金华北山常绿阔叶林群落特征研究
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摘要
中国的常绿阔叶林分布面积大,类型多样,是世界常绿阔叶林的主体,在保护区域生态环境、维持经济社会可持续发展方面具有重要作用。长期以来由于人们对其功能和作用的认识不足,曾遭到严重破坏,导致土壤退化、功能衰退、病虫害频繁等一系列问题。因此,在各地开展常绿阔叶林群落特征的研究工作就显得尤为重要和迫切。本研究以金华北山常绿阔叶林为研究对象,在野外调查的基础上,综合应用数量生态学的方法,对金华北山常绿阔叶林群落的植物区系、群落外貌、结构特征、空间分布、种群更新、种内种间竞争以及物种多样性进行研究,得到以下研究结果：
(1)金华北山常绿阔叶林群落共有维管植物66科115属144种,其中蕨类植物8种,裸子植物2科2属2种；被子植物58科105属134种。在81种木本植物中,以壳斗科、樟科、山茶科等树种为主的常绿树种占有重要优势。这些均体现了我国亚热带常绿阔叶林群落的结构和组成特征。金华北山常绿阔叶林群落种子植物的区系成分复杂,具有明显的过渡性质。优势科的区系分析中,以热带成分为主,温带成分也占有一定比例；在属的区系分析中,热带成分占有重要优势,但温带成分更多,超过了热带成分的比例。群落中高位芽植物占有绝对优势,占总数的78.68%。叶的性质分析表明,金华北山常绿阔叶林群落以中、小型的全缘、草质、单叶植物为主。群落的区系组成和外貌特征都体现出中亚热带北缘常绿阔叶林的特征,反映出相对湿润温暖的气候特点。
(2)整个群落的垂直结构可分为四个层次,其中乔木层植物有49种,灌木层植物有84种,草本植物有26种,层间植物有34种；采用Shannon-Wiener指数、Simpson指数、Pielou指数对群落各层次的物种多样性进行测定,结果表明：各指数变化趋势相同,均表现为灌木层>乔木层>层间>草本层,符合亚热带常绿阔叶林群落的一般规律。对优势树种的重要值分析得知,群落的建群树种为木荷(Schima superba)、苦槠(Castanopsis sclerophylla)、枫香(Liquidambar formosana)和马尾松(Pinus massoniana),它们都是亚热带地区植被的典型代表,反映出群落的基本面貌,体现了亚热带常绿阔叶林的特点。应用聚块性指数、扩散系数、负二项式分布等指标研究各群落优势种群的分布格局,结果表明：金华北山常绿阔叶林群落水平结构的聚集性特征明显,15个优势树种中有14个种群表现为集群分布,其中的常绿树种木荷、苦槠、香樟(Cinnamomum camphora)、女贞(Ligustrum lucidum)、山矾(Symplocos sumuntia)、格药柃(Eurya muricata)等聚集程度较高,而麻栎(Quercus acutissima)、枫香、白栎(Quercus fabri)、杉木(Cunninghamia lanceolata)等聚集程度相对较低,马尾松更是呈现出均匀分布特征。该空间格局的形成与各树种的生物学特性以及群落演替过程中个体间的种内、种间竞争有关。
(3)采用空间代替时间的方法,将优势种群的年龄结构分为四种更新类型,并结合其生物学和生态学特性,对群落所处的演替阶段进行了诊断,结果为：逆-J字型的木荷、苦槠、香樟,都是耐荫性强的常绿阔叶乔木,群落中除有成熟的大径级个体外,林下还有大量的幼树和幼苗的存在,是该区亚热带常绿阔叶林顶极群落的优势种和主要组成种；L型的檵木(Loropetalum chinensis)、女贞、乌饭(Vaccinium bracteatum)、山矾、格药柃等均为常绿阔叶小乔木或者大灌木,耐荫性强,小径级个体数量较多,种群结构稳定,是顶极群落亚乔木层和灌木层的主要组成种；间歇型的枫香、麻栎、锥栗(Castanea henryi)、青冈栎(Cyclobalanopsis glauca),是喜光性树种,耐荫性差,经常出现在林窗内或林缘部,更新具有机会性和波动性特征,是林冠组成种中尚未分化完全的顶级性先锋种；纺锤型的白栎、马尾松和杉木均为阳性树种,在样地内只有少量大胸径个体出现,更新个体缺乏,是先锋种或林窗更新种,在演替后期比较稳定的群落内,将最终衰退消失；本群落可视为顶级群落,但只是处于顶级演替的前期,并未达到最终阶段,将进一步向以木荷、苦槠为优势种的方向发展。
(4)根据金华北山常绿阔叶林群落中各树种的重要值大小,采用2×2联列表,综合运用方差分析、χ2检验、联结系数AC、共同出现百分率PC及点相关系数Φ,对15个优势树种进行种间联结性研究。结果表明：15个树种间总体呈现不显著的正联结关系,反映出群落结构的不稳定性；正联结种对数略大于负联结种对数,大多数种对的种间联结性较弱,其中马尾松与杉木,杉木与山矾,乌饭与锥栗的联结关系更是表现为完全独立,这说明各种群间独立性较强,没有形成相对稳定的协调搭配机制。这种种间联结关系的松散性,表明目前群落正处于顶级演替的前期阶段。
(5)运用生态位宽度、生态位重叠值以及生态位相似比例等指标对金华北山常绿阔叶林中15个优势树种的生态位特征进行测定与分析。研究结果表明：在群落的15种优势树种中,木荷、枫香、格药柃、苦槠、马尾松、杉木的Levins和Shannon-Wiener生态位宽度值较大；15个优势树种的生态位重叠和生态相似性比例均较高,由它们组成的105个种对中有15对生态位重叠值大于0.8,有76.2%的种对大于0.4,而且生态位宽度大的树种间一般有较高的生态位重叠和生态位相似性比例。群落中生态位重叠较大的木荷与枫香,苦槠与马尾松,马尾松与格药柃等种对,彼此存在对资源的利用性竞争；而木荷与格药柃、檵木、山矾、乌饭树,位于群落的不同层次,木荷与它们能够共享资源；格药柃、檵木、山矾、乌饭树之间具有相似的生物生态学特性,在资源不足时因竞争而发生生态位重叠,在资源充足时共享资源。随着演替的进行,马尾松、杉木、枫香等阳性先锋树种将逐渐退出群落,木荷、苦槠、格药柃等常绿树种得到进一步发展,各种群生态位也将随之变化直至达到稳定的演替顶级。
Chinese evergreen broad-leaved forest, the main part of the evergreen broad-leaved forest in the world, has the large distribution area and various types, and has an important role in protecting the region ecological environment, maintaining sustainable economic and social development. For a long time, because of inadequate understanding of evergreen broad-leaved forest in its function and effect, it had been damaged and caused series of problems, such as soil degradation, function decline, pests'frequent. Therefore, the research work of evergreen broad-leaved forest community characteristic appears especially important and urgent. This paper is a report about studies on floristic characteristics, community physiognomy, structural features, distribution pattern, intraspecific and interspecific competition, population restoration and species diversity of the evergreen broad-leaved forest in Beishan mountain of Jinhua. The research was based on the field investigation and quantitative ecology methods. The main obtained results are as follows:
(1) There were 144 species,66families and 115 genera of vascular plants, including 8 ferns,2 families,2 genera and 2 species of gymnosperms,58 families,105genera and 134 species in the evergreen broad-leaved forest community of Beishan Mountain in Jinhua. The evergreen trees of Fagaceae, Lauraceae and Theaceae occupied a dominant position in 81 woody plants. And these characters reflected community structure and composition features of subtropical evergreen broad-leaved forest in China. The floristic characteristics of seed plants in the community had complicated composition significant transition properties. The floristic analysis of dominant families showed that tropic elements occupied the dominant position, and the temperate components were also important. The floristic analysis of genera showed tropical components also had important advantages, but more of them were temperate components. The high buds plant in the community had an absolute advantage, total 78.68%. The analysis of leaf characters showed most of the plant were small and medium-sized, orthophyll, entire and sample leaf. Whatever flora composition or community appearance, they were all reflected the characteristics of evergreen broad-leaved forest in subtropical northern margin, and showed the relatively warm and humid climate characteristics.
(2) The vertical structure of the evergreen broad-leaved forest could be divided into four levels, including tree layer with 49 species, shrub layer with 84 species, herb layer with 26 species and inter layer with 34 species. The species diversity of the four layers was analyzed by using Shannon-Wiener index, Simpson index, and Pielou index. The results showed that:each index had the same change trend, and represented general rules which were shrub layer>tree layer>inter layer>herb layer in the subtropical evergreen broad-leaved forest community. The important value analysis of superiority species showed that the building-group trees in the community were Schima superba, Castanopsis sclerophylla, Liquidambar formosana, Pinus massoniana, and as representative trees, they reflected the basic appearance and characteristics of subtropical evergreen broad-leaved forest community. The distribution pattern of dominate trees was studied by using patchiness index, dispersion index, negative binomial parameter et al. The results showed that aggregation features of level structure in the forest were obvious, and there were 14 cluster distribution populations in 15 dominate trees, and the gathered degree of Schima superba, Castanopsis sclerophylla, Cinnamomum camphora, Ligustrum lucidum, Eurya muricata et al. were higher, while Quercus acutissima, Liquidambar formosana, Quercus fabric and Cunninghamia lanceolata showed lower gathered degree, especially, Pinus massoniana showed uniform distribution. The distribution pattern was concerned with the biological characteristics, and the intraspecific and interspecific competition in community succession process.
(3) The age structure and development of dominant population were categorized into four regeneration types by the method of space for time sere. After that, combining the biological with ecological characteristics of that species, the diagnosis of succession stage of this community was carried out. The results were as follows:(1) Inverse-J type species, such as Schima superba, Castanopsis sclerophylla and Cinnamomum camphora, were strong shade-tolerant and dominant species in climax forest, they could regenerate through seedling bank or sprouting under the closed forest canopy; L type species, such as Loropetalum chinensis, Ligustrum lucidum and Ewya muricata, were broad-leaved evergreen small trees or large shrubs, and they were the main components of Iaccio wood layer and shrub layer species in climax community; Sporadic (multimodal) type species, such as Liquidambar formosana, Quercus acutissima and Castanea henryi, had the opportunity and volatility to the population structure, reflecting the characteristics of top pioneer species; Spindle type species, such as Quercus fabri, Pinus massoniana and Cunninghamia lanceolata, were high positive and the pioneer species in the successional sequence, but they had showed a recessionary signs because of the lack of updated population in the community. (2) The successional stage of the forest was classified as a late seral stage community that would develop into a climax community dominated by Schima superba and Castanopsis sclerophylla.
(4) Based on the important value and 2×2 contingency table, interspecific association of 15 tree species in the evergreen broad-leaved forest of Beishan mountain were examined by using variance ratio (VR),χ2 text, association coefficient (AC), co-occurrence percentage (PC) and point related coefficient (Φ). The results showed that the interspecific relation of 15 species took on a not significantly positive coupling, which reflected instability of the community structure, and the number of positive coupling was slightly more than the negative coupling, most of the interspecific relation had weak coupling and some even had no relationship, such as Pinus massoniana and Cunninghamia lanceolata, Cunninghamia lanceolata and Symplocos sumuntia, Vaccinium bracteatum and Castanea henryi. This laxity feature between various groups reflected the community had not form a relatively stable harmonious collocation mechanism and was at the prophase of the top succession stage.
(5) The niche characteristics of 15 dominate species in the evergreen broad-leaved forest were determined and analyzed by using niche breath, niche overlap and niche similarity proportion. The results showed that Schima superba, Liquidambar formosana, Eurya muricata, Castanopsis sclerophylla, Pinus massoniana and Cunninghamia lanceolata had bigger niche breath value of Levins and Shannon-Wiener in the community, the niche overlap and niche similarity proportion of dominate trees were higher, and the number of niche overlap value which was greater than 0.8 was 15 in 105 pairs constituted by 15 dominate species and 76.2% was greater than 0.4 and the trees which had big niche breadth often had higher niche overlap and niche similarity proportion value. Schima superba and Liquidambar formosana, Castanopsis sclerophylla and Pinus massoniana, Pinus massoniana and Eurya muricata which had bigger niche overlap value in the community, they existed competition of resources utilization each other. Schima superba and Eurya muricata, Loropetalum chinensis, Symplocos sumuntia, Vaccinium bracteatum located in different levels in the community, schima superba could share resources with them. And Eurya muricata, Loropetalum chinensis, Symplocos sumuntia, Vaccinium bracteatum had similar bioecology characteristics; they would compete with each other, when the resources were short, but share in enough resources. The positive pioneer species such as Pinus massoniana, Cunninghamia lanceolata, and Liquidambar formosana would gradually withdraw from the community, while the evergreen trees such as Schima superba, Castanopsis sclerophylla and Eurya muricata would have further development, and the niches of various populations would also change until the community reaches the stable succession climax.
(1)金华北山常绿阔叶林群落共有维管植物66科115属144种,其中蕨类植物8种,裸子植物2科2属2种；被子植物58科105属134种。在81种木本植物中,以壳斗科、樟科、山茶科等树种为主的常绿树种占有重要优势。这些均体现了我国亚热带常绿阔叶林群落的结构和组成特征。金华北山常绿阔叶林群落种子植物的区系成分复杂,具有明显的过渡性质。优势科的区系分析中,以热带成分为主,温带成分也占有一定比例；在属的区系分析中,热带成分占有重要优势,但温带成分更多,超过了热带成分的比例。群落中高位芽植物占有绝对优势,占总数的78.68%。叶的性质分析表明,金华北山常绿阔叶林群落以中、小型的全缘、草质、单叶植物为主。群落的区系组成和外貌特征都体现出中亚热带北缘常绿阔叶林的特征,反映出相对湿润温暖的气候特点。
(2)整个群落的垂直结构可分为四个层次,其中乔木层植物有49种,灌木层植物有84种,草本植物有26种,层间植物有34种；采用Shannon-Wiener指数、Simpson指数、Pielou指数对群落各层次的物种多样性进行测定,结果表明：各指数变化趋势相同,均表现为灌木层>乔木层>层间>草本层,符合亚热带常绿阔叶林群落的一般规律。对优势树种的重要值分析得知,群落的建群树种为木荷(Schima superba)、苦槠(Castanopsis sclerophylla)、枫香(Liquidambar formosana)和马尾松(Pinus massoniana),它们都是亚热带地区植被的典型代表,反映出群落的基本面貌,体现了亚热带常绿阔叶林的特点。应用聚块性指数、扩散系数、负二项式分布等指标研究各群落优势种群的分布格局,结果表明：金华北山常绿阔叶林群落水平结构的聚集性特征明显,15个优势树种中有14个种群表现为集群分布,其中的常绿树种木荷、苦槠、香樟(Cinnamomum camphora)、女贞(Ligustrum lucidum)、山矾(Symplocos sumuntia)、格药柃(Eurya muricata)等聚集程度较高,而麻栎(Quercus acutissima)、枫香、白栎(Quercus fabri)、杉木(Cunninghamia lanceolata)等聚集程度相对较低,马尾松更是呈现出均匀分布特征。该空间格局的形成与各树种的生物学特性以及群落演替过程中个体间的种内、种间竞争有关。
(3)采用空间代替时间的方法,将优势种群的年龄结构分为四种更新类型,并结合其生物学和生态学特性,对群落所处的演替阶段进行了诊断,结果为：逆-J字型的木荷、苦槠、香樟,都是耐荫性强的常绿阔叶乔木,群落中除有成熟的大径级个体外,林下还有大量的幼树和幼苗的存在,是该区亚热带常绿阔叶林顶极群落的优势种和主要组成种；L型的檵木(Loropetalum chinensis)、女贞、乌饭(Vaccinium bracteatum)、山矾、格药柃等均为常绿阔叶小乔木或者大灌木,耐荫性强,小径级个体数量较多,种群结构稳定,是顶极群落亚乔木层和灌木层的主要组成种；间歇型的枫香、麻栎、锥栗(Castanea henryi)、青冈栎(Cyclobalanopsis glauca),是喜光性树种,耐荫性差,经常出现在林窗内或林缘部,更新具有机会性和波动性特征,是林冠组成种中尚未分化完全的顶级性先锋种；纺锤型的白栎、马尾松和杉木均为阳性树种,在样地内只有少量大胸径个体出现,更新个体缺乏,是先锋种或林窗更新种,在演替后期比较稳定的群落内,将最终衰退消失；本群落可视为顶级群落,但只是处于顶级演替的前期,并未达到最终阶段,将进一步向以木荷、苦槠为优势种的方向发展。
(4)根据金华北山常绿阔叶林群落中各树种的重要值大小,采用2×2联列表,综合运用方差分析、χ2检验、联结系数AC、共同出现百分率PC及点相关系数Φ,对15个优势树种进行种间联结性研究。结果表明：15个树种间总体呈现不显著的正联结关系,反映出群落结构的不稳定性；正联结种对数略大于负联结种对数,大多数种对的种间联结性较弱,其中马尾松与杉木,杉木与山矾,乌饭与锥栗的联结关系更是表现为完全独立,这说明各种群间独立性较强,没有形成相对稳定的协调搭配机制。这种种间联结关系的松散性,表明目前群落正处于顶级演替的前期阶段。
(5)运用生态位宽度、生态位重叠值以及生态位相似比例等指标对金华北山常绿阔叶林中15个优势树种的生态位特征进行测定与分析。研究结果表明：在群落的15种优势树种中,木荷、枫香、格药柃、苦槠、马尾松、杉木的Levins和Shannon-Wiener生态位宽度值较大；15个优势树种的生态位重叠和生态相似性比例均较高,由它们组成的105个种对中有15对生态位重叠值大于0.8,有76.2%的种对大于0.4,而且生态位宽度大的树种间一般有较高的生态位重叠和生态位相似性比例。群落中生态位重叠较大的木荷与枫香,苦槠与马尾松,马尾松与格药柃等种对,彼此存在对资源的利用性竞争；而木荷与格药柃、檵木、山矾、乌饭树,位于群落的不同层次,木荷与它们能够共享资源；格药柃、檵木、山矾、乌饭树之间具有相似的生物生态学特性,在资源不足时因竞争而发生生态位重叠,在资源充足时共享资源。随着演替的进行,马尾松、杉木、枫香等阳性先锋树种将逐渐退出群落,木荷、苦槠、格药柃等常绿树种得到进一步发展,各种群生态位也将随之变化直至达到稳定的演替顶级。
Chinese evergreen broad-leaved forest, the main part of the evergreen broad-leaved forest in the world, has the large distribution area and various types, and has an important role in protecting the region ecological environment, maintaining sustainable economic and social development. For a long time, because of inadequate understanding of evergreen broad-leaved forest in its function and effect, it had been damaged and caused series of problems, such as soil degradation, function decline, pests'frequent. Therefore, the research work of evergreen broad-leaved forest community characteristic appears especially important and urgent. This paper is a report about studies on floristic characteristics, community physiognomy, structural features, distribution pattern, intraspecific and interspecific competition, population restoration and species diversity of the evergreen broad-leaved forest in Beishan mountain of Jinhua. The research was based on the field investigation and quantitative ecology methods. The main obtained results are as follows:
(1) There were 144 species,66families and 115 genera of vascular plants, including 8 ferns,2 families,2 genera and 2 species of gymnosperms,58 families,105genera and 134 species in the evergreen broad-leaved forest community of Beishan Mountain in Jinhua. The evergreen trees of Fagaceae, Lauraceae and Theaceae occupied a dominant position in 81 woody plants. And these characters reflected community structure and composition features of subtropical evergreen broad-leaved forest in China. The floristic characteristics of seed plants in the community had complicated composition significant transition properties. The floristic analysis of dominant families showed that tropic elements occupied the dominant position, and the temperate components were also important. The floristic analysis of genera showed tropical components also had important advantages, but more of them were temperate components. The high buds plant in the community had an absolute advantage, total 78.68%. The analysis of leaf characters showed most of the plant were small and medium-sized, orthophyll, entire and sample leaf. Whatever flora composition or community appearance, they were all reflected the characteristics of evergreen broad-leaved forest in subtropical northern margin, and showed the relatively warm and humid climate characteristics.
(2) The vertical structure of the evergreen broad-leaved forest could be divided into four levels, including tree layer with 49 species, shrub layer with 84 species, herb layer with 26 species and inter layer with 34 species. The species diversity of the four layers was analyzed by using Shannon-Wiener index, Simpson index, and Pielou index. The results showed that:each index had the same change trend, and represented general rules which were shrub layer>tree layer>inter layer>herb layer in the subtropical evergreen broad-leaved forest community. The important value analysis of superiority species showed that the building-group trees in the community were Schima superba, Castanopsis sclerophylla, Liquidambar formosana, Pinus massoniana, and as representative trees, they reflected the basic appearance and characteristics of subtropical evergreen broad-leaved forest community. The distribution pattern of dominate trees was studied by using patchiness index, dispersion index, negative binomial parameter et al. The results showed that aggregation features of level structure in the forest were obvious, and there were 14 cluster distribution populations in 15 dominate trees, and the gathered degree of Schima superba, Castanopsis sclerophylla, Cinnamomum camphora, Ligustrum lucidum, Eurya muricata et al. were higher, while Quercus acutissima, Liquidambar formosana, Quercus fabric and Cunninghamia lanceolata showed lower gathered degree, especially, Pinus massoniana showed uniform distribution. The distribution pattern was concerned with the biological characteristics, and the intraspecific and interspecific competition in community succession process.
(3) The age structure and development of dominant population were categorized into four regeneration types by the method of space for time sere. After that, combining the biological with ecological characteristics of that species, the diagnosis of succession stage of this community was carried out. The results were as follows:(1) Inverse-J type species, such as Schima superba, Castanopsis sclerophylla and Cinnamomum camphora, were strong shade-tolerant and dominant species in climax forest, they could regenerate through seedling bank or sprouting under the closed forest canopy; L type species, such as Loropetalum chinensis, Ligustrum lucidum and Ewya muricata, were broad-leaved evergreen small trees or large shrubs, and they were the main components of Iaccio wood layer and shrub layer species in climax community; Sporadic (multimodal) type species, such as Liquidambar formosana, Quercus acutissima and Castanea henryi, had the opportunity and volatility to the population structure, reflecting the characteristics of top pioneer species; Spindle type species, such as Quercus fabri, Pinus massoniana and Cunninghamia lanceolata, were high positive and the pioneer species in the successional sequence, but they had showed a recessionary signs because of the lack of updated population in the community. (2) The successional stage of the forest was classified as a late seral stage community that would develop into a climax community dominated by Schima superba and Castanopsis sclerophylla.
(4) Based on the important value and 2×2 contingency table, interspecific association of 15 tree species in the evergreen broad-leaved forest of Beishan mountain were examined by using variance ratio (VR),χ2 text, association coefficient (AC), co-occurrence percentage (PC) and point related coefficient (Φ). The results showed that the interspecific relation of 15 species took on a not significantly positive coupling, which reflected instability of the community structure, and the number of positive coupling was slightly more than the negative coupling, most of the interspecific relation had weak coupling and some even had no relationship, such as Pinus massoniana and Cunninghamia lanceolata, Cunninghamia lanceolata and Symplocos sumuntia, Vaccinium bracteatum and Castanea henryi. This laxity feature between various groups reflected the community had not form a relatively stable harmonious collocation mechanism and was at the prophase of the top succession stage.
(5) The niche characteristics of 15 dominate species in the evergreen broad-leaved forest were determined and analyzed by using niche breath, niche overlap and niche similarity proportion. The results showed that Schima superba, Liquidambar formosana, Eurya muricata, Castanopsis sclerophylla, Pinus massoniana and Cunninghamia lanceolata had bigger niche breath value of Levins and Shannon-Wiener in the community, the niche overlap and niche similarity proportion of dominate trees were higher, and the number of niche overlap value which was greater than 0.8 was 15 in 105 pairs constituted by 15 dominate species and 76.2% was greater than 0.4 and the trees which had big niche breadth often had higher niche overlap and niche similarity proportion value. Schima superba and Liquidambar formosana, Castanopsis sclerophylla and Pinus massoniana, Pinus massoniana and Eurya muricata which had bigger niche overlap value in the community, they existed competition of resources utilization each other. Schima superba and Eurya muricata, Loropetalum chinensis, Symplocos sumuntia, Vaccinium bracteatum located in different levels in the community, schima superba could share resources with them. And Eurya muricata, Loropetalum chinensis, Symplocos sumuntia, Vaccinium bracteatum had similar bioecology characteristics; they would compete with each other, when the resources were short, but share in enough resources. The positive pioneer species such as Pinus massoniana, Cunninghamia lanceolata, and Liquidambar formosana would gradually withdraw from the community, while the evergreen trees such as Schima superba, Castanopsis sclerophylla and Eurya muricata would have further development, and the niches of various populations would also change until the community reaches the stable succession climax.
引文
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