海南岛热带山地雨林植物功能群划分及生态关键种的确定
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摘要
热带森林对全球生物多样性保护和生态系统功能的维系起着非常重要的作用。然而，热带林极高的物种多样性和结构复杂性，使得生态分析非常困难，若将其物种按照一定规则划归为不同的功能群，将十分有利于热带林的科学研究和经营管理。本文以海南岛霸王岭林区作为海南岛热带林的典型区域，以反映不同环境因子、干扰类型和恢复时间的热带山地雨林为研究对象，在大面积(15hm~2)样地调查的基础上，依据物种的7个功能特性(喜光性、潜在高度、板根大小、木材密度、种子生物量、传播方式和落叶／常绿性)，分别把物种划分为不同类别的功能群，应用NMS分析了各功能群随不同干扰类型和恢复时间热带山地雨林群落的变化规律。应用PCA分析了各功能特性因子之间的相关关系，并应用TWINSPAN综合多个功能特性因子将热带山地雨林中的植物物种划分为13类功能群，最后应用DCCA排序阐述了各功能群随不同环境因子、干扰类型和恢复时间的变化规律，最后应用不同指标确定了各功能群的生态关键种。通过研究得出以下主要结论：
1．海南岛的热带山地雨林按照其立地条件，可大致划分为坡地山地雨林和沟谷山地雨林两大类，按照其受干扰的情况又可划分为老龄林和次生林(主要是由于采伐和刀耕火种形成的)等类型。海南岛热带山地雨林中乔木和灌木的相对多度最高，其次是草本和藤本植物，而附生和棕榈植物很低。乔木的相对多度与土壤有机质和离老龄林的距离成正相关，在采伐后恢复的山地雨林次生林中的分布最高。灌木的相对多度与生境的石砾含量成负相关，在沟谷山地雨林老龄林中的分布最低。草本、棕榈和附生植物的相对多度与生境的湿度、土壤石砾含量、pH、全P和K正相关，主要分布于沟谷山地雨林老龄林中，且随着恢复时间的增加而增加。藤本植物在热带不同类型山地雨林中的相对多度分布差异不明显。
2．耐荫性功能群在热带山地雨林中的相对生物量和相对多度最高，其中强耐荫性的功能群主要分布于沟谷山地雨林老龄林，中性或偏荫性功能群则主要分布于坡地山地雨林老龄林。喜光性功能群在山地雨林次生林中的相对生物量和相对多度最高，尤其是刀耕火种后恢复起来的次生林。随着刀耕火种后恢复起来的次生林恢复时间的增加，强耐荫性功能群的相对生物量和相对多度增加，而喜光性功能群则逐渐减少。强耐荫性功能群的相对生物量与恢复时间、土壤石砾含量、pH、全P和速效K成正相关，与海拔高度和土壤有机质成负相关性，而喜光性功能群的相对生物量与采伐强度成正相关性，与土壤速效N和全N成负相关性。
3．潜在高度中等(15-25m)的功能群在热带山地雨林中有最高的相对生物量和相对多度，潜在高度较高的功能群主要分布于沟谷山地雨林老龄林，其次是坡地山地雨林老龄林。随着刀耕火种后山地雨林次尘林的恢复时间增加，高冠层功能群的相对生物量和多度增加，其它功能群发生的变化不明显。高冠层功能群的相对生物量与土壤的石砾含量、pH、全P和速效K成正相关，与土壤有机质负相关。
4．大板根功能群在林冠层有最高的相对生物量和相对多度，尤其是沟谷山地雨林老龄林的林冠层。无板根的功能群分布较广，以山地雨林次生林最高。随着刀耕火种后山地雨林次生林的恢复时间增加，大板根功能群的相对生物量和相对多度增加，而小板根功能群减少。大板根功能群的相对生物量与干扰后的恢复时间、石砾含量、土壤pH、全P和速效K含量成正相关，而与海拔高度、土壤速效P和有机质成负相关。
5．小木材密度功能群在干扰后处于不同恢复阶段的山地雨林次生林中有最高的相对生物量和相对多度，其次是沟谷山地雨林老龄林。随着刀耕火种后山地雨林次生林的恢复时间增加，小木材密度功能群的相对生物量和相对多度减少，而大材密度的功能群增加。大木材密度功能群的相对生物量与海拔高度和土壤有机质成正相关，与采伐强度、恢复时间、土壤石砾含量和pH成负相关。
6．小种子功能群在刀耕火种后恢复起来的山地雨林次生林和沟谷山地雨林老龄林有最高的相对生物量和相对多度，而大种子的功能群主要分布于采伐后恢复起来的山地雨林次生林和坡地山地雨林老龄林，水分差异对种子大小的影响远大于干扰类型和恢复时间。随着刀耕火种后山地雨林次生林的恢复时间增加，小种子功能群的相对生物量和相对多度减少，而大种子功能群增加。
7．动物传播种子的功能群在热带山地雨林中的相对生物量和相对多度较高，尤其是坡地山地雨林老龄林最高。自体和风传播种子功能群的相对生物量和相对多度极低，其中自体传播种子的功能群主要分布于沟谷山地雨林老龄林或刀耕火种后恢复的山地雨林次生林，而风传播种子的功能群主要分布于采伐后恢复的山地雨林次生林。随着刀耕火种后山地雨林次生林的恢复时间增加，自体传播种子功能群的相对生物量和相对多度增加，而风传播种子功能群减少。动物传播种子功能群的相对生物量与采伐强度和土壤速效K成负相关，而自体传播种子功能群的相对生物量与土壤石砾含量和全P成正相关，风传播种子功能群的相对生物量与采伐强度成正相关。
8，常绿性功能群在热带不同山地雨林中有最高的相对生物量和相对多度，而落叶性功能群的分布极低且主要分布于山地雨林次生林中。随着刀耕火种后山地雨林次生林的恢复时间增加，常绿性功能群的相对生物量和相对多度增加，而落叶性功能群减少。常绿性功能群的相对生物量与海拔高度、坡向、土壤速效P和土壤有机质成正相关，与土壤pH和速效K成负相关，而落叶性功能群的相对生物量与采伐强度成正相关。
9．综合考虑物种的7个功能特性，可将热带山地雨林的所有物种划分为13类功能群，主要贡献因子是潜在高度、板根大小、种子大和木材密度，其次是物种的喜光性和落叶／常绿性，而种子传播方式最低。其中，潜在高度与种子生物量和板根大小之间成正相关；种子生物量与喜光性、自体和风传播方式成负相关，与木材密度和动物传播方式成正相关。此外，喜光性、木材密度、种子生物量和种子传播方式均与物种的落叶与常绿性相关。
10．沟谷比坡地山地雨林老龄林的功能群有更大的潜在高度和板根，且含有一定数量依赖于风或自体传播种子的落叶性物种，而坡地比沟谷山地雨林老龄林的功能群有更大的种子和木材密度，主要由动物传播种子的常绿性物种组成．随着刀耕火种后山地雨林次生林的恢复时间增加，功能群的潜在高度、种子生物量及木材密度增加，且由依赖于风传播种子的落叶性物种过度到动物传播种子的常绿性物种。
11．林冠层比其下层林的功能群有更高的潜在高度、板根、种子生物量和木材密度，可有效地指示生境的N、P、K、pH、水分和海拔高度的梯度变化，而低冠层或下层林较大的板根或种子生物量的功能群也能够在一定程度上指示生境的变化，但明显弱于冠层林的功能群。
12．应用物种的相对生物量、相对胸高断面积、相对多度、相对频度和重要值等5种方法，分别确定了各功能群的关键种。除了受种源影响较大的少数物种(如广东胡椒和毛荔枝)外，各关键种随不同干扰类型及恢复时间的变化趋势，均与对应的功能群相类似，说明了本文确定的关键种对各功能群较强的指示性。
The tropical rain forests have complicated structures, contain rich species and play an important role in conservation of global biodiversity and maintaining universal ecosystem functions. A classification of diverse species in tropical forests based on their different functional traits will greatly contribute to the scientific study and management for tropical forests. In this study, field investigations were conducted in totally 15 ha sample plots of recovery (second-growth) and old-growth forest vegetation in the Bawangling forest region of Hainan Island, South China. The recovery vegetation included different stages of fallows on the shifting cultivation secondary forest stands and logged forests under different logging systems. Based on the field investigations and each functional trait (including life form, potential height, buttress size, wood density, deciduous versus evergreen, seed mass and dispersal agents), plant species in the tropical montane rain forest were classified into different functional groups respectively. NMS (nommetric multidimensional scaling)was used to analyze their change with different environmental factors, disturbance types and recovery time. Then the relationships between different functional traits were analyzed by PCA (principal component analysis). Focusing on all these functional traits, 13 functional groups were identified by TWINSPAN (two-way indicator species analysis), and their change with different environmental factors, disturbance types and recovery time was ordinated by DCCA(detrended canonical correspondence analysis). At last, ecological keystone species in each functional group were identified from five aspects. The main results were as follows: Based on landform, the tropical montane rain forest can be classified into two forest stand types including mountain slope forest stands and ravine forest stands. Based on disturbance types, the tropical montane rain forest can be classified into old-growth forest stands and secondary forest stands (mainly including the logging and shifting cultivation).Trees and shrubs were the most abundant components of tropical montane rain forest, secondarily grasses and lianas, while epiphytes and palms were least in relative abundance. Relative abundance of trees was positively correlated with soil organic matter and the distance to old-growth forest, which had the highest distribution in the secondary forest stands. Relative abundance of shrubs was negatively correlated with stone content, which had high distributions in each forest stands, except for ravine old-growth forest stands. Relative abundance of the life forms such as grasses, palms and epiphytes had positively correlation with soil moisture content, soil stone content, pH, total P and available K, which had more abundance in the ravine old-growth forest stands than in mountain slope old-growth forest stands. Relative abundances of lianas weren't correlated with any environmental factors, which had no difference distribution in different forest stands.
1. Shade-tolerant functional groups had the most relative biomass and abundance in each forest stands. Among them, strong shade-tolerant functional groups were mainly distributed in the ravine old-growth forest stands, while less shade-tolerant functional groups were mainly in mountain slope old-growth forest stands. Shade-intolerant functional groups were mainly distributed in secondary forest stands, especially in recovery secondary forest stands after shifting cultivation. With the increasing of recovery time after shifting cultivation, shade-tolerant functional groups increased in relative biomass and abundance, while shade-intolerant functional groups decreased. Relative biomass of stong shade-tolerant functional groups was positively correlated with recovery time, stone content, pH, total P and available K, and negatively with elevation and soil organic matter, while those of shade-intolerant functional groups were negatively correlated with soil available N and total N.
2, Functional groups with moderate height (15-25m) had the most relative biomass and abundance in each forest stands, while those with maximal height were mainly distributed in the ravine old-growth forest stands, secondarily in mountain slope old-growth forest stands. With the increasing of recovery time after shifting cultivation; functional groups in the canopy layer of forest stands increased in relative biomass and abundance, while other functional groups had no significant chance. Relative biomass of functional groups in the canopy layer of forest stands were positively correlated with soil stone content, pH, total P, available K and recovery time, and negatively with soil organic matter.
3. Functional groups with larger buttress had the most relative biomass and abundace in the canopy layer of forest satnds, especially in the canopy layer of ravine old-growth forest stands, while those with smaller buttress were mainly in secondary forest stands. With the increasing of recovery time after shifting cultivation, functional groups with larger buttress increased in relative biomass and abundance, while smaller buttress decreased. Relative biomass of the functional groups with the largest buttress size was positively correlated with recovery time, stone content, soil pH, total P and available K, and negatively correlated with elevation, soil P and soil organic matter.
4. Functional groups with lighter wood density had the most relative biomass and abundace in secondary forest stands, secondarily in ravine old-growth forest stands, while those with heavier wood density were in mountain slope old-growth forest stands. With the increasing of recovery time after shifting cultivation, functional groups with lighter wood density decreased in relative biomass and abundance, while those with heavier wood density increased. Relative biomass of functional groups with heaviest wood density was positively correlated with elevation and soil organic matter, and negatively correlated with logging intensity, recovery time, soil stone content and pH.
5. Functional groups with lighter seed mass had the most relative biomass and abundance in shifting cultivation secondary forest stands and ravine old-growth forest stands, while those with heavier seed mass appeared in logged secondary forest stands and mountain slope old-growth forest stands, which showed water difference had more influence on see mass than both disturbance types and recovery time. With the increasing of recovery time after Shifting cultivation, the functional groups with lighter seed mass decreased in relative biomass and abundance, while those with heavier ones increased.
6. Functional groups by animal dispersal had the most relative biomass and abundance in each forest stands, especially in old-growth forest stands, while functional groups by auto and wind dispersal had the less. Functional groups by auto dispersal were mainly distributed in ravine old-growth forest stands or recovery secondary forest stands after shifting cultivation, while those by wind dispersal mostly appeared in secondary forest stands. With the increasing of recovery time after shifting cultivation, the functional groups by atuo dispersal increased in relative biomass and abundance, while those by wind dispersal decreased. Relative biomass of functional groups by animal dispersal were negatively correlated with logging intensity and soil available K, while those by auto were positively correlated with soil stone content and total P, and those by wind dispersal were positively correlated with logging intensity.
7. Evergreen functional groups had the most relative biomass and abundance in each forest Stands, while deciduous functional groups had the least and mostly appeared in the secondary forest stands. With the increasing of recovery time after shifting cultivation, evergreen functional groups increased in relative biomass and abundance, while the deciduous ones decreased. Relative biomass of evergreen functional groups were positively correlated with elevation, aspect, soil available P and organic matter, and negatively with soil pH and available K, while those of deciduous functional groups were positively correlated with logging intensity.
8. Based on the 7 functional traits, major plant species in the tropical montane rain forests could be classified into 13 functional groups. The most important factors (traits) were potential height, buttress size and seed mass and wood density, secondarily shade-intolerant and defoliation/evergreen, while seed dispersal was least in importance. Among them, potential height was positively correlated with seed mass and buttress size (P＜0.05); and seed mass was negatively correlated with shade-intolerant, auto and wind dispersal, and positively correlated with wood density and animal dispersal (P<0.05); and the factors such as shade-intolerant, seed mass, wood density and dispersal agents were all singificantly correlated with defoliation/evergreen (P<0.05).
9. The functional groups in ravine old-growth forest stands, partially composed of deciduous species by auto or wind dispersal, had higher potential height and larger buttress than those in mountain slope ones, while those in mountain slope old-growth forest stands, mainly composed of evergreen species by animal dispersal, had heavier seed mass and wood density. With the increasing of recovery time, the potential height, seed mass and wood density of functional groups increased, and deciduous species by wind dispersal changed to evergreen species by animal dispersal.
10. Functional groups in the canopy layer of forest stands had higher potential height, larger buttress size, heavier seed mass and wood density than those in the lower layer, which can greatly reflect the gradient change of environmental factors including soilavailable N, available P, available K, pH, organic matter, moisture content and elevation. However, the functional groups in the lower canopy or understorey layer, which had smaller buttress or lighter seed mass, had weak indication to environmental changes.
11. To identify the ecological keystone species, five methods were adopted including relative biomass, relative basal area, relative abundance, relative frequency and importance value. The results showed that the ecological keystone species in each functional group could well resemble their repective functional groups' change with environmental and disturbance characteristics, except for a few species that were greatly influenced by seed sources. The ecological keystone species identified in this study can well represent the main characteristics of their functional groups,
1．海南岛的热带山地雨林按照其立地条件，可大致划分为坡地山地雨林和沟谷山地雨林两大类，按照其受干扰的情况又可划分为老龄林和次生林(主要是由于采伐和刀耕火种形成的)等类型。海南岛热带山地雨林中乔木和灌木的相对多度最高，其次是草本和藤本植物，而附生和棕榈植物很低。乔木的相对多度与土壤有机质和离老龄林的距离成正相关，在采伐后恢复的山地雨林次生林中的分布最高。灌木的相对多度与生境的石砾含量成负相关，在沟谷山地雨林老龄林中的分布最低。草本、棕榈和附生植物的相对多度与生境的湿度、土壤石砾含量、pH、全P和K正相关，主要分布于沟谷山地雨林老龄林中，且随着恢复时间的增加而增加。藤本植物在热带不同类型山地雨林中的相对多度分布差异不明显。
2．耐荫性功能群在热带山地雨林中的相对生物量和相对多度最高，其中强耐荫性的功能群主要分布于沟谷山地雨林老龄林，中性或偏荫性功能群则主要分布于坡地山地雨林老龄林。喜光性功能群在山地雨林次生林中的相对生物量和相对多度最高，尤其是刀耕火种后恢复起来的次生林。随着刀耕火种后恢复起来的次生林恢复时间的增加，强耐荫性功能群的相对生物量和相对多度增加，而喜光性功能群则逐渐减少。强耐荫性功能群的相对生物量与恢复时间、土壤石砾含量、pH、全P和速效K成正相关，与海拔高度和土壤有机质成负相关性，而喜光性功能群的相对生物量与采伐强度成正相关性，与土壤速效N和全N成负相关性。
3．潜在高度中等(15-25m)的功能群在热带山地雨林中有最高的相对生物量和相对多度，潜在高度较高的功能群主要分布于沟谷山地雨林老龄林，其次是坡地山地雨林老龄林。随着刀耕火种后山地雨林次尘林的恢复时间增加，高冠层功能群的相对生物量和多度增加，其它功能群发生的变化不明显。高冠层功能群的相对生物量与土壤的石砾含量、pH、全P和速效K成正相关，与土壤有机质负相关。
4．大板根功能群在林冠层有最高的相对生物量和相对多度，尤其是沟谷山地雨林老龄林的林冠层。无板根的功能群分布较广，以山地雨林次生林最高。随着刀耕火种后山地雨林次生林的恢复时间增加，大板根功能群的相对生物量和相对多度增加，而小板根功能群减少。大板根功能群的相对生物量与干扰后的恢复时间、石砾含量、土壤pH、全P和速效K含量成正相关，而与海拔高度、土壤速效P和有机质成负相关。
5．小木材密度功能群在干扰后处于不同恢复阶段的山地雨林次生林中有最高的相对生物量和相对多度，其次是沟谷山地雨林老龄林。随着刀耕火种后山地雨林次生林的恢复时间增加，小木材密度功能群的相对生物量和相对多度减少，而大材密度的功能群增加。大木材密度功能群的相对生物量与海拔高度和土壤有机质成正相关，与采伐强度、恢复时间、土壤石砾含量和pH成负相关。
6．小种子功能群在刀耕火种后恢复起来的山地雨林次生林和沟谷山地雨林老龄林有最高的相对生物量和相对多度，而大种子的功能群主要分布于采伐后恢复起来的山地雨林次生林和坡地山地雨林老龄林，水分差异对种子大小的影响远大于干扰类型和恢复时间。随着刀耕火种后山地雨林次生林的恢复时间增加，小种子功能群的相对生物量和相对多度减少，而大种子功能群增加。
7．动物传播种子的功能群在热带山地雨林中的相对生物量和相对多度较高，尤其是坡地山地雨林老龄林最高。自体和风传播种子功能群的相对生物量和相对多度极低，其中自体传播种子的功能群主要分布于沟谷山地雨林老龄林或刀耕火种后恢复的山地雨林次生林，而风传播种子的功能群主要分布于采伐后恢复的山地雨林次生林。随着刀耕火种后山地雨林次生林的恢复时间增加，自体传播种子功能群的相对生物量和相对多度增加，而风传播种子功能群减少。动物传播种子功能群的相对生物量与采伐强度和土壤速效K成负相关，而自体传播种子功能群的相对生物量与土壤石砾含量和全P成正相关，风传播种子功能群的相对生物量与采伐强度成正相关。
8，常绿性功能群在热带不同山地雨林中有最高的相对生物量和相对多度，而落叶性功能群的分布极低且主要分布于山地雨林次生林中。随着刀耕火种后山地雨林次生林的恢复时间增加，常绿性功能群的相对生物量和相对多度增加，而落叶性功能群减少。常绿性功能群的相对生物量与海拔高度、坡向、土壤速效P和土壤有机质成正相关，与土壤pH和速效K成负相关，而落叶性功能群的相对生物量与采伐强度成正相关。
9．综合考虑物种的7个功能特性，可将热带山地雨林的所有物种划分为13类功能群，主要贡献因子是潜在高度、板根大小、种子大和木材密度，其次是物种的喜光性和落叶／常绿性，而种子传播方式最低。其中，潜在高度与种子生物量和板根大小之间成正相关；种子生物量与喜光性、自体和风传播方式成负相关，与木材密度和动物传播方式成正相关。此外，喜光性、木材密度、种子生物量和种子传播方式均与物种的落叶与常绿性相关。
10．沟谷比坡地山地雨林老龄林的功能群有更大的潜在高度和板根，且含有一定数量依赖于风或自体传播种子的落叶性物种，而坡地比沟谷山地雨林老龄林的功能群有更大的种子和木材密度，主要由动物传播种子的常绿性物种组成．随着刀耕火种后山地雨林次生林的恢复时间增加，功能群的潜在高度、种子生物量及木材密度增加，且由依赖于风传播种子的落叶性物种过度到动物传播种子的常绿性物种。
11．林冠层比其下层林的功能群有更高的潜在高度、板根、种子生物量和木材密度，可有效地指示生境的N、P、K、pH、水分和海拔高度的梯度变化，而低冠层或下层林较大的板根或种子生物量的功能群也能够在一定程度上指示生境的变化，但明显弱于冠层林的功能群。
12．应用物种的相对生物量、相对胸高断面积、相对多度、相对频度和重要值等5种方法，分别确定了各功能群的关键种。除了受种源影响较大的少数物种(如广东胡椒和毛荔枝)外，各关键种随不同干扰类型及恢复时间的变化趋势，均与对应的功能群相类似，说明了本文确定的关键种对各功能群较强的指示性。
The tropical rain forests have complicated structures, contain rich species and play an important role in conservation of global biodiversity and maintaining universal ecosystem functions. A classification of diverse species in tropical forests based on their different functional traits will greatly contribute to the scientific study and management for tropical forests. In this study, field investigations were conducted in totally 15 ha sample plots of recovery (second-growth) and old-growth forest vegetation in the Bawangling forest region of Hainan Island, South China. The recovery vegetation included different stages of fallows on the shifting cultivation secondary forest stands and logged forests under different logging systems. Based on the field investigations and each functional trait (including life form, potential height, buttress size, wood density, deciduous versus evergreen, seed mass and dispersal agents), plant species in the tropical montane rain forest were classified into different functional groups respectively. NMS (nommetric multidimensional scaling)was used to analyze their change with different environmental factors, disturbance types and recovery time. Then the relationships between different functional traits were analyzed by PCA (principal component analysis). Focusing on all these functional traits, 13 functional groups were identified by TWINSPAN (two-way indicator species analysis), and their change with different environmental factors, disturbance types and recovery time was ordinated by DCCA(detrended canonical correspondence analysis). At last, ecological keystone species in each functional group were identified from five aspects. The main results were as follows: Based on landform, the tropical montane rain forest can be classified into two forest stand types including mountain slope forest stands and ravine forest stands. Based on disturbance types, the tropical montane rain forest can be classified into old-growth forest stands and secondary forest stands (mainly including the logging and shifting cultivation).Trees and shrubs were the most abundant components of tropical montane rain forest, secondarily grasses and lianas, while epiphytes and palms were least in relative abundance. Relative abundance of trees was positively correlated with soil organic matter and the distance to old-growth forest, which had the highest distribution in the secondary forest stands. Relative abundance of shrubs was negatively correlated with stone content, which had high distributions in each forest stands, except for ravine old-growth forest stands. Relative abundance of the life forms such as grasses, palms and epiphytes had positively correlation with soil moisture content, soil stone content, pH, total P and available K, which had more abundance in the ravine old-growth forest stands than in mountain slope old-growth forest stands. Relative abundances of lianas weren't correlated with any environmental factors, which had no difference distribution in different forest stands.
1. Shade-tolerant functional groups had the most relative biomass and abundance in each forest stands. Among them, strong shade-tolerant functional groups were mainly distributed in the ravine old-growth forest stands, while less shade-tolerant functional groups were mainly in mountain slope old-growth forest stands. Shade-intolerant functional groups were mainly distributed in secondary forest stands, especially in recovery secondary forest stands after shifting cultivation. With the increasing of recovery time after shifting cultivation, shade-tolerant functional groups increased in relative biomass and abundance, while shade-intolerant functional groups decreased. Relative biomass of stong shade-tolerant functional groups was positively correlated with recovery time, stone content, pH, total P and available K, and negatively with elevation and soil organic matter, while those of shade-intolerant functional groups were negatively correlated with soil available N and total N.
2, Functional groups with moderate height (15-25m) had the most relative biomass and abundance in each forest stands, while those with maximal height were mainly distributed in the ravine old-growth forest stands, secondarily in mountain slope old-growth forest stands. With the increasing of recovery time after shifting cultivation; functional groups in the canopy layer of forest stands increased in relative biomass and abundance, while other functional groups had no significant chance. Relative biomass of functional groups in the canopy layer of forest stands were positively correlated with soil stone content, pH, total P, available K and recovery time, and negatively with soil organic matter.
3. Functional groups with larger buttress had the most relative biomass and abundace in the canopy layer of forest satnds, especially in the canopy layer of ravine old-growth forest stands, while those with smaller buttress were mainly in secondary forest stands. With the increasing of recovery time after shifting cultivation, functional groups with larger buttress increased in relative biomass and abundance, while smaller buttress decreased. Relative biomass of the functional groups with the largest buttress size was positively correlated with recovery time, stone content, soil pH, total P and available K, and negatively correlated with elevation, soil P and soil organic matter.
4. Functional groups with lighter wood density had the most relative biomass and abundace in secondary forest stands, secondarily in ravine old-growth forest stands, while those with heavier wood density were in mountain slope old-growth forest stands. With the increasing of recovery time after shifting cultivation, functional groups with lighter wood density decreased in relative biomass and abundance, while those with heavier wood density increased. Relative biomass of functional groups with heaviest wood density was positively correlated with elevation and soil organic matter, and negatively correlated with logging intensity, recovery time, soil stone content and pH.
5. Functional groups with lighter seed mass had the most relative biomass and abundance in shifting cultivation secondary forest stands and ravine old-growth forest stands, while those with heavier seed mass appeared in logged secondary forest stands and mountain slope old-growth forest stands, which showed water difference had more influence on see mass than both disturbance types and recovery time. With the increasing of recovery time after Shifting cultivation, the functional groups with lighter seed mass decreased in relative biomass and abundance, while those with heavier ones increased.
6. Functional groups by animal dispersal had the most relative biomass and abundance in each forest stands, especially in old-growth forest stands, while functional groups by auto and wind dispersal had the less. Functional groups by auto dispersal were mainly distributed in ravine old-growth forest stands or recovery secondary forest stands after shifting cultivation, while those by wind dispersal mostly appeared in secondary forest stands. With the increasing of recovery time after shifting cultivation, the functional groups by atuo dispersal increased in relative biomass and abundance, while those by wind dispersal decreased. Relative biomass of functional groups by animal dispersal were negatively correlated with logging intensity and soil available K, while those by auto were positively correlated with soil stone content and total P, and those by wind dispersal were positively correlated with logging intensity.
7. Evergreen functional groups had the most relative biomass and abundance in each forest Stands, while deciduous functional groups had the least and mostly appeared in the secondary forest stands. With the increasing of recovery time after shifting cultivation, evergreen functional groups increased in relative biomass and abundance, while the deciduous ones decreased. Relative biomass of evergreen functional groups were positively correlated with elevation, aspect, soil available P and organic matter, and negatively with soil pH and available K, while those of deciduous functional groups were positively correlated with logging intensity.
8. Based on the 7 functional traits, major plant species in the tropical montane rain forests could be classified into 13 functional groups. The most important factors (traits) were potential height, buttress size and seed mass and wood density, secondarily shade-intolerant and defoliation/evergreen, while seed dispersal was least in importance. Among them, potential height was positively correlated with seed mass and buttress size (P＜0.05); and seed mass was negatively correlated with shade-intolerant, auto and wind dispersal, and positively correlated with wood density and animal dispersal (P<0.05); and the factors such as shade-intolerant, seed mass, wood density and dispersal agents were all singificantly correlated with defoliation/evergreen (P<0.05).
9. The functional groups in ravine old-growth forest stands, partially composed of deciduous species by auto or wind dispersal, had higher potential height and larger buttress than those in mountain slope ones, while those in mountain slope old-growth forest stands, mainly composed of evergreen species by animal dispersal, had heavier seed mass and wood density. With the increasing of recovery time, the potential height, seed mass and wood density of functional groups increased, and deciduous species by wind dispersal changed to evergreen species by animal dispersal.
10. Functional groups in the canopy layer of forest stands had higher potential height, larger buttress size, heavier seed mass and wood density than those in the lower layer, which can greatly reflect the gradient change of environmental factors including soilavailable N, available P, available K, pH, organic matter, moisture content and elevation. However, the functional groups in the lower canopy or understorey layer, which had smaller buttress or lighter seed mass, had weak indication to environmental changes.
11. To identify the ecological keystone species, five methods were adopted including relative biomass, relative basal area, relative abundance, relative frequency and importance value. The results showed that the ecological keystone species in each functional group could well resemble their repective functional groups' change with environmental and disturbance characteristics, except for a few species that were greatly influenced by seed sources. The ecological keystone species identified in this study can well represent the main characteristics of their functional groups,
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