海南岛热带云雾林群落结构及组配机制研究
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摘要
热带森林是地球上物种最丰富且结构最复杂的陆地生态系统,其物种共存机制是近年来群落生态学研究的一个重要内容。相对于较低海拔的热带林,物种共存机制的研究在高海拔的热带云雾林群落中还几乎没有开展。热带云雾林的环境条件和物种组成与较低海拔的热带森林明显不同,如云雾出现频率较高、风多且强烈、气温偏低、空气湿度较大;树木高度和径级明显减小,树干常弯曲但多缺乏板根,叶片革质且面积较小,附生植物丰富等。因此,热带云雾林的群落结构与组配机制可能有其独特性。热带云雾林相对矮小的树木使得精确测定其功能性状成为可能。海南岛的热带云雾林主要分布在霸王岭、尖峰岭、五指山和鹦哥岭等林区海拔1200 m以上范围,包括热带山地常绿林和热带山顶矮林两种群落类型。本文以海南岛霸王岭热带山地常绿林和热带山顶矮林为对象,通过典型样方调查、环境因子观测和主要功能性状测定,系统比较了两种森林类型的环境因子、群落结构和物种多样性特征,分析了物种共有度、空间格局和竞争作用规律及其在群落组配中的作用;以比叶面积和树种高度这两个功能性状为基础,采用数量比较和模型检验方法,分析了功能性状在物种内、物种间和样地间的变化格局、尺度效应及环境筛驱动机制,讨论了随机与生态位过程在群落组配中的作用。主要研究结果如下:
(一)、热带云雾林群落的环境和结构特征研究
1.热带山地常绿林和热带山顶矮林一天中光合有效辐射呈单峰型曲线变化,热带山地常绿林各时段的光合有效辐射比热带山顶矮林显著低;湿季(5～10月)两群落类型日平均气温分别为(21.76±2.44)℃和(19.33±1.03)℃,且随时间呈单峰型曲线变化,热带山地常绿林日平均气温比热带山顶矮林显著高;湿季两群落类型日平均空气相对湿度分别为(88.44±2.90)%和(97.71±0.80)%,且随时间呈“倒S型”曲线变化,热带山地常绿林各月日平均空气相对湿度比热带山顶矮林显著小;与热带山顶矮林相比,热带山地常绿林的土壤全氮、全磷、速效氮、有机质、pH和土壤厚度显著大而全钾和有效磷含量显著低;热带山地常绿林的坡度、地表岩石裸露比例和海拔高度都比热带山顶矮林显著小;主成分和相关性分析表明:空气温度、土壤有效磷、全钾、全氮及地形因子是热带云雾林的主导环境因子。
2.热带山地常绿林的优势种为线枝蒲桃(Syzygium araiocladum)、蚊母树(Distylium racemosum)、碟斗青冈(Cyclobalanopsis disciformis)、九节(Psychotria rubra)和碎叶蒲桃(S. buxifolium)等,热带山顶矮林的优势种为蚊母树、碎叶蒲桃、黄杞(Engelhardtia roxburghiana)、九节和光叶山矾(S. lancifolia)等;两森林类型的优势科都为樟科(Lauraceae)、山矾科(Symplocaceae)、茜草科(Rubiaceae)、壳斗科(Fagaceae)和木犀科(Oleaceae),优势属都为山矾属(Symplocos)、青冈属(Cyclobalanopsis)、柯属(Lithocarpus)和琼楠属(Beilschmiedia)。两森林类型间的S?rensen相似性指数为0.71。热带山地常绿林幼树(1 cm≤dbh < 5 cm)和小树(5 cm≤dbh < 10 cm)的平均密度比热带山顶矮林显著小,而成年树(dbh≥10 cm)平均密度无显著差异;前者的小树和成年树的平均胸径比后者显著大,而幼树平均胸径比后者显著小;前者的幼树、小树和成年树的平均高度比后者显著大。
(二)、基于物种物种多样性的热带云雾林群落组配机制研究
3.热带山地常绿林群落的物种丰富度观测值、一阶刀切法指数、二阶刀切法指数和抽样法物种丰富度指数都比热带山顶矮林显著高,但群落多度比热带山顶矮林显著低。应用幂律模型、指数模型和逻辑斯蒂模型对热带山地常绿林和热带山顶矮林的物种―面积关系进行拟合,发现逻辑斯蒂模型是两森林类型中物种―面积关系的最优模型。应用分割线段模型、生态为优先模型、Zipf模型、Zipf-Mandelbrot模型和中性理论模型对热带山地常绿林和热带山顶矮林种―多度分布进行拟合,发现Zipf-Mandelbrot模型是热带山地常绿林种―多度分布的最优模型,而生态位优先模型和Zipf-Mandelbrot模型是热带山顶矮林种―多度分布的最优模型。广义线性模型分析表明光合有效辐射、空气温度、土壤氮和磷、坡度等环境筛对群落的物种丰富度及多度有显著影响。
4.应用4种共有度指数(the number of checkerboard species pairs、C score、种间联结数和V ratio)研究热带云雾林物种在5 m×5 m、10 m×10 m、20 m×20 m和30 m×30 m样方尺度上的共有度(co-occurrence)格局。与随机期望值相比,热带山地常绿林群落的所有物种、不同多度及径级物种在5 m和10 m样方尺度上呈现较少的共有度格局,且该格局在5 m样方尺度上最明显。显示物种间显著地非随机分离,说明物种间竞争作用于群落组配,且该作用在较小尺度上最强。热带山顶矮林群落的所有物种、不同多度及径级物种在20 m和30 m样方尺度上均呈现较多的共有度格局,且该格局在20 m尺度上最明显。显示物种间显著地非随机聚集,说明正相互作用(如促进作用facilitation)作用于群落组配,且该作用在较大尺度上最强。
5.应用Donnelly最近邻体指数分析物种分布类型,发现热带山地常绿林和热带山顶矮林不同分布类型物种百分数的大小顺序为规则分布>随机分布>聚集分布;随径级增大,聚集分布和随机分布的物种百分数呈减小趋势,而规则分布的物种百分数呈增大趋势。两森林类型中共同出现的优势物种蚊母树和碎叶蒲桃的幼树呈聚集分布,小树呈聚集分布或随机分布,而大树基本上呈随机分布。应用单变量和双变量O-ring函数分析热带山地常绿林和热带山顶矮林中碎叶蒲桃和蚊母树的不同径级种群的空间格局及其相互关联,发现蚊母树的幼苗和小树主要在小于10 m样方尺度上聚集分布,而碎叶蒲桃的幼树和小树在大于23 m样方尺度上聚集分布,不同物种幼树的空间分布与种子扩散方式有关。蚊母树的小树与幼树、大树与幼树及大树与小树的空间格局主要在小于5 m样方尺度上正关联;碎叶蒲桃的小树与幼树、大树与小树的空间格局负关联。
6.与零假设模型比较,热带山地常绿林和热带山顶矮林分别有(41±4)%和(37±2)%的物种与不同物种的个体间、(43±4)%和(38±7)%的物种与相同物种的个体间、(28±2)%和(44±5)%的物种与所有物种的个体间呈现显著的胸径―最近邻体距离相关关系,说明非随机过程在群落组配中有重要作用;两森林类型分别有(23±3)%和(26±5)%的不同物种的个体间、(27±5)%和(19±9)%的相同物种的个体间、(17±8)%和(27±7)%的所有物种的个体间呈现显著的胸径―最近邻体距离正相关关系,说明了竞争作用驱动群落组配;物种间大小―距离的显著负相关关系反映了促进作用驱动群落组配;热带山地常绿林中物种竞争作用的重要性和强度比热带山顶矮林小,竞争作用的重要性和强度随土壤肥力减小而增大。
(三)、基于植物功能性状的热带云雾林群落组配机制研究
7、热带山地常绿林群落中多度加权比叶面积平均值和多度加权树木高度平均值比热带山顶矮林显著大,而比叶面积和高度可塑性比热带山顶矮林显著小;多元逐步回归表明比叶面积大小及其可塑性与土壤全磷及空气温度显著相关,高度大小仅与空气温度显著相关,高度可塑性与上述两个环境因子都显著相关。说明了空气温度和土壤磷等环境筛作用于热带云雾林物种功能性状变化,从而影响群落物种组成和结构。8.以热带山顶矮林为例分析物种功能性状变化与群落组配的关系。热带山顶矮林中三个地点(松林顶、雅加松顶和斧头岭)的种内比叶面积和种间比叶面积随光照强度增大而减小。种内比叶面积和种间比叶面积在三个地点间有显著差异,广义混合模型分析表明样地间比叶面积差异仅与空气温度显著相关。alpha比叶面积值随群落内光照强度增大而减小,beta比叶面积值在不同样地间有显著差异。说明热带山顶矮林通过群落内光照和群落间空气温度两种环境筛作用,使物种按照功能性状变化在群落内和群落间进行组配。
9.以热带山顶矮林为例,以功能性状为基础在4个样方尺度(5 m×5 m、10 m×10 m、20 m×20 m和30 m×30 m)上对群落组配进行零假设检验。与零假设模型比较,热带山顶矮林中比叶面积小的物种在20 m和30 m样方尺度上有生存优势(over-representation),而高度大的物种在4个样方尺度上都有生存优势。线性回归表明比叶面积平均值的观测值与期望值的效应大小(effect size)仅与日平均气温及日平均最高气温显著正相关,最大物种高度平均值的观测值与期望值的效应大小仅与光合有效辐射正相关。说明热带山顶矮林物种根据比叶面积和高度大小变化进行非随机组配,此组配过程主要由空气温度和物种对光照的竞争作用驱动。
10.综合本文的研究结果可以看出:低温和低磷等环境因子对不同功能性状的物种具有筛选作用,影响其在群落中的分布与多度;在同一群落内促进作用和物种对光照等环境因子的竞争作用又影响着物种的空间配置与个体比例。因此,热带云雾林的群落结构主要由非随机的生态位过程组配形成。
Tropical forests are terrestrial ecosystems harboring the most abundant species and complicated structures, and play important roles in the regional and global biodiversity conservation and ecological function maintenance. Studies on the mechanisms of species coexistence in these ecosystems have been a vitally important topic in community ecology in recent years. Although some studies have been carried out in the low altitudinal tropical forests, species assembly and community structuring in the high altitudinal tropical cloud forests are poorly understood. Compared with the lower altitude tropical forests, environmental conditions in tropical cloud forests are quite different, characterized by frequent fog, low temperature, high humidity and strong winds. Moreover, trees in these forests are typically more deformed and elfin with small sturdy leaves, few buttress,and thick cover of epiphytes. These special environmental and physiognomical features may confine a unique rule of community assembly. The small stature of trees in the tropical cloud forests make it possible to precisely measure some important functional traits (such as specific leaf area and maximum species height). Tropical cloud forests in Hainan Island are distributed at more than 1200 m altitude in Bawangling Mt., Jianfengling Mt., Wuzhishan Mt., Yinggeling Mt., etc, including tropical montane evergreen forest (TMEF) and tropical (montane) dwarf forest (TDF or TMDF). To explore the structuring and assembly rules of tropical cloud forest communities, we investigated the species diversity and environmental conditions, and measured the specific leaf area (SLA), dbh and height for 5765 individuals trees and shrubs (dbh≥1 cm) in TMEF and TMDF in Bawangling National Natural Reserve, Hainan island, South China. Then, we compared the environmetal conditions, community stucture and assembly rules for the two cloud forest types. The main results are as follows.
(Ⅰ) Environmental conditions and community features of the tropical cloud forests
1. Daily photosynthetically active radiation (PAR) showed a unimodal curve both in TMEF and TMDF, but PAR was significantly lower in TMEF than TMDF. From May to October, mean daily air temperature differed significantly between TMEF and TMDF and showed a unimodal curve in the two forests, with average values of (21.76±2.44)°C and (19.33±1.03)°C, respectively. Additionally, mean daily relative humidity differed significantly between TMEF and TMDF and showed an“inverse S”curve; average values were (88.44±2.90) % and (97.71±0.80) %, respectively. TMEF had higher total nitrogen, total phosphorous, available nitrogen, organic matter, pH and soil thickness, but lower total potassium and available phosphorous than TMDF. Slope, cover of exposed rock and altitude were lower in TMEF than TMDF. Principal component analysis and Pearson’s correlation analysis indicated that air temperature, soil phosphorous, potassium, nitrogen and the three topographic factors were the most important predictors of distribution of these tropical cloud forests.
2. The dominant species in TMEF were Syzygium araiocladum, Distylium racemosum, Cyclobalanopsis disciformis, Psychotria rubra and S. buxifolium, and the dominant species in TMDF were D. racemosum, S. buxifolium, Engelhardtia roxburghiana, P. rubra and S. lancifolia. The common dominant families for the two forest types were Lauraceae, Symplocaceae, Rubiaceae, Fagaceae and Oleaceae. The common dominant genera for the two forest types were Symplocos, Cyclobalanopsis, Lithocarpus and Beilschmiedia. The S?rensen species similarity index for the two forest types was 0.71. The mean stem density for saplings (1 cm≤dbh﹤5 cm) and small trees (5 cm≤dbh﹤10 cm) was significantly lower in TMEF than TMDF, while there were no differences in mean stem density for adult trees (10 cm≤dbh) between these two forest types. Mean dbh for small trees and adult trees were significantly higher in TMEF than TMDF, while mean dbh for saplings were significantly lower in TMEF than TMDF. Mean plant height for saplings, small trees and adult trees were significantly higher in TMEF than TMDF.
(Ⅱ) Community assembly based on species diversity
3. The observed species richness values, as well as the species richness values predicted by 1st order Jackknife estimator, 2nd order Jackknife estimator and bootstrap estimator, were significantly higher in TMEF than TMDF; while the individual abundance was significantly lower in TMEF than TMDF. Compared with power curve and exponential curve, logistic curve was the optimal model simulating the species-area relation for the two forest types. After Brokenstick model, Niche preemption model, Zipf model, Zipf-Mandelbrot model and Neutral theory model were used to simulate the species-abundance distribution for TMEF community and TMDF community, it was found that Zipf-Mandelbrot model was the optimal model predicting the species-abundance distribution in TMEF, while Niche preemption model and Zipf-Mandelbrot model were the optimal models predicting the species-abundance distribution in TMDF. A general linear anaysis model revealed that PAR, air temperature, soil nitrogen and phosphorus were the major environmental filters predicting the species richness and individual abundance between the two forest communities..
4. Patterns of species co-occurrence in TMEF and TMDF were assessed at 5 m×5 m, 10 m×10 m, 20 m×20 m and 30 m×30 m plot sizes using four species co-occurrence indices, including the number of checkerboard species pairs, the checkerboard score of matrix, the number of species combinations and the variance ratio. All combined species, species with different abundance classes and species with different dbh classes in TMEF all showed less co-occurrence patterns than null model tests at 5 m and 10 m plot sizes, indicating that species were non-randomly segregated, and competition probably impacted on the community assembly in this forest. The species co-occurrence indices were the highest at 5 m plot size. In contrast, all combined species, species with different abundance classes and species with different dbh classes in TMDF all showed more co-occurrence patterns than null model tests at 20 m and 30 m plot sizes, indicating that species were non-randomly aggregated, and facilitation probably impacted on the community assembly in this forest. The species co-occurrence indcies were the highest at 20 m plot size.
5. Patterns of species distribution in tropical cloud forests were assessed using Donnelly nearest neighbor distance index. Percentage of species in regular distribution was higher than that of random distribution and clumped distribution, while percentage of species in clumped distribution was the lowest; percentage of species in random distribution and clumped distribution decreased, while percengate of species in regular distribution increased with increasing dbh. Patterns of distribution of saplings for the common dominant species, S. buxifolium and D. racemosum in both TMEF and TMDF, showed a clumped distribution, their small tress showed a clumped or random distribution, and their adult trees showed a random distribution. A univariate O-ring function analysis revealed that saplings and small trees for D. racemosum were aggregated at less than 10 m plot size, while saplings and small trees for S. buxifolium were aggregated at more than 23 m plot size. A bivariate O-ring function analysis showed that the spatial patterns between small trees and saplings, adult trees and saplings, and adult trees and small trees for D. racemosum were positively associated at less than 5 m plot size. On the contrary, the spatial patterns between small trees and saplings, and adult trees and small trees for S. buxifolium were negatively associated.
6. Based on null model tests, (41±4) % and (37±2) % heterospecific trees, (43±4) % and (38±7) % conspecific trees, and (28±2) % and (44±5) % all combined tree species in TMEF and TMDF, showed significant correlations between dbh and nearest neighbor distance, indicating that non-random processes affected on community assembly in tropical cloud forests. (23±3) % and (26±5) % heterospecific trees, (27±5) % and (19±9) % conspecific trees, (17±8) % and (27±7) % all combined tree species in TMEF and TMDF, showed significantly positive correlations between dbh and nearest neighbor distance, indicating that competition impacted on community assembly in the tropical cloud forests. However, negative correlations between dbh and nearest neighbor distance based on null model tests showed that facilitation affected community assembly in the tropical cloud forests. Both importance and intensity of competition statistically increased with decreasing forest productivity from TMEF to TMDF for all the shared species spanning these two forests. (Ⅲ) Community assembly based on functional traits
7. Both abundance-weighted mean SLA and plant height were significantly higher in TMEF than TMDF, while phenotypic plasticity in SLA and height was significantly lower in TMEF than TMDF. Multiple linear regression analyses indicated that among the measured environmental factors, both air temperature and soil total phosphorus were significantly correlated with mean SLA and its plasticity, and only air temperature was correlated with mean height. But both air temperature and soil total phosphorus was significantly correlated with the plasticity index of plant height. These results indicate that air temperature and soil phosphorus acted as environmental filters on decreasing SLA and plant height, while increasing the plasticity of the functional traits from TMEF to TMDF, and thus impact on species composition and community structure in tropical cloud forests.
8. SLA decreased significantly with increasing solar irradiance within the three study sites in TMDF, and differed significantly among the three sites both for within and among species comparisons. Mean plot SLA, accounting for both within and among species across the three sites, increased significantly in relation to air temperature but not local PAR and soil total phosphorus. Alpha SLA decreased significantly with increasing solar irradiance within the three sites and beta SLA differed significantly among the three sites. The strong relationship between both intra- and interspecific variation in SLA and environmental conditions strongly confirms the role of trait variation in the assembly of plant species into tropical cloud forest communities via environment filtering related to light availability and air temperature.
9. Community assembly were examined based on null model tests at 5×5 m, 10×10 m, 20×20 m and 30×30 m plot sizes using a trait-based approach in TMDF. Lower SLA growing forest plants were over-represented within forest communities at 20 m and 30 m plot sizes, and taller growing forest plants were over-represented within forest communities at the four plot sizes. The correlation between effect size of the test statistic (i.e. mean trait value) of null model tests for specific leaf area and air temperature was positive, and that for maximum species height and percentage reduction of PAR was positive, revealing that there is a stress-tolerator advantage for lower specific leaf area species to be adapted to low temperature, and a size-advantage for taller species competing for light. Thus species are non-randomly assembled with respect to both specific leaf area and maximum species height in the tropical montane dwarf forest community, and that these processes are driven by both the temperature stress and biotic competition.
10. In conclusion, the low air temperature and low soil phosphorus act as environmental filters on species with different functional traits, affecting their distribution and abundance in tropical cloud forests; meanwhile, both competition and facilitation processes affect the spatial patterning and proportions of individual abundance within a community. The tropical cloud forest communities are mainly assembled by niche-based non-random processes.
(一)、热带云雾林群落的环境和结构特征研究
1.热带山地常绿林和热带山顶矮林一天中光合有效辐射呈单峰型曲线变化,热带山地常绿林各时段的光合有效辐射比热带山顶矮林显著低;湿季(5～10月)两群落类型日平均气温分别为(21.76±2.44)℃和(19.33±1.03)℃,且随时间呈单峰型曲线变化,热带山地常绿林日平均气温比热带山顶矮林显著高;湿季两群落类型日平均空气相对湿度分别为(88.44±2.90)%和(97.71±0.80)%,且随时间呈“倒S型”曲线变化,热带山地常绿林各月日平均空气相对湿度比热带山顶矮林显著小;与热带山顶矮林相比,热带山地常绿林的土壤全氮、全磷、速效氮、有机质、pH和土壤厚度显著大而全钾和有效磷含量显著低;热带山地常绿林的坡度、地表岩石裸露比例和海拔高度都比热带山顶矮林显著小;主成分和相关性分析表明:空气温度、土壤有效磷、全钾、全氮及地形因子是热带云雾林的主导环境因子。
2.热带山地常绿林的优势种为线枝蒲桃(Syzygium araiocladum)、蚊母树(Distylium racemosum)、碟斗青冈(Cyclobalanopsis disciformis)、九节(Psychotria rubra)和碎叶蒲桃(S. buxifolium)等,热带山顶矮林的优势种为蚊母树、碎叶蒲桃、黄杞(Engelhardtia roxburghiana)、九节和光叶山矾(S. lancifolia)等;两森林类型的优势科都为樟科(Lauraceae)、山矾科(Symplocaceae)、茜草科(Rubiaceae)、壳斗科(Fagaceae)和木犀科(Oleaceae),优势属都为山矾属(Symplocos)、青冈属(Cyclobalanopsis)、柯属(Lithocarpus)和琼楠属(Beilschmiedia)。两森林类型间的S?rensen相似性指数为0.71。热带山地常绿林幼树(1 cm≤dbh < 5 cm)和小树(5 cm≤dbh < 10 cm)的平均密度比热带山顶矮林显著小,而成年树(dbh≥10 cm)平均密度无显著差异;前者的小树和成年树的平均胸径比后者显著大,而幼树平均胸径比后者显著小;前者的幼树、小树和成年树的平均高度比后者显著大。
(二)、基于物种物种多样性的热带云雾林群落组配机制研究
3.热带山地常绿林群落的物种丰富度观测值、一阶刀切法指数、二阶刀切法指数和抽样法物种丰富度指数都比热带山顶矮林显著高,但群落多度比热带山顶矮林显著低。应用幂律模型、指数模型和逻辑斯蒂模型对热带山地常绿林和热带山顶矮林的物种―面积关系进行拟合,发现逻辑斯蒂模型是两森林类型中物种―面积关系的最优模型。应用分割线段模型、生态为优先模型、Zipf模型、Zipf-Mandelbrot模型和中性理论模型对热带山地常绿林和热带山顶矮林种―多度分布进行拟合,发现Zipf-Mandelbrot模型是热带山地常绿林种―多度分布的最优模型,而生态位优先模型和Zipf-Mandelbrot模型是热带山顶矮林种―多度分布的最优模型。广义线性模型分析表明光合有效辐射、空气温度、土壤氮和磷、坡度等环境筛对群落的物种丰富度及多度有显著影响。
4.应用4种共有度指数(the number of checkerboard species pairs、C score、种间联结数和V ratio)研究热带云雾林物种在5 m×5 m、10 m×10 m、20 m×20 m和30 m×30 m样方尺度上的共有度(co-occurrence)格局。与随机期望值相比,热带山地常绿林群落的所有物种、不同多度及径级物种在5 m和10 m样方尺度上呈现较少的共有度格局,且该格局在5 m样方尺度上最明显。显示物种间显著地非随机分离,说明物种间竞争作用于群落组配,且该作用在较小尺度上最强。热带山顶矮林群落的所有物种、不同多度及径级物种在20 m和30 m样方尺度上均呈现较多的共有度格局,且该格局在20 m尺度上最明显。显示物种间显著地非随机聚集,说明正相互作用(如促进作用facilitation)作用于群落组配,且该作用在较大尺度上最强。
5.应用Donnelly最近邻体指数分析物种分布类型,发现热带山地常绿林和热带山顶矮林不同分布类型物种百分数的大小顺序为规则分布>随机分布>聚集分布;随径级增大,聚集分布和随机分布的物种百分数呈减小趋势,而规则分布的物种百分数呈增大趋势。两森林类型中共同出现的优势物种蚊母树和碎叶蒲桃的幼树呈聚集分布,小树呈聚集分布或随机分布,而大树基本上呈随机分布。应用单变量和双变量O-ring函数分析热带山地常绿林和热带山顶矮林中碎叶蒲桃和蚊母树的不同径级种群的空间格局及其相互关联,发现蚊母树的幼苗和小树主要在小于10 m样方尺度上聚集分布,而碎叶蒲桃的幼树和小树在大于23 m样方尺度上聚集分布,不同物种幼树的空间分布与种子扩散方式有关。蚊母树的小树与幼树、大树与幼树及大树与小树的空间格局主要在小于5 m样方尺度上正关联;碎叶蒲桃的小树与幼树、大树与小树的空间格局负关联。
6.与零假设模型比较,热带山地常绿林和热带山顶矮林分别有(41±4)%和(37±2)%的物种与不同物种的个体间、(43±4)%和(38±7)%的物种与相同物种的个体间、(28±2)%和(44±5)%的物种与所有物种的个体间呈现显著的胸径―最近邻体距离相关关系,说明非随机过程在群落组配中有重要作用;两森林类型分别有(23±3)%和(26±5)%的不同物种的个体间、(27±5)%和(19±9)%的相同物种的个体间、(17±8)%和(27±7)%的所有物种的个体间呈现显著的胸径―最近邻体距离正相关关系,说明了竞争作用驱动群落组配;物种间大小―距离的显著负相关关系反映了促进作用驱动群落组配;热带山地常绿林中物种竞争作用的重要性和强度比热带山顶矮林小,竞争作用的重要性和强度随土壤肥力减小而增大。
(三)、基于植物功能性状的热带云雾林群落组配机制研究
7、热带山地常绿林群落中多度加权比叶面积平均值和多度加权树木高度平均值比热带山顶矮林显著大,而比叶面积和高度可塑性比热带山顶矮林显著小;多元逐步回归表明比叶面积大小及其可塑性与土壤全磷及空气温度显著相关,高度大小仅与空气温度显著相关,高度可塑性与上述两个环境因子都显著相关。说明了空气温度和土壤磷等环境筛作用于热带云雾林物种功能性状变化,从而影响群落物种组成和结构。8.以热带山顶矮林为例分析物种功能性状变化与群落组配的关系。热带山顶矮林中三个地点(松林顶、雅加松顶和斧头岭)的种内比叶面积和种间比叶面积随光照强度增大而减小。种内比叶面积和种间比叶面积在三个地点间有显著差异,广义混合模型分析表明样地间比叶面积差异仅与空气温度显著相关。alpha比叶面积值随群落内光照强度增大而减小,beta比叶面积值在不同样地间有显著差异。说明热带山顶矮林通过群落内光照和群落间空气温度两种环境筛作用,使物种按照功能性状变化在群落内和群落间进行组配。
9.以热带山顶矮林为例,以功能性状为基础在4个样方尺度(5 m×5 m、10 m×10 m、20 m×20 m和30 m×30 m)上对群落组配进行零假设检验。与零假设模型比较,热带山顶矮林中比叶面积小的物种在20 m和30 m样方尺度上有生存优势(over-representation),而高度大的物种在4个样方尺度上都有生存优势。线性回归表明比叶面积平均值的观测值与期望值的效应大小(effect size)仅与日平均气温及日平均最高气温显著正相关,最大物种高度平均值的观测值与期望值的效应大小仅与光合有效辐射正相关。说明热带山顶矮林物种根据比叶面积和高度大小变化进行非随机组配,此组配过程主要由空气温度和物种对光照的竞争作用驱动。
10.综合本文的研究结果可以看出:低温和低磷等环境因子对不同功能性状的物种具有筛选作用,影响其在群落中的分布与多度;在同一群落内促进作用和物种对光照等环境因子的竞争作用又影响着物种的空间配置与个体比例。因此,热带云雾林的群落结构主要由非随机的生态位过程组配形成。
Tropical forests are terrestrial ecosystems harboring the most abundant species and complicated structures, and play important roles in the regional and global biodiversity conservation and ecological function maintenance. Studies on the mechanisms of species coexistence in these ecosystems have been a vitally important topic in community ecology in recent years. Although some studies have been carried out in the low altitudinal tropical forests, species assembly and community structuring in the high altitudinal tropical cloud forests are poorly understood. Compared with the lower altitude tropical forests, environmental conditions in tropical cloud forests are quite different, characterized by frequent fog, low temperature, high humidity and strong winds. Moreover, trees in these forests are typically more deformed and elfin with small sturdy leaves, few buttress,and thick cover of epiphytes. These special environmental and physiognomical features may confine a unique rule of community assembly. The small stature of trees in the tropical cloud forests make it possible to precisely measure some important functional traits (such as specific leaf area and maximum species height). Tropical cloud forests in Hainan Island are distributed at more than 1200 m altitude in Bawangling Mt., Jianfengling Mt., Wuzhishan Mt., Yinggeling Mt., etc, including tropical montane evergreen forest (TMEF) and tropical (montane) dwarf forest (TDF or TMDF). To explore the structuring and assembly rules of tropical cloud forest communities, we investigated the species diversity and environmental conditions, and measured the specific leaf area (SLA), dbh and height for 5765 individuals trees and shrubs (dbh≥1 cm) in TMEF and TMDF in Bawangling National Natural Reserve, Hainan island, South China. Then, we compared the environmetal conditions, community stucture and assembly rules for the two cloud forest types. The main results are as follows.
(Ⅰ) Environmental conditions and community features of the tropical cloud forests
1. Daily photosynthetically active radiation (PAR) showed a unimodal curve both in TMEF and TMDF, but PAR was significantly lower in TMEF than TMDF. From May to October, mean daily air temperature differed significantly between TMEF and TMDF and showed a unimodal curve in the two forests, with average values of (21.76±2.44)°C and (19.33±1.03)°C, respectively. Additionally, mean daily relative humidity differed significantly between TMEF and TMDF and showed an“inverse S”curve; average values were (88.44±2.90) % and (97.71±0.80) %, respectively. TMEF had higher total nitrogen, total phosphorous, available nitrogen, organic matter, pH and soil thickness, but lower total potassium and available phosphorous than TMDF. Slope, cover of exposed rock and altitude were lower in TMEF than TMDF. Principal component analysis and Pearson’s correlation analysis indicated that air temperature, soil phosphorous, potassium, nitrogen and the three topographic factors were the most important predictors of distribution of these tropical cloud forests.
2. The dominant species in TMEF were Syzygium araiocladum, Distylium racemosum, Cyclobalanopsis disciformis, Psychotria rubra and S. buxifolium, and the dominant species in TMDF were D. racemosum, S. buxifolium, Engelhardtia roxburghiana, P. rubra and S. lancifolia. The common dominant families for the two forest types were Lauraceae, Symplocaceae, Rubiaceae, Fagaceae and Oleaceae. The common dominant genera for the two forest types were Symplocos, Cyclobalanopsis, Lithocarpus and Beilschmiedia. The S?rensen species similarity index for the two forest types was 0.71. The mean stem density for saplings (1 cm≤dbh﹤5 cm) and small trees (5 cm≤dbh﹤10 cm) was significantly lower in TMEF than TMDF, while there were no differences in mean stem density for adult trees (10 cm≤dbh) between these two forest types. Mean dbh for small trees and adult trees were significantly higher in TMEF than TMDF, while mean dbh for saplings were significantly lower in TMEF than TMDF. Mean plant height for saplings, small trees and adult trees were significantly higher in TMEF than TMDF.
(Ⅱ) Community assembly based on species diversity
3. The observed species richness values, as well as the species richness values predicted by 1st order Jackknife estimator, 2nd order Jackknife estimator and bootstrap estimator, were significantly higher in TMEF than TMDF; while the individual abundance was significantly lower in TMEF than TMDF. Compared with power curve and exponential curve, logistic curve was the optimal model simulating the species-area relation for the two forest types. After Brokenstick model, Niche preemption model, Zipf model, Zipf-Mandelbrot model and Neutral theory model were used to simulate the species-abundance distribution for TMEF community and TMDF community, it was found that Zipf-Mandelbrot model was the optimal model predicting the species-abundance distribution in TMEF, while Niche preemption model and Zipf-Mandelbrot model were the optimal models predicting the species-abundance distribution in TMDF. A general linear anaysis model revealed that PAR, air temperature, soil nitrogen and phosphorus were the major environmental filters predicting the species richness and individual abundance between the two forest communities..
4. Patterns of species co-occurrence in TMEF and TMDF were assessed at 5 m×5 m, 10 m×10 m, 20 m×20 m and 30 m×30 m plot sizes using four species co-occurrence indices, including the number of checkerboard species pairs, the checkerboard score of matrix, the number of species combinations and the variance ratio. All combined species, species with different abundance classes and species with different dbh classes in TMEF all showed less co-occurrence patterns than null model tests at 5 m and 10 m plot sizes, indicating that species were non-randomly segregated, and competition probably impacted on the community assembly in this forest. The species co-occurrence indices were the highest at 5 m plot size. In contrast, all combined species, species with different abundance classes and species with different dbh classes in TMDF all showed more co-occurrence patterns than null model tests at 20 m and 30 m plot sizes, indicating that species were non-randomly aggregated, and facilitation probably impacted on the community assembly in this forest. The species co-occurrence indcies were the highest at 20 m plot size.
5. Patterns of species distribution in tropical cloud forests were assessed using Donnelly nearest neighbor distance index. Percentage of species in regular distribution was higher than that of random distribution and clumped distribution, while percentage of species in clumped distribution was the lowest; percentage of species in random distribution and clumped distribution decreased, while percengate of species in regular distribution increased with increasing dbh. Patterns of distribution of saplings for the common dominant species, S. buxifolium and D. racemosum in both TMEF and TMDF, showed a clumped distribution, their small tress showed a clumped or random distribution, and their adult trees showed a random distribution. A univariate O-ring function analysis revealed that saplings and small trees for D. racemosum were aggregated at less than 10 m plot size, while saplings and small trees for S. buxifolium were aggregated at more than 23 m plot size. A bivariate O-ring function analysis showed that the spatial patterns between small trees and saplings, adult trees and saplings, and adult trees and small trees for D. racemosum were positively associated at less than 5 m plot size. On the contrary, the spatial patterns between small trees and saplings, and adult trees and small trees for S. buxifolium were negatively associated.
6. Based on null model tests, (41±4) % and (37±2) % heterospecific trees, (43±4) % and (38±7) % conspecific trees, and (28±2) % and (44±5) % all combined tree species in TMEF and TMDF, showed significant correlations between dbh and nearest neighbor distance, indicating that non-random processes affected on community assembly in tropical cloud forests. (23±3) % and (26±5) % heterospecific trees, (27±5) % and (19±9) % conspecific trees, (17±8) % and (27±7) % all combined tree species in TMEF and TMDF, showed significantly positive correlations between dbh and nearest neighbor distance, indicating that competition impacted on community assembly in the tropical cloud forests. However, negative correlations between dbh and nearest neighbor distance based on null model tests showed that facilitation affected community assembly in the tropical cloud forests. Both importance and intensity of competition statistically increased with decreasing forest productivity from TMEF to TMDF for all the shared species spanning these two forests. (Ⅲ) Community assembly based on functional traits
7. Both abundance-weighted mean SLA and plant height were significantly higher in TMEF than TMDF, while phenotypic plasticity in SLA and height was significantly lower in TMEF than TMDF. Multiple linear regression analyses indicated that among the measured environmental factors, both air temperature and soil total phosphorus were significantly correlated with mean SLA and its plasticity, and only air temperature was correlated with mean height. But both air temperature and soil total phosphorus was significantly correlated with the plasticity index of plant height. These results indicate that air temperature and soil phosphorus acted as environmental filters on decreasing SLA and plant height, while increasing the plasticity of the functional traits from TMEF to TMDF, and thus impact on species composition and community structure in tropical cloud forests.
8. SLA decreased significantly with increasing solar irradiance within the three study sites in TMDF, and differed significantly among the three sites both for within and among species comparisons. Mean plot SLA, accounting for both within and among species across the three sites, increased significantly in relation to air temperature but not local PAR and soil total phosphorus. Alpha SLA decreased significantly with increasing solar irradiance within the three sites and beta SLA differed significantly among the three sites. The strong relationship between both intra- and interspecific variation in SLA and environmental conditions strongly confirms the role of trait variation in the assembly of plant species into tropical cloud forest communities via environment filtering related to light availability and air temperature.
9. Community assembly were examined based on null model tests at 5×5 m, 10×10 m, 20×20 m and 30×30 m plot sizes using a trait-based approach in TMDF. Lower SLA growing forest plants were over-represented within forest communities at 20 m and 30 m plot sizes, and taller growing forest plants were over-represented within forest communities at the four plot sizes. The correlation between effect size of the test statistic (i.e. mean trait value) of null model tests for specific leaf area and air temperature was positive, and that for maximum species height and percentage reduction of PAR was positive, revealing that there is a stress-tolerator advantage for lower specific leaf area species to be adapted to low temperature, and a size-advantage for taller species competing for light. Thus species are non-randomly assembled with respect to both specific leaf area and maximum species height in the tropical montane dwarf forest community, and that these processes are driven by both the temperature stress and biotic competition.
10. In conclusion, the low air temperature and low soil phosphorus act as environmental filters on species with different functional traits, affecting their distribution and abundance in tropical cloud forests; meanwhile, both competition and facilitation processes affect the spatial patterning and proportions of individual abundance within a community. The tropical cloud forest communities are mainly assembled by niche-based non-random processes.
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