异不显著(P＞0.05)。
采用样线法和定点观察法对矮岩羊2007年和2008年集群进行了研究。结果表明,2007年观察到34群217只矮岩羊,集群平均大小为6.4±4.9只;2008年16群135只羊,集群平均大小为8.4±7.7只。集群分为雄性群(2007年:8.8%;2008年:6.3%)、雌性群(2.9%;0)、混合群(70.6%:81.2%)、母仔群(8.8%;12.5%)和孤羊(8.8%;0)5种类型。在不同季节,5种集群类型比例都有所变化,差异极显著(P＜0.01)。不同大小集群分布之间和不同类型集群大小之间也有显著或极显著差异,其中2～10只集群的集群数在2007年占73.5%、在2008年占75.0%,呈集小群特征。汇总不同年份的集群,ANVOA表明各年集群平均大小差异显著(P＜0.05),LSD法多重比较发现,相对2000年集群平均大小2007年(P＜0.05)和2008年(P＜0.01)均显著上升。年龄组结构中,成体、亚成体和幼体个体数比为100∶26.4∶28.6(2007年)和100∶24.7∶20.4(2008年)。雌雄性比2007年为100∶55.7、2008年为100∶56.6,年间差异不显著(P＞0.05)。
采用样线调查法对矮岩羊生境选择季节性交化进行了研究。结果表明,矮岩羊秋冬季对17种生境因子有选择性,倾向选择中等以上避风性,石质或便石质活动基底,灌丛盖度10%～40%、种数＞4种、高度＞100 cm,草种数＞5种、高度＜20 cm或＞40 cm,坡度＞60°的南坡或西坡,隐蔽级＜30%或＞60%,水源距离＞500 m,裸岩距离＜2m,林缘距离1000～1500 m,1级风,海拔高度＜2800 m,人为干扰距离500～1000 m,动物干扰距离80～150 m的生境。而春夏季对18种生境因子有选择性,倾向选择凹坡或凸坡,中等及以下避风性,石质或便石质活动基底,灌丛盖度10%～40%、种数4～5种、高度＞100 cm,草盖度＞30%,种数4～5种、高度＜20cm或＞40 cm,坡度＞60°的东坡、西坡或东南坡,隐蔽级＞60%,水源距离500～1000 m,裸岩距离＜2m,林缘距离＜1000 m,海拔高度＞2600 m,人为干扰距离＞500 m,动物干扰距离＞80 m的生境。因子分析表明,影响秋冬季生境选择的主要因子为人为干扰距离、海拔高度、林缘距离、水源距离、灌丛高度、隐蔽级、风级和动物干扰距离等生境因子,而影响春夏季生境选择的主要因子为人为干扰距离、海拔高度、林缘距离、草高度、草盖度、坡度、坡形等生境因子。
As an endangered species,dwarf blue sheep was listed in ClassⅡof the National Key Protected Animals in China.The sheep is an endemic species only inhabiting in Jinsha River gorge at the southeastern Tibetan Plateau(mainly in Zhubalong,south of Batang County).We conducted an investigation into population ecology of dwarf blue sheep in Sichuan Zhubalong Nature Reserve in August 2006 and July-December 2007 and March-June 2008.We described and studied the ethogram,PAE behavior coding,activity rhythms,time budget,group pattern of dwarf blue sheep and seasonal change in habitat selection.The main results were summarized as follows:
We established a behavioral functional classification and coding system for the behavior of dwarf blue sheep.'Posture','Act' and 'Environment' were coded with PAE via distinguish basic unit of animal behavior and analyze behavioral hierarchy.11 postures,75 acts and 92 behaviors of dwarf blue sheep were recorded and classified as follows:feeding,ruminating,resting,moving, urinating/defecating,thermo-regulating,rutting and estrous behavior,mating,parental behavior, communication behavior,aggregation behavior and miscellaneous behavior.Frequency and seasonal occurrence of different individual behavior within sex-age classes were also recorded in PAE system.
Population individuals of dwarf blue sheep were divided into 4 sex-age classes according to their ages and sexes.Activities of the sheep such as feeding,lying,moving,standing and other behavior,were recorded by using a modified version of traditional focal and instantaneous sampling. We used nonparametric Mann-Whitney U and Kruskal-Wallis H tests to compare differences in the time budgets among different sex-age classes and different seasons.The sheep spent the majority of their time feeding and lying,44.56±2.72%and 25.54±3.19%of all activities respectively.The remnant time was devoted to other three activities,such as standing(15.15±0.60%),moving(11.39±0.55%) and other behaviors(3.40±0.31%).The percentage of daytime spent in feeding reached its lowest level(40.80%) in August and highest level(49.66%) in December.In contrast,lying reached its highest level(29.35%) in August and lowest level(18.85%) in December.Dwarf blue sheep had two daily peaks of feeding(early morning and late afternoon) and one peak of lying in midday during spring,autumn and winter,but also showed three peaks of feeding(early morning, midday and late afternoon) and two peaks of lying(at 10:00-11:30 and 13:00-14:30) during summer. Feeding remained at relatively higher levels(47.01%) and lying kept at relatively lower levels (21.95%) in winter,and contrarily in summer.No significant differences in general activity budgets between adult males and females and between adults and subadults were found,but adult males were found to spend more time in other behaviors than adult females and subadults.Kids spent significant more time in lying and less time in feeding and standing than did other sex-age classes,but no significant differences in daytime spent moving among all sex-age classes.
We investigated group patterns of dwarf blue sheep by using methods of transect censuses and fixed point observations.We observed 34 groups consisting of 217 individuals with a mean group size of 6.4±4.9 in 2007 and 16 groups comprising 135 individuals with a mean group size of 8.4±7.7 in 2008.Sheep groups can be classified into five types:male only(2007:8.8%of total groups,2008:6.3%),female only(2.9%,0),mixed(70.6%,81.2%),mother-offspring(8.8%,12.5%), and solitary individual(8.8%,0).These group types varied seasonally in proportion(P＜0.01).These types also differed in distribution(P＜0.01) and in size(P＜0.01).Groups of 2-10 individuals were predominant(73.5%of all groups in 2007 and 75.0%in 2008).We collected population data from several recent articles(1997,1998,2000,2006) to analyze mean group size and sex ratio dynamics using the ANOVA technique.The results indicated that group size was very variable(P＜0.05) and the mean group size was significantly larger in 2007(P＜0.05) and in 2008(P＜0.01) than in 2000. The adult:juvenile:lamb ratio was 100:26.4:28.6 in 2007 and 100:24.7:20.4 in 2008.The female:male ratio(100:55.7 in 2007 and 100:56.6 in 2008) remained stable(P＞0.05).Group patterns of the sheep can be influenced by benefits in group living,biological characteristics, breeding cycle,competition for resources,habitat type or topography,predation risk,and human disturbance,group patterns of the sheep implied a survival strategy.
We set up 10 line transects to investigate habitat selection of dwarf blue sheep and the Chi-Square Test and Mann-Whitey U Test were used to analyze the raw data.The sheep preferred areas during autumn-winter with the following attributes:well-sheltered from wind,lithoid or partially lithoid substrates,10%-40%shrub canopy cover,5 or more shrub species,tall shrubs(＞100 cm),6 or more herbaceous species,short(＜20 cm) or tall(＞40 cm) grasses,cliffy(＞60°) and sunny slope,low(＜30%) or high(＞60%) levels of concealment,moderate distance to water source(＞500 m),short distance to bare rock(＜2 m),moderate distance to forest edge(1000-1500 m),low altitude (＜2800 m),moderate distance to human disturbance(500-1000 m) and animal disturbance(80-150 m).While the sheep preferred areas during spring-summer with the following attributes:concave or convex slope form,well or worse -sheltered from wind,lithoid or partially lithoid substrates. 10%-40%shrub canopy cover,biggish herbaceous canopy(＞30%),4-5 shrub species,tall shrubs (＞100 cm),4-5 herbaceous species,short(＜20 cm) or tall grasses(＞40 cm),steep(＞60°) and sunny slope,moderate distance to water source(＞500 m),long distance to animal disturbance(＞80 m), long distance to human disturbance(＞500 m),high(＞60%) levels of concealment,short distance to bare rock(＜2 m),short distance to forest edge(＜1000 m),medium altitude(＞2600 m).The results of factor analysis indicated that autumn-winter habitat selection was mainly influenced by distance to human disturbance,altitude,distance to forest edge,distance to water source,shrub height,level of concealment,wind speed and distance to animal disturbance.While spring-summer habitat selection was mainly influenced by distance to human disturbance,altitude,distance to forest edge. herbaceous height,herbaceous canopy,slope gradient,slope form.
针对行为的3要素,建立了矮岩羊行为的三位编码系统:依据矮岩羊的姿势、动作和环境建立P码、A码和E码,对一种特定的行为依其发生的环境,涉及的姿势和动作建立PAE编码系统。观察分辨和记录到矮岩羊的11种姿势,75种动作,14种活动环境,92种行为,分列于取食、反刍、休息、运动、排遗、调温、发情、交配、育幼、冲突、通讯、聚群和杂类等13大类行为。标明各种行为在雄性、雌性和幼体之间的相对发生频次以及发生季节,并建立了各行为要素码与行为序位码之间的联系。
按年龄、性别把矮岩羊种群分为成雄、成雌、亚成体和幼体4个组,采用焦点取样-瞬时记录法记录一次岩羊的行为。观察记录矮岩羊的取食、卧息、站立、运动和其它行为等5种行为,采用非参数检验的Mann-Whitney U检验和Kruskal-Wallis H检验对矮岩羊不同性别-年龄组以及季节变化差异进行了比较分析。结果表明:矮岩羊的日活动时间分配,取食(44.56±2.72%)时间比例最高,其次是卧息(25.54±3.19%),再次是站立(15.15±0.60%)和运动(11.39±0.55%),其它行为(3.40±0.31%)最低;一年中,8月取食时间比例最低(40.80%)、卧息最高(29.35%),12月取食最高(49.66%)、卧息最低(18.85%),站立、运动和其它行为变化不大;春秋冬3季均出现2个取食高峰和1个卧息高峰,夏季则出呈现3个取食高峰和2个卧息高峰,冬季取食时间比例(47.01%)最高、卧息(21.95%)最低,夏季则与之相反;成年雄性(4.40%)与成年雌性(2.80%)、成体(3.60%)与亚成体(2.70%)仅在其它行为上前者显著高于后者,而在取食、卧息、站立和运动上均无显著差异(P＞0.05);幼体在3个年龄组中取食(40.70%)和站立(12.10%)时间比例上均最低,卧息最高(32.30%),而在运动时间分配上成体、亚成体和幼体差异不显著(P＞0.05)。
采用样线法和定点观察法对矮岩羊2007年和2008年集群进行了研究。结果表明,2007年观察到34群217只矮岩羊,集群平均大小为6.4±4.9只;2008年16群135只羊,集群平均大小为8.4±7.7只。集群分为雄性群(2007年:8.8%;2008年:6.3%)、雌性群(2.9%;0)、混合群(70.6%:81.2%)、母仔群(8.8%;12.5%)和孤羊(8.8%;0)5种类型。在不同季节,5种集群类型比例都有所变化,差异极显著(P＜0.01)。不同大小集群分布之间和不同类型集群大小之间也有显著或极显著差异,其中2～10只集群的集群数在2007年占73.5%、在2008年占75.0%,呈集小群特征。汇总不同年份的集群,ANVOA表明各年集群平均大小差异显著(P＜0.05),LSD法多重比较发现,相对2000年集群平均大小2007年(P＜0.05)和2008年(P＜0.01)均显著上升。年龄组结构中,成体、亚成体和幼体个体数比为100∶26.4∶28.6(2007年)和100∶24.7∶20.4(2008年)。雌雄性比2007年为100∶55.7、2008年为100∶56.6,年间差异不显著(P＞0.05)。
采用样线调查法对矮岩羊生境选择季节性交化进行了研究。结果表明,矮岩羊秋冬季对17种生境因子有选择性,倾向选择中等以上避风性,石质或便石质活动基底,灌丛盖度10%～40%、种数＞4种、高度＞100 cm,草种数＞5种、高度＜20 cm或＞40 cm,坡度＞60°的南坡或西坡,隐蔽级＜30%或＞60%,水源距离＞500 m,裸岩距离＜2m,林缘距离1000～1500 m,1级风,海拔高度＜2800 m,人为干扰距离500～1000 m,动物干扰距离80～150 m的生境。而春夏季对18种生境因子有选择性,倾向选择凹坡或凸坡,中等及以下避风性,石质或便石质活动基底,灌丛盖度10%～40%、种数4～5种、高度＞100 cm,草盖度＞30%,种数4～5种、高度＜20cm或＞40 cm,坡度＞60°的东坡、西坡或东南坡,隐蔽级＞60%,水源距离500～1000 m,裸岩距离＜2m,林缘距离＜1000 m,海拔高度＞2600 m,人为干扰距离＞500 m,动物干扰距离＞80 m的生境。因子分析表明,影响秋冬季生境选择的主要因子为人为干扰距离、海拔高度、林缘距离、水源距离、灌丛高度、隐蔽级、风级和动物干扰距离等生境因子,而影响春夏季生境选择的主要因子为人为干扰距离、海拔高度、林缘距离、草高度、草盖度、坡度、坡形等生境因子。
As an endangered species,dwarf blue sheep was listed in ClassⅡof the National Key Protected Animals in China.The sheep is an endemic species only inhabiting in Jinsha River gorge at the southeastern Tibetan Plateau(mainly in Zhubalong,south of Batang County).We conducted an investigation into population ecology of dwarf blue sheep in Sichuan Zhubalong Nature Reserve in August 2006 and July-December 2007 and March-June 2008.We described and studied the ethogram,PAE behavior coding,activity rhythms,time budget,group pattern of dwarf blue sheep and seasonal change in habitat selection.The main results were summarized as follows:
We established a behavioral functional classification and coding system for the behavior of dwarf blue sheep.'Posture','Act' and 'Environment' were coded with PAE via distinguish basic unit of animal behavior and analyze behavioral hierarchy.11 postures,75 acts and 92 behaviors of dwarf blue sheep were recorded and classified as follows:feeding,ruminating,resting,moving, urinating/defecating,thermo-regulating,rutting and estrous behavior,mating,parental behavior, communication behavior,aggregation behavior and miscellaneous behavior.Frequency and seasonal occurrence of different individual behavior within sex-age classes were also recorded in PAE system.
Population individuals of dwarf blue sheep were divided into 4 sex-age classes according to their ages and sexes.Activities of the sheep such as feeding,lying,moving,standing and other behavior,were recorded by using a modified version of traditional focal and instantaneous sampling. We used nonparametric Mann-Whitney U and Kruskal-Wallis H tests to compare differences in the time budgets among different sex-age classes and different seasons.The sheep spent the majority of their time feeding and lying,44.56±2.72%and 25.54±3.19%of all activities respectively.The remnant time was devoted to other three activities,such as standing(15.15±0.60%),moving(11.39±0.55%) and other behaviors(3.40±0.31%).The percentage of daytime spent in feeding reached its lowest level(40.80%) in August and highest level(49.66%) in December.In contrast,lying reached its highest level(29.35%) in August and lowest level(18.85%) in December.Dwarf blue sheep had two daily peaks of feeding(early morning and late afternoon) and one peak of lying in midday during spring,autumn and winter,but also showed three peaks of feeding(early morning, midday and late afternoon) and two peaks of lying(at 10:00-11:30 and 13:00-14:30) during summer. Feeding remained at relatively higher levels(47.01%) and lying kept at relatively lower levels (21.95%) in winter,and contrarily in summer.No significant differences in general activity budgets between adult males and females and between adults and subadults were found,but adult males were found to spend more time in other behaviors than adult females and subadults.Kids spent significant more time in lying and less time in feeding and standing than did other sex-age classes,but no significant differences in daytime spent moving among all sex-age classes.
We investigated group patterns of dwarf blue sheep by using methods of transect censuses and fixed point observations.We observed 34 groups consisting of 217 individuals with a mean group size of 6.4±4.9 in 2007 and 16 groups comprising 135 individuals with a mean group size of 8.4±7.7 in 2008.Sheep groups can be classified into five types:male only(2007:8.8%of total groups,2008:6.3%),female only(2.9%,0),mixed(70.6%,81.2%),mother-offspring(8.8%,12.5%), and solitary individual(8.8%,0).These group types varied seasonally in proportion(P＜0.01).These types also differed in distribution(P＜0.01) and in size(P＜0.01).Groups of 2-10 individuals were predominant(73.5%of all groups in 2007 and 75.0%in 2008).We collected population data from several recent articles(1997,1998,2000,2006) to analyze mean group size and sex ratio dynamics using the ANOVA technique.The results indicated that group size was very variable(P＜0.05) and the mean group size was significantly larger in 2007(P＜0.05) and in 2008(P＜0.01) than in 2000. The adult:juvenile:lamb ratio was 100:26.4:28.6 in 2007 and 100:24.7:20.4 in 2008.The female:male ratio(100:55.7 in 2007 and 100:56.6 in 2008) remained stable(P＞0.05).Group patterns of the sheep can be influenced by benefits in group living,biological characteristics, breeding cycle,competition for resources,habitat type or topography,predation risk,and human disturbance,group patterns of the sheep implied a survival strategy.
We set up 10 line transects to investigate habitat selection of dwarf blue sheep and the Chi-Square Test and Mann-Whitey U Test were used to analyze the raw data.The sheep preferred areas during autumn-winter with the following attributes:well-sheltered from wind,lithoid or partially lithoid substrates,10%-40%shrub canopy cover,5 or more shrub species,tall shrubs(＞100 cm),6 or more herbaceous species,short(＜20 cm) or tall(＞40 cm) grasses,cliffy(＞60°) and sunny slope,low(＜30%) or high(＞60%) levels of concealment,moderate distance to water source(＞500 m),short distance to bare rock(＜2 m),moderate distance to forest edge(1000-1500 m),low altitude (＜2800 m),moderate distance to human disturbance(500-1000 m) and animal disturbance(80-150 m).While the sheep preferred areas during spring-summer with the following attributes:concave or convex slope form,well or worse -sheltered from wind,lithoid or partially lithoid substrates. 10%-40%shrub canopy cover,biggish herbaceous canopy(＞30%),4-5 shrub species,tall shrubs (＞100 cm),4-5 herbaceous species,short(＜20 cm) or tall grasses(＞40 cm),steep(＞60°) and sunny slope,moderate distance to water source(＞500 m),long distance to animal disturbance(＞80 m), long distance to human disturbance(＞500 m),high(＞60%) levels of concealment,short distance to bare rock(＜2 m),short distance to forest edge(＜1000 m),medium altitude(＞2600 m).The results of factor analysis indicated that autumn-winter habitat selection was mainly influenced by distance to human disturbance,altitude,distance to forest edge,distance to water source,shrub height,level of concealment,wind speed and distance to animal disturbance.While spring-summer habitat selection was mainly influenced by distance to human disturbance,altitude,distance to forest edge. herbaceous height,herbaceous canopy,slope gradient,slope form.
引文
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