三峡库区森林植物多样性研究
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摘要
以132块临时样地、146块固定样地的调查数据和1999-2003年三峡库区森林资源二类调查数据为基础,以三峡库区森林植物多样性研究为主线,应用植被数量分析方法,并将定性和定量方法相结合,从种子植物区系特征、植物群落数量分类与排序及其环境解释、群落物种多样性及环境梯度特征、不同演替阶段森林土壤养分特征、典型针叶林的结构以及林下植物物种的生态位及种间联结特征等方面,比较系统地研究了三峡库区森林植物多样性特征,并分析了库区珍稀濒危物种的现状及其保护对策,对指导三峡库区的生物多样性保护、森林恢复与可持续经营具有重要的理论和现实意义。
根据吴征镒先生对中国种子植物分布类型的划分,该区种子植物属的分布类型可划分为15个类型及其变型,以热带性质的属分布较多,温带性质的属次之,具有较为明显的过渡性。同时,该区域植物区系起源古老,植物种类丰富多样,共有种子植物5688种,占全国种子植物总数的21%。其中珍稀濒危植物56种,分属34科,48属,约占全国总数的10%。三峡库区珍稀濒危植物具有过渡性、分散性和特有性等特征,而且受人为干扰严重。
TWINSPAN数量分类结果表明三峡库区森林植物群落可化分为巴山松(Pinus henryi)林、华山松(P. armandii)林、马尾松(P. massoniana)林、杉木(Cunnighamia lanceolata)林、柏木(Cupessus funebris)林、马尾松+栓皮栎(Pinus massoniana+Quercus variabilis)林、马尾松+栲(Pinus massoniana +Castanopsis fargesii)林、马尾松+大头茶(Pinus massoniana+ Grdonia axillaris)林、杉木+栲(Cunnighamia lanceolata+Castamopsis fargesii)林、短柄枹栎（Quercus glandulifera var. brevipetiolata)林、栓皮栎(Q. variabilis)林、白栎(Q. albus)林、槲栎+栓皮栎(Q. aliena+Q. variabilis)林、西南槲树(Q. denlata var. oxyloba)林、亮叶水青冈(Fagus lucida)林、茅栗(Castanea seguinii)林、化香(Platycarya strobilacea)林、红桦(Betaula albo-sinensis)林、糙皮桦(Betula utilis)林、四照花(Dendrobenthamia japonica var. chinensis)林、枫香(Liquidambar formosana)林、华中樱桃+刺叶栎(Cerasus conrodinae+Quercus spinosa)林、连香树+细齿稠李(Cercidiphyllum japonicum+Padus obtusata)林、灯台树(Cornus controversa)林、朴树(Celtis sinensis)林、包槲柯+锥栗(Lithocarpus cleistocarpus+Castanea henryi)林、化香+曼青冈(Platycarya strobilacea+Cyclobalanopsis oxyodon)林、石栎(Lithocarpus glaber)林、甜槠(Castanopsis eyeri)林、红豆树(Ormosia henryi)林、润楠(Machilus pingii)林、大头茶(Gordoniaasillaris)林、刺叶栎(Quercus spinosa)林共33个群落类型;DCA排序结果较好地解释了库区森林植物群落分布及其与环境因子的关系,反映了植物群落所在地环境的温度、海拔和土壤水分的综合梯度,总体趋势表现为:随着海拔高度逐渐升高,水分逐渐减小,气温逐渐降低,植被类型依次为常绿阔叶林、常绿落叶阔叶混交林、落叶阔叶林、针阔叶混交林、针叶林。
三峡库区森林植物群落中灌木层的物种最丰富,其次是草本层,但不同群落之间是波动的。目前,由于环境差异及强烈的人为干扰活动,三峡库区森林植物群落物种多样性在海拔梯度上没有明显的规律。随着由灌丛向针叶林、针阔混交林、落叶阔叶林、常绿阔叶林的正向演替,林下土壤pH值逐步降低,有机质、速效磷逐渐升高,而水解氮逐渐降低,但三者均具有较强的表聚性。
在三峡库区针叶林约占库区植被的50%,其中马尾松林占34%,杉木林占10%,柏木林占6%;三种林分的物种多样性的总体趋势与库区森林的物种多样性趋势一致。对马尾松林下物种生态位测度表明,菝葜(Smilax davidiana)、铁仔(Myrsine africana)、细齿柃木(Eurya japonica)、小果蔷薇(Rosa cymosa)、黄栀子(Pittosporum phillyraeoides)、算盘子(Glochidion puberum)、乌饭(Vaccinium serrulatum)等灌木种群和铁芒萁(Dicranopteris lineari)、芒(Miscanthus sinensis)、蕨(Pteridium aquilinum)、白茅(Imperata cylindrica)、野古草(Arundinella anomala)、乌蕨(Stenoloma chusanum)等草本种群的生态位宽度较大;展毛野牡丹(Melastoma normle)、烟管荚迷(Viburnum utile)、肖梵天花(Urena lobata)等灌木种群和毛轴碎米蕨(Cheilosoria chusana)、淡竹叶(Lophatherum gracile)、假淡竹叶(Centotheca lappaceus)、黄背茅(Themeda japonica)、斑茅(Saccharum arunndinaceum)、水竹(Phyllostachys heteroclada)、扇叶铁线蕨(Adiantum flabellulatum)、掌叶铁线蕨(Adiantum pedatum)等草本种群的生态位宽度较小。其中灌木种群中细齿铃木和映山红(Rhododendron simsii)、土茯苓(Smilax glabra)和山胡椒(Lindera glauca)、黄荆(Vitex negundo)和地瓜藤(Ficus tikoua)、乌饭和黄栀子,草本种群中掌叶铁线蕨和阔鳞鳞毛蕨(Dryopteris championii)、乌蕨和马尔参(Miscanthus nepalensis)、扇叶铁线蕨和野古草(Arundinella anomala),生态位重叠较大。另外,同属物种,如烟管荚迷和宜昌荚迷(Viburnum erosum),菝葜、粉背菝葜(Smilax hypoglauca)和土茯苓,丝叶苔草(Carex capilliformis)、披针叶苔草(Carex lanceolata)、苔草(Carex baccans)和密叶苔草(Carex maubertiana),掌叶铁线蕨和扇叶铁线蕨,铁芒萁(Dicranopteris lineari)和芒萁(Dicranopteris dichotoma),金星蕨(Parathelypteris- landuligera)和日本金星蕨(Parathelypteris japonica)等之间的生态位重叠程度降低,由于在一定程度上对环境资源需求分化,更利于它们共存于某一小生境中,具有长期生存适应的生态学意义。
三峡库区马尾松林下灌木中继木和黄栀子为极显著的正关联种对,黄栌、土茯苓、马桑和展毛野牡丹为极显著的负关联。草本中极显著正关联种对为铁芒萁和芒、芒萁和红盖鳞毛蕨(Dryopteris erythrosor)、丝叶苔草和石荠宁(Mosla scabra)等;极显著的负关联为毛轴碎米蕨和淡竹叶、假淡竹叶,黄背茅和水竹等。根据种间联结特征及现实物种分布,将马尾松林下灌木和草本大致分为4个生态种组。第一种组为黄栀子、铁仔、继木、铁芒萁、芒、野古草、蕨等,第二种组为小铁仔、粉背菝葜、地瓜藤、芒萁、狗脊蕨、丝叶苔草、乌蕨等,第三种组为菝葜、细齿铃木、算盘子、毛轴碎米蕨、红盖鳞毛蕨、金星蕨等,第四种组为马桑、烟管荚迷、映山红、苔草、圆果雀稗(Paspalum orbiculare)、刚毛秀竹(Microstegium ciliatum)等。
Based on the investigated data of 132 temporal and 146 permanent plots and the data of national second-class forest resource investigation(1999-2003) in the Three Gorges Reservoir Area, species diversity of the forest communities in the area was mainly studied using the integration of quantitative and qualitative methods, especially quantitative analysis methods. Flora characteristics of seed plants, quantitative classification and ordination of plant communities and their environmental interpretations, plant species diversity and their responses to environment, and changes of soil nutrients along with community succession in the area were analyzed deeply together with the analysis of the community structure and species diversity, niche and interspecific association of under-story species of the main coniferous forest in the area. Current status, existing problems and conservation strategies of the rare and endangered species in the area were also analyzed finally. This study will be significance theoretically and practically of restoration, rehabilitation, and sustainable management for the degraded forest in the area.
Areal types of genera of seed plants in the area were divided in 15 types according to the method for Areal types’study of Prof. Wu Zhengyi. The dominant type was tropical areal type, then temperate areal type, and the transitional feature was obvious. The origin of the flora was ancient and there were plenty of plant species, including 5688 seed plant species, accounting for 22% of total species in China. There were 56 rare and endangered plant species, belonging to 34 families and 48 genera, and accounting for 10% of the total number in China approximately. They had transitional, decentralized and endemic characteristics and have been disturbed by human activities seriously.
Plant communities were divided into 33 types by the quantitative method TWINSPAN (TWo INdicator Species ANalysis) in the area. They were Form. Pinus henryi, Form. P. armandii, Form. P. massoniana, Form. Cunnighamia lanceolata, Form. Cupessus funebris, Form. Pinus massoniana+Quercus variabilis, Form. Pinus massoniana +Castanopsis fargesii, Form. Pinus massoniana+ Grdonia axillaris, Form. Cunnighamia lanceolata+Castamopsis fargesii, Form. Quercus glandulifera var. brevipetiolata, Form. Q. variabilis, Form. Q. albus, Form. Q. aliena+Q. variabilis, Form. Q. dentata var. oxyloba, Form. Fagus lucida, Form. Dendrobenthamia japonica var. chinensis, Form. Liquidambar taiwaniana, Form. Castanea seguinii, Form. Platycarya strobilacea, Form. Betaula albo-sinensis, Form. B. utilis, Form. Cerasus conrodinae+Quercus spinosa, Form. Cercidiphyllum japonicum+Padus obtusata, Form. Cornus controversa, Form. Celtis sinensis, Form. Lithocarpus cleistocarpus+Castanea henryi, Form. Platycarya strobilacea+Cyclobalanopsis oxyodon, Form. Lithocarpus glaber, Form. Castanopsis eyeri, Form. Ormosia henryi, Form. Machilus pingii, Form. Gordonia asillaris and Form. Quercus spinosa. Distribution of the plant communities and the relationships with the environmental factors were interpreted well by the result of DCA (Detrended Correspondence Analysis) ordination of the communities. Integration of temperature, elevation, soil water content of the community sites was reflected obviously by the DAC result. The main trend was that soil water content and temperature was decreasing with the elevation increasing, the vegetation types were evergreen broad- leaved forest, evergreen and deciduous broad-leaved mixed forest, deciduous broad-leaved forest, coniferous and board-leaved mixed forest, and coniferous forest respectively.
Plant species richness index of shrub was the highest and that of herb was higher, but they were fluctuant in different communities. Because of the environmental heterogeneity and the strong artificial disturbance, there was no obvious trend of the diversity indices of the communities along with the elevation. With the positive succession of the vegetation, shrub—coniferous forest—coniferous and broad-leaved mixed forest—deciduous broad-leaved forest—evergreen broad-leaved forest in the area, soil pH value decreased gradually, organic matter and available phosphorus of the soil increased while hydrolysable nitrogen decreased gradually, though all of them distributed principally in surface layer.
Coniferous forest occupied approximately 50% in the middle and low mountain area in the Three Gorges Reservoir Area, Pinus massoniana forest 34%, Cunnighamia lanceolata forest 10%, and Cupessus funebris forest 6%. The main trend of the species diversity of this three forest types was as same as that of the forest in the area. The niche breadths of the shrub species populations, Smilax davidiana,Myrsine Africana, Eurya japonica, Rosa cymosa, Pittosporum phillyraeoides, Glochidion puberum and Vaccinium serrulatum, and the herb species populations, Dicranopteris linearis, Miscanthus sinensis, Pteridium aquilinum, Imperata cylindrica, Arundinella anomala and Stenoloma chusanum, were higher, while that of the shrub species populations, Melastoma normale, Viburnum utile and Urena lobata, and the herb species populations, Cheilosoria chusana, Lophatherum gracile, Centotheca lappacea, Themeda japonica, Saccharum arunndinaceum, Phyllostachys heteroclada, Adiantum flabellulatum and Adiantum pedatum, was lower relatively in Pinus massoniana community. The niche overlaps between the shrub species populations of Eurya japonica and Rhododendron mucronulatum, Smilax glabra and Lindera glauca, Vitex negundo and Ficus tikoua, Vaccinium serrulatum and Pittosporum phillyraeoides, and herb species populations of Adiantum pedatum and Dryopteris championii, Stenoloma chusanum and Miscanthus nepalensis, Adiantum flabellulatum and Arundinella anomala, were higher in Pinus massoniana community. In addition, the overlaps among the species populations belonging to the same genus, Viburnum utile and V. erosum, Smilax davidiana, S. hypoglauca and S. glabra, Carex capilliformis, C. lanceolata, C. baccans and C. maubertiana, Adiantum pedatum and A. flabellulatum, Dicranopteris linearis and D. dichotoma, Parathelypteris glanduligera and P. japonica, were lower relatively. This could be helpful for the coexistence of the species in a small area.
The interspecific association between the shrub species populations, Loropetalum chinense and Pittosporum phillyraeoides, was positive significantly, while the associations between the populations of Cotinus coggygria, Smilax glabra, Coriaria nepalensis and Melastoma normle were negative significantly in Pinus massoniana community. The interspecific association between the herb species populations, Dicranopteris linearis and Miscanthus sinensis, D. dichotoma and Dryopteris erythrosora, C. capilliformis and Mosla scabra, were positive significantly, while the associations between the populations of Cheilosoria chusana and Lophatherum gracile, Centotheca lappacea, Themeda japonica and Phyllostachys heteroclada, were negative significantly. Understory species of the Pinus massoniana community were divided into four ecological species groups based on their intrespecific association characteristics. The first group included Pittosporum phillyraeoides, Myrsine africana, Loropetalum chinense, Rosa cymosa, Dicranopteris linearis, Miscanthus sinensis, Arundinella anomala and Pteridium aquilinum etc.; the second included Myrsine semiserrata, Smilax hypoglauca, Ficus tikoua, Dicranopteris dichotoma, Woodwardia japonica, Carex capilliformis and Stenoloma chusanum etc.; the third included Smilax davidiana, Eurya japonica, Glochidion puberum, Cheilosoria chusana, Dryopteris erythrosora and Parathelypteris glanduligera etc.; and the fourth included Coriaria nepalensis, Viburnum utile, Rhododendron mucronulatum, Carex baccans, Paspalum orbiculare and Microstegium ciliatum etc..
根据吴征镒先生对中国种子植物分布类型的划分,该区种子植物属的分布类型可划分为15个类型及其变型,以热带性质的属分布较多,温带性质的属次之,具有较为明显的过渡性。同时,该区域植物区系起源古老,植物种类丰富多样,共有种子植物5688种,占全国种子植物总数的21%。其中珍稀濒危植物56种,分属34科,48属,约占全国总数的10%。三峡库区珍稀濒危植物具有过渡性、分散性和特有性等特征,而且受人为干扰严重。
TWINSPAN数量分类结果表明三峡库区森林植物群落可化分为巴山松(Pinus henryi)林、华山松(P. armandii)林、马尾松(P. massoniana)林、杉木(Cunnighamia lanceolata)林、柏木(Cupessus funebris)林、马尾松+栓皮栎(Pinus massoniana+Quercus variabilis)林、马尾松+栲(Pinus massoniana +Castanopsis fargesii)林、马尾松+大头茶(Pinus massoniana+ Grdonia axillaris)林、杉木+栲(Cunnighamia lanceolata+Castamopsis fargesii)林、短柄枹栎（Quercus glandulifera var. brevipetiolata)林、栓皮栎(Q. variabilis)林、白栎(Q. albus)林、槲栎+栓皮栎(Q. aliena+Q. variabilis)林、西南槲树(Q. denlata var. oxyloba)林、亮叶水青冈(Fagus lucida)林、茅栗(Castanea seguinii)林、化香(Platycarya strobilacea)林、红桦(Betaula albo-sinensis)林、糙皮桦(Betula utilis)林、四照花(Dendrobenthamia japonica var. chinensis)林、枫香(Liquidambar formosana)林、华中樱桃+刺叶栎(Cerasus conrodinae+Quercus spinosa)林、连香树+细齿稠李(Cercidiphyllum japonicum+Padus obtusata)林、灯台树(Cornus controversa)林、朴树(Celtis sinensis)林、包槲柯+锥栗(Lithocarpus cleistocarpus+Castanea henryi)林、化香+曼青冈(Platycarya strobilacea+Cyclobalanopsis oxyodon)林、石栎(Lithocarpus glaber)林、甜槠(Castanopsis eyeri)林、红豆树(Ormosia henryi)林、润楠(Machilus pingii)林、大头茶(Gordoniaasillaris)林、刺叶栎(Quercus spinosa)林共33个群落类型;DCA排序结果较好地解释了库区森林植物群落分布及其与环境因子的关系,反映了植物群落所在地环境的温度、海拔和土壤水分的综合梯度,总体趋势表现为:随着海拔高度逐渐升高,水分逐渐减小,气温逐渐降低,植被类型依次为常绿阔叶林、常绿落叶阔叶混交林、落叶阔叶林、针阔叶混交林、针叶林。
三峡库区森林植物群落中灌木层的物种最丰富,其次是草本层,但不同群落之间是波动的。目前,由于环境差异及强烈的人为干扰活动,三峡库区森林植物群落物种多样性在海拔梯度上没有明显的规律。随着由灌丛向针叶林、针阔混交林、落叶阔叶林、常绿阔叶林的正向演替,林下土壤pH值逐步降低,有机质、速效磷逐渐升高,而水解氮逐渐降低,但三者均具有较强的表聚性。
在三峡库区针叶林约占库区植被的50%,其中马尾松林占34%,杉木林占10%,柏木林占6%;三种林分的物种多样性的总体趋势与库区森林的物种多样性趋势一致。对马尾松林下物种生态位测度表明,菝葜(Smilax davidiana)、铁仔(Myrsine africana)、细齿柃木(Eurya japonica)、小果蔷薇(Rosa cymosa)、黄栀子(Pittosporum phillyraeoides)、算盘子(Glochidion puberum)、乌饭(Vaccinium serrulatum)等灌木种群和铁芒萁(Dicranopteris lineari)、芒(Miscanthus sinensis)、蕨(Pteridium aquilinum)、白茅(Imperata cylindrica)、野古草(Arundinella anomala)、乌蕨(Stenoloma chusanum)等草本种群的生态位宽度较大;展毛野牡丹(Melastoma normle)、烟管荚迷(Viburnum utile)、肖梵天花(Urena lobata)等灌木种群和毛轴碎米蕨(Cheilosoria chusana)、淡竹叶(Lophatherum gracile)、假淡竹叶(Centotheca lappaceus)、黄背茅(Themeda japonica)、斑茅(Saccharum arunndinaceum)、水竹(Phyllostachys heteroclada)、扇叶铁线蕨(Adiantum flabellulatum)、掌叶铁线蕨(Adiantum pedatum)等草本种群的生态位宽度较小。其中灌木种群中细齿铃木和映山红(Rhododendron simsii)、土茯苓(Smilax glabra)和山胡椒(Lindera glauca)、黄荆(Vitex negundo)和地瓜藤(Ficus tikoua)、乌饭和黄栀子,草本种群中掌叶铁线蕨和阔鳞鳞毛蕨(Dryopteris championii)、乌蕨和马尔参(Miscanthus nepalensis)、扇叶铁线蕨和野古草(Arundinella anomala),生态位重叠较大。另外,同属物种,如烟管荚迷和宜昌荚迷(Viburnum erosum),菝葜、粉背菝葜(Smilax hypoglauca)和土茯苓,丝叶苔草(Carex capilliformis)、披针叶苔草(Carex lanceolata)、苔草(Carex baccans)和密叶苔草(Carex maubertiana),掌叶铁线蕨和扇叶铁线蕨,铁芒萁(Dicranopteris lineari)和芒萁(Dicranopteris dichotoma),金星蕨(Parathelypteris- landuligera)和日本金星蕨(Parathelypteris japonica)等之间的生态位重叠程度降低,由于在一定程度上对环境资源需求分化,更利于它们共存于某一小生境中,具有长期生存适应的生态学意义。
三峡库区马尾松林下灌木中继木和黄栀子为极显著的正关联种对,黄栌、土茯苓、马桑和展毛野牡丹为极显著的负关联。草本中极显著正关联种对为铁芒萁和芒、芒萁和红盖鳞毛蕨(Dryopteris erythrosor)、丝叶苔草和石荠宁(Mosla scabra)等;极显著的负关联为毛轴碎米蕨和淡竹叶、假淡竹叶,黄背茅和水竹等。根据种间联结特征及现实物种分布,将马尾松林下灌木和草本大致分为4个生态种组。第一种组为黄栀子、铁仔、继木、铁芒萁、芒、野古草、蕨等,第二种组为小铁仔、粉背菝葜、地瓜藤、芒萁、狗脊蕨、丝叶苔草、乌蕨等,第三种组为菝葜、细齿铃木、算盘子、毛轴碎米蕨、红盖鳞毛蕨、金星蕨等,第四种组为马桑、烟管荚迷、映山红、苔草、圆果雀稗(Paspalum orbiculare)、刚毛秀竹(Microstegium ciliatum)等。
Based on the investigated data of 132 temporal and 146 permanent plots and the data of national second-class forest resource investigation(1999-2003) in the Three Gorges Reservoir Area, species diversity of the forest communities in the area was mainly studied using the integration of quantitative and qualitative methods, especially quantitative analysis methods. Flora characteristics of seed plants, quantitative classification and ordination of plant communities and their environmental interpretations, plant species diversity and their responses to environment, and changes of soil nutrients along with community succession in the area were analyzed deeply together with the analysis of the community structure and species diversity, niche and interspecific association of under-story species of the main coniferous forest in the area. Current status, existing problems and conservation strategies of the rare and endangered species in the area were also analyzed finally. This study will be significance theoretically and practically of restoration, rehabilitation, and sustainable management for the degraded forest in the area.
Areal types of genera of seed plants in the area were divided in 15 types according to the method for Areal types’study of Prof. Wu Zhengyi. The dominant type was tropical areal type, then temperate areal type, and the transitional feature was obvious. The origin of the flora was ancient and there were plenty of plant species, including 5688 seed plant species, accounting for 22% of total species in China. There were 56 rare and endangered plant species, belonging to 34 families and 48 genera, and accounting for 10% of the total number in China approximately. They had transitional, decentralized and endemic characteristics and have been disturbed by human activities seriously.
Plant communities were divided into 33 types by the quantitative method TWINSPAN (TWo INdicator Species ANalysis) in the area. They were Form. Pinus henryi, Form. P. armandii, Form. P. massoniana, Form. Cunnighamia lanceolata, Form. Cupessus funebris, Form. Pinus massoniana+Quercus variabilis, Form. Pinus massoniana +Castanopsis fargesii, Form. Pinus massoniana+ Grdonia axillaris, Form. Cunnighamia lanceolata+Castamopsis fargesii, Form. Quercus glandulifera var. brevipetiolata, Form. Q. variabilis, Form. Q. albus, Form. Q. aliena+Q. variabilis, Form. Q. dentata var. oxyloba, Form. Fagus lucida, Form. Dendrobenthamia japonica var. chinensis, Form. Liquidambar taiwaniana, Form. Castanea seguinii, Form. Platycarya strobilacea, Form. Betaula albo-sinensis, Form. B. utilis, Form. Cerasus conrodinae+Quercus spinosa, Form. Cercidiphyllum japonicum+Padus obtusata, Form. Cornus controversa, Form. Celtis sinensis, Form. Lithocarpus cleistocarpus+Castanea henryi, Form. Platycarya strobilacea+Cyclobalanopsis oxyodon, Form. Lithocarpus glaber, Form. Castanopsis eyeri, Form. Ormosia henryi, Form. Machilus pingii, Form. Gordonia asillaris and Form. Quercus spinosa. Distribution of the plant communities and the relationships with the environmental factors were interpreted well by the result of DCA (Detrended Correspondence Analysis) ordination of the communities. Integration of temperature, elevation, soil water content of the community sites was reflected obviously by the DAC result. The main trend was that soil water content and temperature was decreasing with the elevation increasing, the vegetation types were evergreen broad- leaved forest, evergreen and deciduous broad-leaved mixed forest, deciduous broad-leaved forest, coniferous and board-leaved mixed forest, and coniferous forest respectively.
Plant species richness index of shrub was the highest and that of herb was higher, but they were fluctuant in different communities. Because of the environmental heterogeneity and the strong artificial disturbance, there was no obvious trend of the diversity indices of the communities along with the elevation. With the positive succession of the vegetation, shrub—coniferous forest—coniferous and broad-leaved mixed forest—deciduous broad-leaved forest—evergreen broad-leaved forest in the area, soil pH value decreased gradually, organic matter and available phosphorus of the soil increased while hydrolysable nitrogen decreased gradually, though all of them distributed principally in surface layer.
Coniferous forest occupied approximately 50% in the middle and low mountain area in the Three Gorges Reservoir Area, Pinus massoniana forest 34%, Cunnighamia lanceolata forest 10%, and Cupessus funebris forest 6%. The main trend of the species diversity of this three forest types was as same as that of the forest in the area. The niche breadths of the shrub species populations, Smilax davidiana,Myrsine Africana, Eurya japonica, Rosa cymosa, Pittosporum phillyraeoides, Glochidion puberum and Vaccinium serrulatum, and the herb species populations, Dicranopteris linearis, Miscanthus sinensis, Pteridium aquilinum, Imperata cylindrica, Arundinella anomala and Stenoloma chusanum, were higher, while that of the shrub species populations, Melastoma normale, Viburnum utile and Urena lobata, and the herb species populations, Cheilosoria chusana, Lophatherum gracile, Centotheca lappacea, Themeda japonica, Saccharum arunndinaceum, Phyllostachys heteroclada, Adiantum flabellulatum and Adiantum pedatum, was lower relatively in Pinus massoniana community. The niche overlaps between the shrub species populations of Eurya japonica and Rhododendron mucronulatum, Smilax glabra and Lindera glauca, Vitex negundo and Ficus tikoua, Vaccinium serrulatum and Pittosporum phillyraeoides, and herb species populations of Adiantum pedatum and Dryopteris championii, Stenoloma chusanum and Miscanthus nepalensis, Adiantum flabellulatum and Arundinella anomala, were higher in Pinus massoniana community. In addition, the overlaps among the species populations belonging to the same genus, Viburnum utile and V. erosum, Smilax davidiana, S. hypoglauca and S. glabra, Carex capilliformis, C. lanceolata, C. baccans and C. maubertiana, Adiantum pedatum and A. flabellulatum, Dicranopteris linearis and D. dichotoma, Parathelypteris glanduligera and P. japonica, were lower relatively. This could be helpful for the coexistence of the species in a small area.
The interspecific association between the shrub species populations, Loropetalum chinense and Pittosporum phillyraeoides, was positive significantly, while the associations between the populations of Cotinus coggygria, Smilax glabra, Coriaria nepalensis and Melastoma normle were negative significantly in Pinus massoniana community. The interspecific association between the herb species populations, Dicranopteris linearis and Miscanthus sinensis, D. dichotoma and Dryopteris erythrosora, C. capilliformis and Mosla scabra, were positive significantly, while the associations between the populations of Cheilosoria chusana and Lophatherum gracile, Centotheca lappacea, Themeda japonica and Phyllostachys heteroclada, were negative significantly. Understory species of the Pinus massoniana community were divided into four ecological species groups based on their intrespecific association characteristics. The first group included Pittosporum phillyraeoides, Myrsine africana, Loropetalum chinense, Rosa cymosa, Dicranopteris linearis, Miscanthus sinensis, Arundinella anomala and Pteridium aquilinum etc.; the second included Myrsine semiserrata, Smilax hypoglauca, Ficus tikoua, Dicranopteris dichotoma, Woodwardia japonica, Carex capilliformis and Stenoloma chusanum etc.; the third included Smilax davidiana, Eurya japonica, Glochidion puberum, Cheilosoria chusana, Dryopteris erythrosora and Parathelypteris glanduligera etc.; and the fourth included Coriaria nepalensis, Viburnum utile, Rhododendron mucronulatum, Carex baccans, Paspalum orbiculare and Microstegium ciliatum etc..
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