秦岭大熊猫主要栖息地植物群落特征及与生境对应关系分析
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摘要
佛坪、长青国家自然保护区是秦岭大熊猫主要栖息地，由于独特的地理位置和生态环境，孕育了种类独特、丰富多样的生物资源和植物群落。然而，目前对整个栖息地植被的研究仍然停留在具体的某一个群落类型或者具体地段，缺乏对整个栖息地区系组成、群落类型结构以及生长状况全面系统的研究和认识。为了揭示栖息地植被分布规律以及与生境因子之间的关系，本研究在上述两个国家自然保护区海拔900-3071m范围内各设置一条样线，每条样线上布设40-60个标准样地，共计107个，对样地内植物群落物种组成、生长状态、分布环境、土壤等方面进行全面系统调查。在分析样地调查数据和土壤分析数据的基础上，分别从植物区系组成结构、区系海拔格局、植物群落物种多样性、群落复杂性、物种多样性与复杂性的海拔格局及其关系、群落稳定性评价等7个方面进行系统、深入研究，以期揭示秦岭大熊猫栖息地植物群落分布特征及其与环境的关系，从而为栖息地物种多样性保护与可持续利用、植被恢复与重建、保护和维持大熊猫活动领域等方面提供理论依据。其主要研究结果如下：
（1）秦岭大熊猫栖息地种子植物共有2031种，分别隶属于148科，694属，占整个秦岭种子植物总数59.11%，占全国种子植物总数的7.45%，该区植物种类繁多，区系成分极为丰富。其中，中国特有种36种，珍稀保护物种15种，在植物区系成分中，原始种类、孑遗成分较多，尤其在中国特有种和国家保护物种中古老成分所占比例较大，体现出植物区系的古老性。整个研究区植物区系以温带成分为主，并且具有亚热带向温带过渡的特点，是亚热带和温带植物区系的重要交汇地区。
（2）对秦岭大熊猫栖息地种子植物区系海拔格局分析表明，科、属、种数量沿着海拔升高呈偏正态分布，海拔1200-1300m范围是科、属、种最丰富的区域。利用热带成分比重与温带成分比重的比值，判断其区系平衡点所处区域。通过分析发现，该研究区理论上不存在植被亚热带-暖温带分界线（热带和温带成分区系平衡点），其理论分界线应该在海拔530m区域。海拔2800m是热带成分分布临界高度。
（3）分别采用双向指示种分析（TWINSPAN）、除趋势对应分析（DCA）和除趋势典范对应分析（DCCA）对秦岭南坡大熊猫栖息地群落进行分类与排序，结果表明TWINSPAN双向聚类分析能较好地将不同的群落类型截然分开，研究区107个群落样地被分成太白红杉（Larix chinensis）、巴山冷杉（Abies fargesii）、红桦（Betulaalbo-sinensis）、牛皮桦（Betula albo-sinensis var. septentrionalis）等44个群落（群系）类型；利用DCA对107个样地进行排序，结果显示不同类型样地呈现聚集分布，其结果充分验证了TWINSPAN对群落类型划分的科学性。除此之外，通过对107个样地除趋势典范对应分析（DCCA）和Monte Carlo显著性检验发现，在研究涉及的12个环境因子中制约秦岭南坡大熊猫栖息地群落类型、植物种分布格局的主要因素是海拔、坡度、土壤有效磷与全磷。
（4）不同植被型、群落类型以及各个层次的多样性存在明显差异。就植被型而言，常绿和落叶混交林的物种多样性最高，竹林最低；而对于群落而言，椆（Cyclobalanopsisglauca）+锐齿槲栎（Quercus aliena var. acuteserrata）混交林物种多样性最高，巴山木竹林（Bashania fargesii）最低。植被型和群落各个层次间物种多样性的高低没有一致顺序，这主要与环境因子以及灌木层密度影响有关。在群落各层物种多样性相互关系中，乔木层和灌木层存在显著相关性，其它层之间没有显著关系。深入分析12个环境因子与物种多样性关系发现，土壤、地形、物种多样性三者存在相互制约关系，其中海拔、土壤全钾和有效磷分别与物种多样性存在着显著相关关系。
（5）利用Huffman编码对不同群落类型以及群落内各层次进行复杂性评价发现，不同群落类型复杂性存在明显差异，并且各个层次间也存在明显差异，乔木层总复杂性、无序结构复杂性与灌木层有显著相关性。进一步分析群落复杂性与环境因子、群落优势度、变异系数之间的关系发现，土壤因子和地形因子与群落结构复杂性有显著相关性，尤其是海拔、坡向、有效磷、全磷对群落结构复杂性有着显著影响；而群落优势度以及变异系数均与群落复杂性呈显著线性关系，变异系数越大，群落有序结构复杂性越高，无序结构复杂性越低；群落优势度越大，群落复杂性越小。
（6）随着海拔升高，群落结构总复杂性表现出与物种多样性极为相似的变化规律，呈现出“低—高—低”的偏正态分布格局，其中海拔1100-1300m范围是植物群落总复杂性和物种多样性最大区域。
植物群落无序结构复杂性与总复杂性沿海拔梯度变化呈现相似的变化规律，即“低—高—低”分布格局，但是群落有序结构复杂性随着海拔升高无明显的变化规律。该区域植物群落物种多样性不仅与海拔、气温、土壤等外界条件变化有关，而且还与竹林分布密度有着密切关系。在2个竹林交汇区域物种多样性及物种数量有着明显提高。相关分析表明，群落总复杂性、无序复杂性和物种丰富度、Shannon-Weiner指数、Simpson指数、均匀度指数均有着极其显著的线性关系。其中，群落总复杂性与Shannon-weiner相关性接近于1，所以物种多样性能够完全表达植物群落总复杂性变化规律。但是有序结构复杂性与物种均匀程度没有显著关系，植物群落有序结构复杂性与物种多样性之间有着明显差异，因而物种多样性不能够完全作为作植物群落有序结构复杂性的测度。
（7）在该研究区选取分布最广、最具有代表性的16个群落类型，并利用模糊数学隶属函数对其优势种更新、林地质量、群落物种多样性、群落复杂性、土壤肥力、保护程度6个方面进行综合评价得到，群落稳定性由大到小顺序为：铁杉（Tsuga chinensis）>短柄枹栎（Quercus glandulifera var.brevipetiolata）>太白杨（Populus purdomii）>锐齿槲栎>华山松（Pinus armandii）>白桦（Betula platyphylla）>巴山冷杉>油松（Pinustabulaeformi）+华山松>红桦>牛皮桦>太白红杉>油松+锐齿槲栎>油松>栓皮栎（Quercusvariabilis）>毛栗（Castanea mollissima）>亮叶桦（Betula luminifera）。
Foping Nature Reserve and Changqing Nature Reserve are the main habitats of the giantpanda in the middle range of the Qinling Mountains. The geographical position and ecologicalforest environment of this region create uniquely and richly biological resources and plantcommunities. Previous studies focused on the study of plant community traits and speciesdiversity in this habitat area, but the flora, community types, and growth status are stillunknown, therefore, the protection and operation of the giant panda habitat exists someblindness.
Plant community character, distribution and the relationship with environment on thehabitat of Giant Panda in the Qinling Mountains were explored in this study. Foping NatureReserve and Changqing Nature Reserve were selected as study areas. Tow line transects wereset between the altitude of900-3071m in the field, and107standard plots (25×20m2)wereinvestigated on transects. Vegetation community, growth status and distribution environmentwere recorded and soil samples were collected in the standard plots. Based on analysis ofplots and soil data, flora characteristics, altitudinal patterns of seed plants, plant speciesdiversity, community complexity, the altitudinal pattern between species diversity andcommunity complexity and community stability evaluation were examined comprehensively.
Our study was trying to explore the plant community distribution and its relationshipwith the environment in the Qinling panda habitat, so as to provide some scientific referencesand theoretical basis for habitat conservation and sustainable utilization of species diversity,vegetation restoration and reconstruction, and the artificial expansion of the giant pandaactivity areas.
The results are as follows:
(1) The habitat contained2031spermatophyte species from148families and694genera.Spermatophyte species accounted for59.11%of the total plant species in the QinlingMountains but only7.45%of the total plant species in China, including36endemic Chinese plant species and15protected species. There were many ancient and relict spermatophytespecies in this region, which represented a large proportion of endemic Chinese species andthose in a nationally conserved status. Tertiary relict plant species are rich in abundance andthey form an important part of plant communities in the research area, which indicates that theorigin of the flora is very ancient. Temperate genera are dominant in the area, but there is atransition from subtropical to temperate types. Thus, the research area is an importantintersection region. At the same time, the research area is also the intersection area betweenSino-Japan forest zone and Sino-Himalaya forest zone.
(2) Analyzing of the altitudinal phytogeography of spermatophytes (seed plants) inQingling Mountains habbitat, the quantity of family, genus and species showed the skewnormal distribution with the increased altitude. The richest family, genus and species wereconcentrated at1200-1300m. According to the ratio of the the tropical and temperate elementsproportion, the balance point of the research area was found out. There was no vegetationsubtropical-warm temperate dividing line(floristic equilibrium point) in study area fromtheory, and the theoretical boundary for the subtropical-warm temperate zone should be at530m above the sea level, and the threshold height of tropical distribution is at2800m.
(3)107standard plots were classified into44plant communities by Two-way IndicatorsSpecies Analysis (TWINSPAN). The result of ordering by Detrended Corresponding Analysis(DCA) showed that the standard plots fit to aggregation distribution and verified thescientificity of TWINSPAN. The test of significance between Detrended CanonicalCorrespondence Analysis (DCCA) and Monte Carlo that the restrictive factors of communitytype and species distribution were altitude, slope, rapidly-available phosphorus and totalphosphorus at the landscape level.
(4) Different vegetation types and community types had different species diversity anddiversity of different layers had significant differences. For vegetation types, the highestdiversity was in evergreen forests and mixed deciduous forest, the lowest one was in bambooforest. For community types, the highest diversity was in Cyclobalanopsis glauca+Q. alienavar. Acuteserrata mixed forest, the lowest was in Bashania fargesii bamboo forest. Differentlayers of vegetation types and community types did not have the same sequence, because ofthe diversities of different layers had a close relationship with environment and shrub density.The diversity of tree layer was significantly correlated with the shrub layer diversity, butwasn’t related with the herb layer. Careful analysis of the relationships between speciesdiversity and environment factors explored there existed mutual restriction relationshipbetween soil, topography and species diversity. Altitude, available phosphorus and totalpotassium had significant correlation with diversity respectively.
(5) The community complexity was evaluated by Huffman Coding. The complexity ofdifferent community types and layers was significantly different. The total complexity anddisorder complexity was significantly correlated with that in shrub layer. Soil and topographyfactors had significant correlation with community structure complexity, especially altitude,slope, available phosphorus, total phosphorus. Community dominance and coefficient ofvariation had the significant linear relationship with community complexity respectively.Community ordered structure complexity was higher, disordered structure complexity waslower; community dominance was higher, and community complexity was lower.
(6) With increased altitude, the species diversity and complexity of plant communitiespresented a "low-high-low" pattern of skewed normal distribution. The altitude between1300and1500m was the largest region of total complexity and diversity of plant community.Community disordered structure complexity was similar to community complexity. However,Community ordered structure complexity did not have regular changes along with the altitude.Plant species diversity of the natural reserve was not only related with the altitude change, butalso had close relationship with the density of Bashania fargesii and Fargesiaqinlingensis.The species diversity and species number had improved significantly in theintersection of two bamboo groves. Correlation analysis proved community complexity andcommunity disordered structure complexity had significant linear relationship with diversityindex respectively. The correlation value between community complexity andShannon-Weiner was close to1, it means the species diversity would express the change ofcommunity complexity. Community ordered structure complexity was not correlated withevenness index, which means it could not be used as the measure method of communitycomplexity.
(7) Community stability of16reprehensive and typical community types were evaluatedby subordinate function of fuzzy mathematics. The evaluation index included the regenerationof dominant species, stand qualities, diversity, community complexity, soil fertility andprotective degree. The stability order of different communities was: Tsuga chinensis>Quercus glandulifera var.brevipetiolata> Populus purdomii> Quercus aliena var.acuteserrata> Pinus armandii> Betula platyphylla> Abies fargesii> Pinus tabulaeformi+Pinus armandii>Betula albo-sinensis> Betula albo-sinensis var. septentrionalis> Larixchinensis> Pinus tabulaeformi+Quercus aliena var. acuteserrata> Pinus tabulaeformi>Quercus variabilis> Castanea mollissima> Betula luminifera.
（1）秦岭大熊猫栖息地种子植物共有2031种，分别隶属于148科，694属，占整个秦岭种子植物总数59.11%，占全国种子植物总数的7.45%，该区植物种类繁多，区系成分极为丰富。其中，中国特有种36种，珍稀保护物种15种，在植物区系成分中，原始种类、孑遗成分较多，尤其在中国特有种和国家保护物种中古老成分所占比例较大，体现出植物区系的古老性。整个研究区植物区系以温带成分为主，并且具有亚热带向温带过渡的特点，是亚热带和温带植物区系的重要交汇地区。
（2）对秦岭大熊猫栖息地种子植物区系海拔格局分析表明，科、属、种数量沿着海拔升高呈偏正态分布，海拔1200-1300m范围是科、属、种最丰富的区域。利用热带成分比重与温带成分比重的比值，判断其区系平衡点所处区域。通过分析发现，该研究区理论上不存在植被亚热带-暖温带分界线（热带和温带成分区系平衡点），其理论分界线应该在海拔530m区域。海拔2800m是热带成分分布临界高度。
（3）分别采用双向指示种分析（TWINSPAN）、除趋势对应分析（DCA）和除趋势典范对应分析（DCCA）对秦岭南坡大熊猫栖息地群落进行分类与排序，结果表明TWINSPAN双向聚类分析能较好地将不同的群落类型截然分开，研究区107个群落样地被分成太白红杉（Larix chinensis）、巴山冷杉（Abies fargesii）、红桦（Betulaalbo-sinensis）、牛皮桦（Betula albo-sinensis var. septentrionalis）等44个群落（群系）类型；利用DCA对107个样地进行排序，结果显示不同类型样地呈现聚集分布，其结果充分验证了TWINSPAN对群落类型划分的科学性。除此之外，通过对107个样地除趋势典范对应分析（DCCA）和Monte Carlo显著性检验发现，在研究涉及的12个环境因子中制约秦岭南坡大熊猫栖息地群落类型、植物种分布格局的主要因素是海拔、坡度、土壤有效磷与全磷。
（4）不同植被型、群落类型以及各个层次的多样性存在明显差异。就植被型而言，常绿和落叶混交林的物种多样性最高，竹林最低；而对于群落而言，椆（Cyclobalanopsisglauca）+锐齿槲栎（Quercus aliena var. acuteserrata）混交林物种多样性最高，巴山木竹林（Bashania fargesii）最低。植被型和群落各个层次间物种多样性的高低没有一致顺序，这主要与环境因子以及灌木层密度影响有关。在群落各层物种多样性相互关系中，乔木层和灌木层存在显著相关性，其它层之间没有显著关系。深入分析12个环境因子与物种多样性关系发现，土壤、地形、物种多样性三者存在相互制约关系，其中海拔、土壤全钾和有效磷分别与物种多样性存在着显著相关关系。
（5）利用Huffman编码对不同群落类型以及群落内各层次进行复杂性评价发现，不同群落类型复杂性存在明显差异，并且各个层次间也存在明显差异，乔木层总复杂性、无序结构复杂性与灌木层有显著相关性。进一步分析群落复杂性与环境因子、群落优势度、变异系数之间的关系发现，土壤因子和地形因子与群落结构复杂性有显著相关性，尤其是海拔、坡向、有效磷、全磷对群落结构复杂性有着显著影响；而群落优势度以及变异系数均与群落复杂性呈显著线性关系，变异系数越大，群落有序结构复杂性越高，无序结构复杂性越低；群落优势度越大，群落复杂性越小。
（6）随着海拔升高，群落结构总复杂性表现出与物种多样性极为相似的变化规律，呈现出“低—高—低”的偏正态分布格局，其中海拔1100-1300m范围是植物群落总复杂性和物种多样性最大区域。
植物群落无序结构复杂性与总复杂性沿海拔梯度变化呈现相似的变化规律，即“低—高—低”分布格局，但是群落有序结构复杂性随着海拔升高无明显的变化规律。该区域植物群落物种多样性不仅与海拔、气温、土壤等外界条件变化有关，而且还与竹林分布密度有着密切关系。在2个竹林交汇区域物种多样性及物种数量有着明显提高。相关分析表明，群落总复杂性、无序复杂性和物种丰富度、Shannon-Weiner指数、Simpson指数、均匀度指数均有着极其显著的线性关系。其中，群落总复杂性与Shannon-weiner相关性接近于1，所以物种多样性能够完全表达植物群落总复杂性变化规律。但是有序结构复杂性与物种均匀程度没有显著关系，植物群落有序结构复杂性与物种多样性之间有着明显差异，因而物种多样性不能够完全作为作植物群落有序结构复杂性的测度。
（7）在该研究区选取分布最广、最具有代表性的16个群落类型，并利用模糊数学隶属函数对其优势种更新、林地质量、群落物种多样性、群落复杂性、土壤肥力、保护程度6个方面进行综合评价得到，群落稳定性由大到小顺序为：铁杉（Tsuga chinensis）>短柄枹栎（Quercus glandulifera var.brevipetiolata）>太白杨（Populus purdomii）>锐齿槲栎>华山松（Pinus armandii）>白桦（Betula platyphylla）>巴山冷杉>油松（Pinustabulaeformi）+华山松>红桦>牛皮桦>太白红杉>油松+锐齿槲栎>油松>栓皮栎（Quercusvariabilis）>毛栗（Castanea mollissima）>亮叶桦（Betula luminifera）。
Foping Nature Reserve and Changqing Nature Reserve are the main habitats of the giantpanda in the middle range of the Qinling Mountains. The geographical position and ecologicalforest environment of this region create uniquely and richly biological resources and plantcommunities. Previous studies focused on the study of plant community traits and speciesdiversity in this habitat area, but the flora, community types, and growth status are stillunknown, therefore, the protection and operation of the giant panda habitat exists someblindness.
Plant community character, distribution and the relationship with environment on thehabitat of Giant Panda in the Qinling Mountains were explored in this study. Foping NatureReserve and Changqing Nature Reserve were selected as study areas. Tow line transects wereset between the altitude of900-3071m in the field, and107standard plots (25×20m2)wereinvestigated on transects. Vegetation community, growth status and distribution environmentwere recorded and soil samples were collected in the standard plots. Based on analysis ofplots and soil data, flora characteristics, altitudinal patterns of seed plants, plant speciesdiversity, community complexity, the altitudinal pattern between species diversity andcommunity complexity and community stability evaluation were examined comprehensively.
Our study was trying to explore the plant community distribution and its relationshipwith the environment in the Qinling panda habitat, so as to provide some scientific referencesand theoretical basis for habitat conservation and sustainable utilization of species diversity,vegetation restoration and reconstruction, and the artificial expansion of the giant pandaactivity areas.
The results are as follows:
(1) The habitat contained2031spermatophyte species from148families and694genera.Spermatophyte species accounted for59.11%of the total plant species in the QinlingMountains but only7.45%of the total plant species in China, including36endemic Chinese plant species and15protected species. There were many ancient and relict spermatophytespecies in this region, which represented a large proportion of endemic Chinese species andthose in a nationally conserved status. Tertiary relict plant species are rich in abundance andthey form an important part of plant communities in the research area, which indicates that theorigin of the flora is very ancient. Temperate genera are dominant in the area, but there is atransition from subtropical to temperate types. Thus, the research area is an importantintersection region. At the same time, the research area is also the intersection area betweenSino-Japan forest zone and Sino-Himalaya forest zone.
(2) Analyzing of the altitudinal phytogeography of spermatophytes (seed plants) inQingling Mountains habbitat, the quantity of family, genus and species showed the skewnormal distribution with the increased altitude. The richest family, genus and species wereconcentrated at1200-1300m. According to the ratio of the the tropical and temperate elementsproportion, the balance point of the research area was found out. There was no vegetationsubtropical-warm temperate dividing line(floristic equilibrium point) in study area fromtheory, and the theoretical boundary for the subtropical-warm temperate zone should be at530m above the sea level, and the threshold height of tropical distribution is at2800m.
(3)107standard plots were classified into44plant communities by Two-way IndicatorsSpecies Analysis (TWINSPAN). The result of ordering by Detrended Corresponding Analysis(DCA) showed that the standard plots fit to aggregation distribution and verified thescientificity of TWINSPAN. The test of significance between Detrended CanonicalCorrespondence Analysis (DCCA) and Monte Carlo that the restrictive factors of communitytype and species distribution were altitude, slope, rapidly-available phosphorus and totalphosphorus at the landscape level.
(4) Different vegetation types and community types had different species diversity anddiversity of different layers had significant differences. For vegetation types, the highestdiversity was in evergreen forests and mixed deciduous forest, the lowest one was in bambooforest. For community types, the highest diversity was in Cyclobalanopsis glauca+Q. alienavar. Acuteserrata mixed forest, the lowest was in Bashania fargesii bamboo forest. Differentlayers of vegetation types and community types did not have the same sequence, because ofthe diversities of different layers had a close relationship with environment and shrub density.The diversity of tree layer was significantly correlated with the shrub layer diversity, butwasn’t related with the herb layer. Careful analysis of the relationships between speciesdiversity and environment factors explored there existed mutual restriction relationshipbetween soil, topography and species diversity. Altitude, available phosphorus and totalpotassium had significant correlation with diversity respectively.
(5) The community complexity was evaluated by Huffman Coding. The complexity ofdifferent community types and layers was significantly different. The total complexity anddisorder complexity was significantly correlated with that in shrub layer. Soil and topographyfactors had significant correlation with community structure complexity, especially altitude,slope, available phosphorus, total phosphorus. Community dominance and coefficient ofvariation had the significant linear relationship with community complexity respectively.Community ordered structure complexity was higher, disordered structure complexity waslower; community dominance was higher, and community complexity was lower.
(6) With increased altitude, the species diversity and complexity of plant communitiespresented a "low-high-low" pattern of skewed normal distribution. The altitude between1300and1500m was the largest region of total complexity and diversity of plant community.Community disordered structure complexity was similar to community complexity. However,Community ordered structure complexity did not have regular changes along with the altitude.Plant species diversity of the natural reserve was not only related with the altitude change, butalso had close relationship with the density of Bashania fargesii and Fargesiaqinlingensis.The species diversity and species number had improved significantly in theintersection of two bamboo groves. Correlation analysis proved community complexity andcommunity disordered structure complexity had significant linear relationship with diversityindex respectively. The correlation value between community complexity andShannon-Weiner was close to1, it means the species diversity would express the change ofcommunity complexity. Community ordered structure complexity was not correlated withevenness index, which means it could not be used as the measure method of communitycomplexity.
(7) Community stability of16reprehensive and typical community types were evaluatedby subordinate function of fuzzy mathematics. The evaluation index included the regenerationof dominant species, stand qualities, diversity, community complexity, soil fertility andprotective degree. The stability order of different communities was: Tsuga chinensis>Quercus glandulifera var.brevipetiolata> Populus purdomii> Quercus aliena var.acuteserrata> Pinus armandii> Betula platyphylla> Abies fargesii> Pinus tabulaeformi+Pinus armandii>Betula albo-sinensis> Betula albo-sinensis var. septentrionalis> Larixchinensis> Pinus tabulaeformi+Quercus aliena var. acuteserrata> Pinus tabulaeformi>Quercus variabilis> Castanea mollissima> Betula luminifera.
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