山地次生林中鸟类巢址的时空结构的研究
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摘要
2005年～2007年的3月到7月,我们在吉林省东部左家自然保护区山地次生林中选取实验样地和对照样地,对鸟类的巢址时空结构进行了分析和研究。以鸟类群落结构研究的一个新视角,揭示山地次生林鸟类巢址的三维时空结构,构建巢址的三维时间和空间结构模式。同时分析了巢址结构在群落结构形成中的作用、巢址时空结构的功能;巢址结构的稳定性。研究结果如下:
一、揭示了山地次生林鸟类巢址的时空结构
结果表明,在山地次生林鸟类群落中,鸟类巢址空间分层分布的现象是比较明显的。综合2005-2007年巢址分析,红脚隼是最高的;环颈雉是最低的;银喉长尾山雀,喜鹊,黄鹂,红隼处于较高的位置;大斑啄木鸟,沼泽山雀,白眉姬鶲,灰头绿啄木鸟,大山雀和北红尾鸲处于较低的位置。普通鳾和麻雀处于中等高度的位置。从时间看,2005-2007年每年每种鸟类巢址空间结构也有不同。
二、分析了山地次生林鸟类巢址结构与植被的关系
从天然巢址看,成功的天然巢树种主要是白皮柳和蒙古栎。从鸟类的种类上看,大山雀,大斑啄木鸟,戴胜,灰椋鸟,普通鳾和银喉长尾山雀这6种鸟的巢树种类中,白皮柳≥50%,而且占统计鸟种类数的50%。80%的红隼选择蒙古栎作为巢树;银喉长尾山雀只选择白皮柳和黑桦两种树种。黄鹂主要选择槺椴，其次是蒙古栎，而且这两种树种占黄鹂巢树总数的83.3%。灰背鸫和沼泽山雀的巢树种类比较分散;灰椋鸟只选择白皮柳和蒙古栎两种树种;100%的喜鹊选择蒙古栎作为它的巢树。黄喉鵐 100%选择地面筑巢。同时,我们还对树龄、巢高、草本植物与巢址结构的关系作了量化分析。
在草本与鸟类巢址的关系分析中,我们发现草本盖度是很好的指示因子。同时对次级洞巢鸟对人工巢箱的利用,包括人工巢箱的使用情况;人工巢箱重复利用的规律;不同年度所使用的人工巢箱高度的变化以及鸟类使用的巢箱总体海拔高度和相对高度分布等等都进行了定量的研究。
三、阐述了山地次生阔叶林鸟类巢址结构中的优势巢址
我们将处于垂直高度在“高”的位置上的鸟类体长均值(37.2)、垂直高度处于“中”和“中低”的均值((19.9+21.3)/2=20.6),以及垂直高度处于“低”的均值(12.8)三个数据进行比较,很显然:37.2 >20.6 >12.8。由此我们发现,鸟类垂直高度的分布与鸟类体长有对应关系。
从群落水平看,L样地(对照样地)的180个鸟巢中,超过10个巢的鸟种分别是:大山雀(32),普通鳾(25),沼泽山雀(18),红隼(14),灰背鸫(13),黄鹂(12)。A样地(实验样地)的118个鸟巢中,超过10个的鸟种分别是:普通鳾(25),喜鹊(19),沼泽山雀(14),大山雀(13),红隼(12),大斑啄木鸟(11)和黄鹂(11)。
虽然对照样地和实验样地的优势巢址略有不同,但是他们有共同的优势巢址。因此次生阔叶林鸟类群落的优势巢址是:大山雀,普通鳾,沼泽山雀,红隼和黄鹂巢址。四、提出了鸟类巢址结构稳定性的理论基础
从系统论、协同论和耗散理论的角度,我们对鸟类群落的巢址结构进行了分析。
从系统论上看,任何鸟类群落都是有一定的鸟类组成,等级序列,功能等。它们具有整体性,关联性,动态平衡性和时序性。比如,从次生洞巢鸟类系统看,构成这个系统的要素是大山雀(3.61%),普通鳾(2.15%)和沼泽山雀(2.06%)。他们各自占有一定的比例,成为次生洞巢鸟类的结构,而且各要素之间即鸟类和鸟类之间都是有联系的。它们彼此共同出现在一定的空间中。无论是在实验样地和对照样地,次级洞巢鸟对天然洞的利用率都是3.61%。这足以说明该次级洞巢鸟群落的动态平衡性。
从耗散理论看,认为鸟类群落的巢址结构也是耗散结构。因为该巢址结构,包括任何处于动态发展中的其他鸟类群落结构,都要吸收养料排出废物,不断进行新陈代谢才能生存。从群落的角度看,一方面群落需要输入食物、足够的阳光、水分和巢材或者新增巢资源,或者引入新种类;另一方面群落要输出子代和排掉废弃物,排除或者竞争排挤掉其他不适种类。群落不断地在变化着,从一种态势到另外一种态势,总是处在保持相对稳定的高度组织化的“暂时”的有序结构。这就是我们所看到的某时刻某林型中所存在的鸟类巢址状态,即巢址结构。我们上述研究结果中的任何一种客观的巢址结构都是如此。同时,我们还分析了洞巢的再利用和人工巢箱的重复利用规律等。
从协同论看,在我们前面的一些研究结果中,例如,不同年份的鸟类巢址立体结构,都可以看做是“功能结构”。这种功能结构是不同垂直高度的鸟类彼此竞争和扑食的一种结果的展示。
总之,我们对山地次生林鸟类巢址结构的形成和稳定性进行了多角度的探讨。这将有助于理解巢洞资源量、生态位重叠、种间竞争,物种多样性等相关问题。同时,对于森林生态系统的平衡、维持生态系统稳定性以及对自然保护区建设、规划和管理都有重要指导意义。
We selected the study and the comparative plots ,in Mountain Zuojia Nature Reserve,a secondary forest,in eastern Jilin Province in March to July from 2005 to 2007,and then we analyzed the bird nest-site structures.Based on the new perspective of bide communities,the study revealed the structure of three-dimensional space-time of the bird nest site and its structure model of constructing the nest site in secondary mountain forestry.We also analyzed the function and stability of nest-site structure. The results are as follows:
First,the study reveals bird nest three-dimensional structure of time and space in the mountain broadleaved forest.
The results show that the phenomenon of layered space of bird nest site in the mountain secondary broad-leaved forest is quite obvious.Through comprehensive analysis of 2005-2007 nest site,Eastern Red-footed Falcon (Falco amurensis) is the highest; Ring-necked Pheasant (Phasianus colchicus karpowi) is the lowest.Long-tailed Tit (Aegithalos caudatus),Common Magpie (Pica pica Sericea),Black-naped Oriole (Oriolus chinensis diffuses) and Common Kestrel (Falco tinnunculus interstinctus) are in a higher position.Great Pied Woodpecker(Picoides major japonicus) , Marsh Tit (Parus palustris brevirostris) ,Yellow-rumped Flycatcher (Ficedula zanthopygia),Grey-headed Woodpecker (Picus canus jessoensis),Great Tit (Parus major minor) and Daurian Redstart (Phoenicurus auroreus auroreus) are at the lower end of the location. Eurasian Nuthatch (Sitta europaea seorsa) and Tree Sparrow (Passer montanus montanus) are common in the middle height position.Each bird nest space distribution of each year from 2005 to 2007 is also different.
Second,the study proposes the relationship between vegetation and birds nest-site structure in the mountain broadleaved forest.
From the look of natural nest sites,the good natural nests are mainly white willow tree (Salix Pierotii Mig ) and Quercus mongolica(Quercus mongolica). From the point of view of bird species,among Great Tit,Great Pied Woodpecker,Eurasian Hoopoe(Upupa epops epops),White-cheeked Starling(Sturnus cineraceus),Eurasian Nuthatch and Long-tailed Tit,
The tree species of them is white willow which is≥50% and accounting for 50% of the number of bird species. The tree species of 80% of Common Kestrel is Quercus;Long-tailed tits select only two kinds of trees, Salix Pierotii Mig and Betula dahurica. Black-naped Oriole would like to choose Tilia mandshurica,followed by Quercus mongolica,which account for 83.3% for the Black-naped Oriole.The nest tree species of Grey-backed Thrush (Turdus hortulorum) and marsh tit scattered;White-cheeked Starling selects only two kinds of trees, Salix Pierotii Mig and Quercus mongolica;100% of Common Magpie select Quercus mongolica as their nest tree. All of Yellow-throated Bunting (Enberiza elegans ticehursti) use ground for their nests. We make quantitative analysis about relationship between the structure of the nest site and the tree age,nest height,herbs .In the analysis of the relationship between bird nests and the grass,we find that herb cover is a good indication factor.Meanwhile we also make quantitative research of the secondary cavity nesters using artificial nest boxes,including the condition of usying artificial nest boxes;reuse rules of artificial nest boxes;the change of artificial nest boxes used at a high degree among different years and the overall elevation and relative height distribution of using nest boxes,and so on.
Third,the study elaborates preponderant nests of birds nest-site structure in the mountain broadleaved forest.
We compared among vertical height in the "high"position,the mean body length of birds (37.2),vertical height in the " middle"and" lower-middle",mean ((19.9 +21.3) / 2 = 20.6),and the vertical height in the "low",average (12.8) ,so that it is clear: 37.2> 20.6> 12.8. Thus we find that there exists a corresponding relationshipbetween the distribution of vertical height of birds and birds body length.
From the view of the community level,there are somebirds who have more than 10 bird nests in 180 nests in L plots (on the usual ground),are: Great Tit (32),Eurasian Nuthatch (25), Marsh Tit (18),Common Kestrel (14), Grey-backed Thrush (13),Black-naped Oriole (12).There are some brids who have more than 10 bird nests in 118 nests in A plot (experimental plots) .they are: Eurasian Nuthatch (25),Common Magpie (19),Marsh Tit (14),Great Tit (13),Common Kestrel (12 ),Great Pied Woodpecker (11) and Black-naped Oriole (11).
Although the advantages of nest-site in the experimental plots and the comparative plots are slightly different,but they still have the same advantages of nest sites. Therefore,bird communities with advantage of nest site in the secondary broad-leaved forest are: Great Tit,Eurasian Nuthatch,Marsh Tit;Common Kestrel and Black-naped Oriole.
Fourth,we analyse the theory of the structural stability of bird nest-site.
From system theory,coordination theory and dissipative theory perspective,we analyzed nest site structure of the bird community.According to system theory,any bird community is composed of certain birds,grade sequence and function. They have integrity,relevance,and timing of dynamic balance.For example,the system of the secondary hole nesting birds,the elements that make up this system are the Great Tit (3.61%),Eurasian Nuthatch (2.15%) and Marsh Tit (2.06%).They respectively occupy a certain percentage in the secondary structure of hole nesting birds,and the elements are that the birds and the birds are all linked each orther. They shared a certain space with each other. Both in the experimental plots and comparative plots,the utilization of natural holes for the secondary cavity nesters is 3.61%.It also illustrates the dynamic balance of the community of the secondary cavity nesters. According to coordination theory,in our previous some research results,for example,three-dimensional structure of bird nests of different years can be seen as "functional structure. "This functional structure is the show of the different vertical height of birds for competition and prey.According to dissipative theory,community structure of bird nest site is also dissipative structure, for the nest-site structure,including any other bird community structure in the dynamic development,must absorb the nutrients , discharge of waste and make continueus metabolism to survive. From the point of view of the community,on the one hand, communities need to get food,adequate sunlight,water and nesting material or nest resources,on the other hand,the community should produce offspring,drain out waste,compete with or drive other inappropriate species.Communities are constantly changing,from one situation to another situation,and always remain in the temporary and ordered structure which is relatively stable and highly organized.As we see,this is the nest site structure,the existence state of bird nest site in a forest,including any above-mentioned objective nest site structure in our results.We also analyzed the re-use law of the nest holes and artificial nest boxes.
In short, we discuss the structure and stability of bird nest on mountain secondary forestry from different angles.This will help to understand the nest resources,niche overlap, interspecific competition,species diversity and other related issues. Meanwhile,these views have important guiding significance to the balance of forest ecosystems, maintain ecosystem stability,construction and management of conservation area.
一、揭示了山地次生林鸟类巢址的时空结构
结果表明,在山地次生林鸟类群落中,鸟类巢址空间分层分布的现象是比较明显的。综合2005-2007年巢址分析,红脚隼是最高的;环颈雉是最低的;银喉长尾山雀,喜鹊,黄鹂,红隼处于较高的位置;大斑啄木鸟,沼泽山雀,白眉姬鶲,灰头绿啄木鸟,大山雀和北红尾鸲处于较低的位置。普通鳾和麻雀处于中等高度的位置。从时间看,2005-2007年每年每种鸟类巢址空间结构也有不同。
二、分析了山地次生林鸟类巢址结构与植被的关系
从天然巢址看,成功的天然巢树种主要是白皮柳和蒙古栎。从鸟类的种类上看,大山雀,大斑啄木鸟,戴胜,灰椋鸟,普通鳾和银喉长尾山雀这6种鸟的巢树种类中,白皮柳≥50%,而且占统计鸟种类数的50%。80%的红隼选择蒙古栎作为巢树;银喉长尾山雀只选择白皮柳和黑桦两种树种。黄鹂主要选择槺椴，其次是蒙古栎，而且这两种树种占黄鹂巢树总数的83.3%。灰背鸫和沼泽山雀的巢树种类比较分散;灰椋鸟只选择白皮柳和蒙古栎两种树种;100%的喜鹊选择蒙古栎作为它的巢树。黄喉鵐 100%选择地面筑巢。同时,我们还对树龄、巢高、草本植物与巢址结构的关系作了量化分析。
在草本与鸟类巢址的关系分析中,我们发现草本盖度是很好的指示因子。同时对次级洞巢鸟对人工巢箱的利用,包括人工巢箱的使用情况;人工巢箱重复利用的规律;不同年度所使用的人工巢箱高度的变化以及鸟类使用的巢箱总体海拔高度和相对高度分布等等都进行了定量的研究。
三、阐述了山地次生阔叶林鸟类巢址结构中的优势巢址
我们将处于垂直高度在“高”的位置上的鸟类体长均值(37.2)、垂直高度处于“中”和“中低”的均值((19.9+21.3)/2=20.6),以及垂直高度处于“低”的均值(12.8)三个数据进行比较,很显然:37.2 >20.6 >12.8。由此我们发现,鸟类垂直高度的分布与鸟类体长有对应关系。
从群落水平看,L样地(对照样地)的180个鸟巢中,超过10个巢的鸟种分别是:大山雀(32),普通鳾(25),沼泽山雀(18),红隼(14),灰背鸫(13),黄鹂(12)。A样地(实验样地)的118个鸟巢中,超过10个的鸟种分别是:普通鳾(25),喜鹊(19),沼泽山雀(14),大山雀(13),红隼(12),大斑啄木鸟(11)和黄鹂(11)。
虽然对照样地和实验样地的优势巢址略有不同,但是他们有共同的优势巢址。因此次生阔叶林鸟类群落的优势巢址是:大山雀,普通鳾,沼泽山雀,红隼和黄鹂巢址。四、提出了鸟类巢址结构稳定性的理论基础
从系统论、协同论和耗散理论的角度,我们对鸟类群落的巢址结构进行了分析。
从系统论上看,任何鸟类群落都是有一定的鸟类组成,等级序列,功能等。它们具有整体性,关联性,动态平衡性和时序性。比如,从次生洞巢鸟类系统看,构成这个系统的要素是大山雀(3.61%),普通鳾(2.15%)和沼泽山雀(2.06%)。他们各自占有一定的比例,成为次生洞巢鸟类的结构,而且各要素之间即鸟类和鸟类之间都是有联系的。它们彼此共同出现在一定的空间中。无论是在实验样地和对照样地,次级洞巢鸟对天然洞的利用率都是3.61%。这足以说明该次级洞巢鸟群落的动态平衡性。
从耗散理论看,认为鸟类群落的巢址结构也是耗散结构。因为该巢址结构,包括任何处于动态发展中的其他鸟类群落结构,都要吸收养料排出废物,不断进行新陈代谢才能生存。从群落的角度看,一方面群落需要输入食物、足够的阳光、水分和巢材或者新增巢资源,或者引入新种类;另一方面群落要输出子代和排掉废弃物,排除或者竞争排挤掉其他不适种类。群落不断地在变化着,从一种态势到另外一种态势,总是处在保持相对稳定的高度组织化的“暂时”的有序结构。这就是我们所看到的某时刻某林型中所存在的鸟类巢址状态,即巢址结构。我们上述研究结果中的任何一种客观的巢址结构都是如此。同时,我们还分析了洞巢的再利用和人工巢箱的重复利用规律等。
从协同论看,在我们前面的一些研究结果中,例如,不同年份的鸟类巢址立体结构,都可以看做是“功能结构”。这种功能结构是不同垂直高度的鸟类彼此竞争和扑食的一种结果的展示。
总之,我们对山地次生林鸟类巢址结构的形成和稳定性进行了多角度的探讨。这将有助于理解巢洞资源量、生态位重叠、种间竞争,物种多样性等相关问题。同时,对于森林生态系统的平衡、维持生态系统稳定性以及对自然保护区建设、规划和管理都有重要指导意义。
We selected the study and the comparative plots ,in Mountain Zuojia Nature Reserve,a secondary forest,in eastern Jilin Province in March to July from 2005 to 2007,and then we analyzed the bird nest-site structures.Based on the new perspective of bide communities,the study revealed the structure of three-dimensional space-time of the bird nest site and its structure model of constructing the nest site in secondary mountain forestry.We also analyzed the function and stability of nest-site structure. The results are as follows:
First,the study reveals bird nest three-dimensional structure of time and space in the mountain broadleaved forest.
The results show that the phenomenon of layered space of bird nest site in the mountain secondary broad-leaved forest is quite obvious.Through comprehensive analysis of 2005-2007 nest site,Eastern Red-footed Falcon (Falco amurensis) is the highest; Ring-necked Pheasant (Phasianus colchicus karpowi) is the lowest.Long-tailed Tit (Aegithalos caudatus),Common Magpie (Pica pica Sericea),Black-naped Oriole (Oriolus chinensis diffuses) and Common Kestrel (Falco tinnunculus interstinctus) are in a higher position.Great Pied Woodpecker(Picoides major japonicus) , Marsh Tit (Parus palustris brevirostris) ,Yellow-rumped Flycatcher (Ficedula zanthopygia),Grey-headed Woodpecker (Picus canus jessoensis),Great Tit (Parus major minor) and Daurian Redstart (Phoenicurus auroreus auroreus) are at the lower end of the location. Eurasian Nuthatch (Sitta europaea seorsa) and Tree Sparrow (Passer montanus montanus) are common in the middle height position.Each bird nest space distribution of each year from 2005 to 2007 is also different.
Second,the study proposes the relationship between vegetation and birds nest-site structure in the mountain broadleaved forest.
From the look of natural nest sites,the good natural nests are mainly white willow tree (Salix Pierotii Mig ) and Quercus mongolica(Quercus mongolica). From the point of view of bird species,among Great Tit,Great Pied Woodpecker,Eurasian Hoopoe(Upupa epops epops),White-cheeked Starling(Sturnus cineraceus),Eurasian Nuthatch and Long-tailed Tit,
The tree species of them is white willow which is≥50% and accounting for 50% of the number of bird species. The tree species of 80% of Common Kestrel is Quercus;Long-tailed tits select only two kinds of trees, Salix Pierotii Mig and Betula dahurica. Black-naped Oriole would like to choose Tilia mandshurica,followed by Quercus mongolica,which account for 83.3% for the Black-naped Oriole.The nest tree species of Grey-backed Thrush (Turdus hortulorum) and marsh tit scattered;White-cheeked Starling selects only two kinds of trees, Salix Pierotii Mig and Quercus mongolica;100% of Common Magpie select Quercus mongolica as their nest tree. All of Yellow-throated Bunting (Enberiza elegans ticehursti) use ground for their nests. We make quantitative analysis about relationship between the structure of the nest site and the tree age,nest height,herbs .In the analysis of the relationship between bird nests and the grass,we find that herb cover is a good indication factor.Meanwhile we also make quantitative research of the secondary cavity nesters using artificial nest boxes,including the condition of usying artificial nest boxes;reuse rules of artificial nest boxes;the change of artificial nest boxes used at a high degree among different years and the overall elevation and relative height distribution of using nest boxes,and so on.
Third,the study elaborates preponderant nests of birds nest-site structure in the mountain broadleaved forest.
We compared among vertical height in the "high"position,the mean body length of birds (37.2),vertical height in the " middle"and" lower-middle",mean ((19.9 +21.3) / 2 = 20.6),and the vertical height in the "low",average (12.8) ,so that it is clear: 37.2> 20.6> 12.8. Thus we find that there exists a corresponding relationshipbetween the distribution of vertical height of birds and birds body length.
From the view of the community level,there are somebirds who have more than 10 bird nests in 180 nests in L plots (on the usual ground),are: Great Tit (32),Eurasian Nuthatch (25), Marsh Tit (18),Common Kestrel (14), Grey-backed Thrush (13),Black-naped Oriole (12).There are some brids who have more than 10 bird nests in 118 nests in A plot (experimental plots) .they are: Eurasian Nuthatch (25),Common Magpie (19),Marsh Tit (14),Great Tit (13),Common Kestrel (12 ),Great Pied Woodpecker (11) and Black-naped Oriole (11).
Although the advantages of nest-site in the experimental plots and the comparative plots are slightly different,but they still have the same advantages of nest sites. Therefore,bird communities with advantage of nest site in the secondary broad-leaved forest are: Great Tit,Eurasian Nuthatch,Marsh Tit;Common Kestrel and Black-naped Oriole.
Fourth,we analyse the theory of the structural stability of bird nest-site.
From system theory,coordination theory and dissipative theory perspective,we analyzed nest site structure of the bird community.According to system theory,any bird community is composed of certain birds,grade sequence and function. They have integrity,relevance,and timing of dynamic balance.For example,the system of the secondary hole nesting birds,the elements that make up this system are the Great Tit (3.61%),Eurasian Nuthatch (2.15%) and Marsh Tit (2.06%).They respectively occupy a certain percentage in the secondary structure of hole nesting birds,and the elements are that the birds and the birds are all linked each orther. They shared a certain space with each other. Both in the experimental plots and comparative plots,the utilization of natural holes for the secondary cavity nesters is 3.61%.It also illustrates the dynamic balance of the community of the secondary cavity nesters. According to coordination theory,in our previous some research results,for example,three-dimensional structure of bird nests of different years can be seen as "functional structure. "This functional structure is the show of the different vertical height of birds for competition and prey.According to dissipative theory,community structure of bird nest site is also dissipative structure, for the nest-site structure,including any other bird community structure in the dynamic development,must absorb the nutrients , discharge of waste and make continueus metabolism to survive. From the point of view of the community,on the one hand, communities need to get food,adequate sunlight,water and nesting material or nest resources,on the other hand,the community should produce offspring,drain out waste,compete with or drive other inappropriate species.Communities are constantly changing,from one situation to another situation,and always remain in the temporary and ordered structure which is relatively stable and highly organized.As we see,this is the nest site structure,the existence state of bird nest site in a forest,including any above-mentioned objective nest site structure in our results.We also analyzed the re-use law of the nest holes and artificial nest boxes.
In short, we discuss the structure and stability of bird nest on mountain secondary forestry from different angles.This will help to understand the nest resources,niche overlap, interspecific competition,species diversity and other related issues. Meanwhile,these views have important guiding significance to the balance of forest ecosystems, maintain ecosystem stability,construction and management of conservation area.
引文
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