贺兰山岩羊的种群动态及警戒行为研究
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摘要
随着越来越多的生态旅游和户外活动,人们对野生动物的干扰在逐步增加。然而,避免人为干扰对野生动物的伤害却不能简单地将有野生动物的地方全部封闭起来。一方面,生态旅游能够为野生动物保护提供部分资金；另一方面,加强环境保护的目的之一也是要能够为人们提供更多更好的休闲娱乐场所,以提高人们的生活质量。所以野生动物管理者和保护生物学工作者都迫切地希望找到一种人和野生动物和谐共处的措施,而要达成这一目标,就必须针对野生动物对人为干扰的反应机制进行研究。
不同的物种具有不一样的对人为干扰的反应机制,同一物种也会因与人类接触历史的不同而产生不一致的逃跑方式。贺兰山是岩羊(Pseudois nayaur)的分布北界,岩羊的种群密度较大,又多活动于低山环境中,因此便于进行不同程度人为扰下的逃跑行为研究。同时,本实验室的研究人员已经对该地区岩羊的集群行为、家域、种群密度等进行了比较充分的工作,积累了比较丰富的有关岩羊的生态学资料。这些资料对岩羊逃跑行为的研究来说,无论是从实验设计,还是对现象的解释,都提供了一个良好的基础。
不同密度的野生动物面临着不同程度的种内竞争,而激烈程度不同的种内竞争又决定了在受到干扰时是否有足够的可替代性适宜生境,这些都影响着动物逃跑时机的决策。因此,种群密度对野生动物的警戒行为至关重要。基于这个原因,笔者也根据前几年所积累的数据,对宁夏贺兰山岩羊的种群动态作出了分析,并在此基础上,分析岩羊的警戒行为模式。现将结果报道如下：
1、从2010年3月到12月,我们在保护区内划定36条样线,并分季节进行了4次调查,运用Distance6.0软件进行数量估计,结果显示保护区内的岩羊密度约为6.08(5.01-7.38)只/km2,各个季节的估计密度从5.63只/km2到6.48只／km2,基本上没有季节上的变化(χ2=0.00,df=3,P=1.00).整个保护区内岩羊在2010年的数量约为11,767(9,696-14,282)头,比2003年的岩羊数量略有增加(Z=1.98,P=0.048)。探测函数的结果显示在距离观测人员较近的地方(200m以内)就能够观测到大部分岩羊。
2、通过2004到2009年对岩羊数量变化的监测,我们发现岩羊在保护区范围内的分布并不均匀,从而根据这6年的整体遇见率将整个保护区划分为非分布区、低密度区、中密度区和高密度区。非分布区主要分布在地势高差太小、人为干扰较大且位于保护区的边缘部分。低密度区主要在保护区的北段,中密度区在保护区的南段,而高密度区集中在保护区的中部,降雨量的大小和植被覆盖率可能是影响岩羊分布密度的重要原因。
3、单因变量多因素方差(general linear models:univariate)分析表明岩羊遇见率的大小主要决定于地区差异(F=55.96,P=0.00),年际间的变化不大(F=0.21,P=0.96),且两者之间没有交互效应(F=0.77,P=0.66)；群大小也没有显著性的年际差异(F=0.48,P=0.79),且与地区之间没有交互效应(F=0.71,P=0.71),但不同密度区之间的群大小却差异显著(F=10.75,P=0.00),高密度区(Mean±SD:8.48+5.55)比中(6.12+5.25)、低密度区(4.60±3.64)岩羊的集群要大(P<0.05)。群大小与遇见率之间存在显著的相关性(线性回归：linear regression model: Y=0.80+0.36遇见率,radj2=0.46, F=138.28, P<0.001).
4、保护区内的岩羊种群数量有较大的年际间变化(χ2=972.21, df=5, P<0.001),这个变化与前一年的降水日数呈显著的正相关(r=0.93,P=0.008)。降水日数可能是该保护区内岩羊种群数量变化的一个重要原因。在连续的6年时间里,雌雄比、雌幼比、成幼比在各个密度区间里都没有太大的变化,但三个密度区间的雌幼比和成幼比却差异明显(P<0.05)。
5、根据保护区内不同地区的人为干扰状况,我们将整个研究区域分为5个部分,2个人为活动较多的公路(A4和A5)及3个人为干扰较少的区域(A1、A2、A3)。采用初始逃跑距离(Flight initiation distance:FID)和停留距离(Finaldistance:FD)对岩羊的逃跑行为进行量度。根据平均群大小将群大小分为4＼个等级。然后用单因变量多因素方差(general linear models:univariate)分析不同地区、不同季节、不同群大小间岩羊所发生的警戒行为差异。通过626群岩羊的逃跑行为研究,我们发现：岩羊在不同的地区有着不一致的初始逃跑殖离(F=89.11,P<0.01)和停留距离(F=98.02,P<0.01)。岩羊在人为干扰最少的A1区域保持着与人最远的距离,而在两个干扰较严重的地区A4和A5,岩羊却有着最近的初始逃跑距离和停留距离。初始逃跑距离在季节间的差异不大(F=0.88,P=0.45),群大小级别间的差异也不大(F=2.27,P=0.08),但停留距离却受这两个因素的影响(季节：F=3.76,P=0.01；群大小：F=3.54,P=0.02)
6、两个人为干扰严重地区的多数岩羊在与人相遇后都是缓慢逃离,而其它三个地区的大多数岩羊却是快速逃离。初始逃跑距离和停留距离均与干扰人数呈显著负相关,却与干扰源距离之间成显著正相关关系。在人为干扰比较严重的时间段(10：00-15：00h),在五个地区间有着相似的遇见率(F=0.57,P=0.69), A3.A4和A5地区岩羊的估计密度相似(P>0.05),但都比A1和A2地区高(F=5.67,P=0.01),而A1和A2间的密度差别不大(P>0.05),所有五个地区岩羊都没有季节上的数量变化(F=0.64,P=0.60)。
With the increasing of off-road and outdoor ecotourism, wildlife encountered more and more harassment from the human. While we cannot only prohibit human go into protected area simply. On the one hand, we should provide more place to the people, on the other hand, ecotourism can produce money on protection. Wildlife managers and conservationists are looking forward to find a way to make the coexistence of the wildlife and ecotourism. But it was necessary to know how about the flight responses of the wildlife at first.
Different species had different flight responses to anthropogenic disturbance, the different history of contacting with human also had effect on the flight responses. Helan Mountain is the north boundary of distribution area for blue sheep (Pseudois nayaur). the density is much higher than any other areas, and was inhabited in lower elevation, so it is easier to contact in this place. At the same time, researchers in our lab had done works on the group type, home range size, population dynamic, life table, and so on, there are some basic data on blue sheep in Ningxia Helan Mountain National Nature Reserve. That can provide ways to explain the flight responses of the species.
Wildlife in different density encountered different intraspecific competition, which had some effect on the decision in flight responses of wildlife, for the substitute place was an important factor on the trade-off of stay or flee. We analyzed population dynamics of blue sheep in recent years, and then observed flight responses of them. The results can be seen below:
1、We surveyed36line transects in the reserve seasonally from October to December,2010, used Distance6.0to estimate the density of blue sheep, the result was6.08(5.01～7.38)sheep/km2. The density from5.63sheep/km2to6.48sheep/km2in four seasons, but had no significant difference in the season of the reserve (χ2=0.00, df=3, P=1.00). Population size about11,767(9,696～14,282) in the reserve, a little higher than that in2003(Z=1.98, P=0.048). Detection function showed that most of blue sheep could be encountered near than200m.
2^Long-term studies of population dynamics are important for conservation biology and wildlife management. We conducted2,790line transects and observed12,516blue sheep from2004to2009in the Ningxia Helan Mountain National Nature Reserve of China, and found the distribution of blue sheep was uneven in the reserve, then classified the reserve into no distribution, low density area, middle density area and high density area according to the encounter rates of the6years. No distribution area lied in the edge of reserve, the difference in elevation was less between the highest and lowest place and with heavier anthropogenic disturbance; low density area mainly in the north part of the reserve, middle density area mainly in the south part, high density area mainly in the middle part. Rainfall and vegetation cover may influent the density of blue sheep in the reserve.
3、Result of general linear models (GLM:univariate) showed that the encounter rates was mainly decided by the area (F=55.96, P=0.00), not varied among years (F=0.21, P=0.96), and there had no interaction between them. Year also had no effects on the group size (F=0.48, P=0.19) and no interaction with density area (F=0.71, P=0.71), but group size different significant among density areas (F=10.75, P=0.00). Group size in high density area (Mean±SD:8.48±5.55) was bigger than that in middle density area (6.12±5.25) and low density area (4.60±3.64). Group size had significant correlations with encounter rates (linear regression model:Y=0.80+0.36encounter rate, radj2=0.46, F=138.28, P<0.001).
4. Population size of the blue sheep fluctuated from12,375in2004to8,188in2006then13,344in2009, but did not ascend or decline continuously. Population size varied among6years in the reserve (χ2=972.21, df=5, P<0.001), the variation accompanied with rainy days during previous year (r=0.93, P=0.008). So rainy days maybe the cause of variation in blue sheep population size. Ratios between female and male, female and juvenile, adult and juvenile had no difference in each density area among6years, but varied among density areas (P<0.05). Adult female blue sheep was the most animal in the reserve.
5^According to the anthropogenic disturbance, we categorized the reserve into5parts:two on-trail area (A4:Dashuigou on-trail; A5:Suyukou on-trail) and3parts with less human disturbance (Al:Dashuigou off-trail; A2:Maliankou; A3: Suyukou off-trail). We used flight initiation distance (FID)and final distance (FD) to evaluate the flight responses of blue sheep. Used general linear models (univariate) to analyze the effects of area, group size and season on the flight responses. With626groups that encountered from March to December,2010, we found blue sheep had different flight initiation distance among areas (F=0.88, P=0.45), also did in the final distance(F=98.02, P<0.01). In Dashuigou off-trail remained farthest from human, but had the nearest distance in two on-trail areas. Flight initiation distance did not vary among group size (F=2.27, P=0.08), either among seasons (F=0.88, P=0.45), but final distance did (season:F=3.76, P=0.01; group size:F=3.54, P=0.02).
6、Majority of the blue sheep fled slowly when encountered with human in the two on-trail area, but majority fled fast in other three areas. Both the flight initiation distance and final distance had negative correlation with number of tourists, and had positive correlation with distance from anthropogenic disturbance. In the time with heavier anthropogenic disturbance (10:00-15:00h), there are similar encounter rates among5areas (F=0.57, P=0.69). Al and A2(with less anthropogenic disturbance) had lower density of blue sheep than other three areas (F=5.67, P=0.01), but had no difference between them (P>0.05). There had no variation in density among seasons (F=0.64, P=0.60).
不同的物种具有不一样的对人为干扰的反应机制,同一物种也会因与人类接触历史的不同而产生不一致的逃跑方式。贺兰山是岩羊(Pseudois nayaur)的分布北界,岩羊的种群密度较大,又多活动于低山环境中,因此便于进行不同程度人为扰下的逃跑行为研究。同时,本实验室的研究人员已经对该地区岩羊的集群行为、家域、种群密度等进行了比较充分的工作,积累了比较丰富的有关岩羊的生态学资料。这些资料对岩羊逃跑行为的研究来说,无论是从实验设计,还是对现象的解释,都提供了一个良好的基础。
不同密度的野生动物面临着不同程度的种内竞争,而激烈程度不同的种内竞争又决定了在受到干扰时是否有足够的可替代性适宜生境,这些都影响着动物逃跑时机的决策。因此,种群密度对野生动物的警戒行为至关重要。基于这个原因,笔者也根据前几年所积累的数据,对宁夏贺兰山岩羊的种群动态作出了分析,并在此基础上,分析岩羊的警戒行为模式。现将结果报道如下：
1、从2010年3月到12月,我们在保护区内划定36条样线,并分季节进行了4次调查,运用Distance6.0软件进行数量估计,结果显示保护区内的岩羊密度约为6.08(5.01-7.38)只/km2,各个季节的估计密度从5.63只/km2到6.48只／km2,基本上没有季节上的变化(χ2=0.00,df=3,P=1.00).整个保护区内岩羊在2010年的数量约为11,767(9,696-14,282)头,比2003年的岩羊数量略有增加(Z=1.98,P=0.048)。探测函数的结果显示在距离观测人员较近的地方(200m以内)就能够观测到大部分岩羊。
2、通过2004到2009年对岩羊数量变化的监测,我们发现岩羊在保护区范围内的分布并不均匀,从而根据这6年的整体遇见率将整个保护区划分为非分布区、低密度区、中密度区和高密度区。非分布区主要分布在地势高差太小、人为干扰较大且位于保护区的边缘部分。低密度区主要在保护区的北段,中密度区在保护区的南段,而高密度区集中在保护区的中部,降雨量的大小和植被覆盖率可能是影响岩羊分布密度的重要原因。
3、单因变量多因素方差(general linear models:univariate)分析表明岩羊遇见率的大小主要决定于地区差异(F=55.96,P=0.00),年际间的变化不大(F=0.21,P=0.96),且两者之间没有交互效应(F=0.77,P=0.66)；群大小也没有显著性的年际差异(F=0.48,P=0.79),且与地区之间没有交互效应(F=0.71,P=0.71),但不同密度区之间的群大小却差异显著(F=10.75,P=0.00),高密度区(Mean±SD:8.48+5.55)比中(6.12+5.25)、低密度区(4.60±3.64)岩羊的集群要大(P<0.05)。群大小与遇见率之间存在显著的相关性(线性回归：linear regression model: Y=0.80+0.36遇见率,radj2=0.46, F=138.28, P<0.001).
4、保护区内的岩羊种群数量有较大的年际间变化(χ2=972.21, df=5, P<0.001),这个变化与前一年的降水日数呈显著的正相关(r=0.93,P=0.008)。降水日数可能是该保护区内岩羊种群数量变化的一个重要原因。在连续的6年时间里,雌雄比、雌幼比、成幼比在各个密度区间里都没有太大的变化,但三个密度区间的雌幼比和成幼比却差异明显(P<0.05)。
5、根据保护区内不同地区的人为干扰状况,我们将整个研究区域分为5个部分,2个人为活动较多的公路(A4和A5)及3个人为干扰较少的区域(A1、A2、A3)。采用初始逃跑距离(Flight initiation distance:FID)和停留距离(Finaldistance:FD)对岩羊的逃跑行为进行量度。根据平均群大小将群大小分为4＼个等级。然后用单因变量多因素方差(general linear models:univariate)分析不同地区、不同季节、不同群大小间岩羊所发生的警戒行为差异。通过626群岩羊的逃跑行为研究,我们发现：岩羊在不同的地区有着不一致的初始逃跑殖离(F=89.11,P<0.01)和停留距离(F=98.02,P<0.01)。岩羊在人为干扰最少的A1区域保持着与人最远的距离,而在两个干扰较严重的地区A4和A5,岩羊却有着最近的初始逃跑距离和停留距离。初始逃跑距离在季节间的差异不大(F=0.88,P=0.45),群大小级别间的差异也不大(F=2.27,P=0.08),但停留距离却受这两个因素的影响(季节：F=3.76,P=0.01；群大小：F=3.54,P=0.02)
6、两个人为干扰严重地区的多数岩羊在与人相遇后都是缓慢逃离,而其它三个地区的大多数岩羊却是快速逃离。初始逃跑距离和停留距离均与干扰人数呈显著负相关,却与干扰源距离之间成显著正相关关系。在人为干扰比较严重的时间段(10：00-15：00h),在五个地区间有着相似的遇见率(F=0.57,P=0.69), A3.A4和A5地区岩羊的估计密度相似(P>0.05),但都比A1和A2地区高(F=5.67,P=0.01),而A1和A2间的密度差别不大(P>0.05),所有五个地区岩羊都没有季节上的数量变化(F=0.64,P=0.60)。
With the increasing of off-road and outdoor ecotourism, wildlife encountered more and more harassment from the human. While we cannot only prohibit human go into protected area simply. On the one hand, we should provide more place to the people, on the other hand, ecotourism can produce money on protection. Wildlife managers and conservationists are looking forward to find a way to make the coexistence of the wildlife and ecotourism. But it was necessary to know how about the flight responses of the wildlife at first.
Different species had different flight responses to anthropogenic disturbance, the different history of contacting with human also had effect on the flight responses. Helan Mountain is the north boundary of distribution area for blue sheep (Pseudois nayaur). the density is much higher than any other areas, and was inhabited in lower elevation, so it is easier to contact in this place. At the same time, researchers in our lab had done works on the group type, home range size, population dynamic, life table, and so on, there are some basic data on blue sheep in Ningxia Helan Mountain National Nature Reserve. That can provide ways to explain the flight responses of the species.
Wildlife in different density encountered different intraspecific competition, which had some effect on the decision in flight responses of wildlife, for the substitute place was an important factor on the trade-off of stay or flee. We analyzed population dynamics of blue sheep in recent years, and then observed flight responses of them. The results can be seen below:
1、We surveyed36line transects in the reserve seasonally from October to December,2010, used Distance6.0to estimate the density of blue sheep, the result was6.08(5.01～7.38)sheep/km2. The density from5.63sheep/km2to6.48sheep/km2in four seasons, but had no significant difference in the season of the reserve (χ2=0.00, df=3, P=1.00). Population size about11,767(9,696～14,282) in the reserve, a little higher than that in2003(Z=1.98, P=0.048). Detection function showed that most of blue sheep could be encountered near than200m.
2^Long-term studies of population dynamics are important for conservation biology and wildlife management. We conducted2,790line transects and observed12,516blue sheep from2004to2009in the Ningxia Helan Mountain National Nature Reserve of China, and found the distribution of blue sheep was uneven in the reserve, then classified the reserve into no distribution, low density area, middle density area and high density area according to the encounter rates of the6years. No distribution area lied in the edge of reserve, the difference in elevation was less between the highest and lowest place and with heavier anthropogenic disturbance; low density area mainly in the north part of the reserve, middle density area mainly in the south part, high density area mainly in the middle part. Rainfall and vegetation cover may influent the density of blue sheep in the reserve.
3、Result of general linear models (GLM:univariate) showed that the encounter rates was mainly decided by the area (F=55.96, P=0.00), not varied among years (F=0.21, P=0.96), and there had no interaction between them. Year also had no effects on the group size (F=0.48, P=0.19) and no interaction with density area (F=0.71, P=0.71), but group size different significant among density areas (F=10.75, P=0.00). Group size in high density area (Mean±SD:8.48±5.55) was bigger than that in middle density area (6.12±5.25) and low density area (4.60±3.64). Group size had significant correlations with encounter rates (linear regression model:Y=0.80+0.36encounter rate, radj2=0.46, F=138.28, P<0.001).
4. Population size of the blue sheep fluctuated from12,375in2004to8,188in2006then13,344in2009, but did not ascend or decline continuously. Population size varied among6years in the reserve (χ2=972.21, df=5, P<0.001), the variation accompanied with rainy days during previous year (r=0.93, P=0.008). So rainy days maybe the cause of variation in blue sheep population size. Ratios between female and male, female and juvenile, adult and juvenile had no difference in each density area among6years, but varied among density areas (P<0.05). Adult female blue sheep was the most animal in the reserve.
5^According to the anthropogenic disturbance, we categorized the reserve into5parts:two on-trail area (A4:Dashuigou on-trail; A5:Suyukou on-trail) and3parts with less human disturbance (Al:Dashuigou off-trail; A2:Maliankou; A3: Suyukou off-trail). We used flight initiation distance (FID)and final distance (FD) to evaluate the flight responses of blue sheep. Used general linear models (univariate) to analyze the effects of area, group size and season on the flight responses. With626groups that encountered from March to December,2010, we found blue sheep had different flight initiation distance among areas (F=0.88, P=0.45), also did in the final distance(F=98.02, P<0.01). In Dashuigou off-trail remained farthest from human, but had the nearest distance in two on-trail areas. Flight initiation distance did not vary among group size (F=2.27, P=0.08), either among seasons (F=0.88, P=0.45), but final distance did (season:F=3.76, P=0.01; group size:F=3.54, P=0.02).
6、Majority of the blue sheep fled slowly when encountered with human in the two on-trail area, but majority fled fast in other three areas. Both the flight initiation distance and final distance had negative correlation with number of tourists, and had positive correlation with distance from anthropogenic disturbance. In the time with heavier anthropogenic disturbance (10:00-15:00h), there are similar encounter rates among5areas (F=0.57, P=0.69). Al and A2(with less anthropogenic disturbance) had lower density of blue sheep than other three areas (F=5.67, P=0.01), but had no difference between them (P>0.05). There had no variation in density among seasons (F=0.64, P=0.60).
引文
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