尖峰岭热带山地雨林优势树种白颜树空间分布格局

doi:10.3724/SP.J.1258.2014.00122

摘要/Abstract

摘要：

优势树种对群落的构建和形成起主要作用。该文以海南岛尖峰岭60 hm²样地内重要值第二大的树种白颜树(Gironniera subaequalis)为研究对象, 探索其种群分布格局特征, 讨论环境异质性、密度依赖死亡、扩散限制等机制在格局形成过程中所起的作用。将白颜树10022个植株分为6个径级, 分别归属于幼树、中龄树、成年树三个生活史阶段, 采用成对相关函数分析各径级的空间分布特征; 双变量成对相关函数和标签关联函数分析不同生活史阶段之间的空间关系; Berman检验方法检验3个地形因子分别对幼树、中龄树、成年树分布影响的显著度。结果表明: 白颜树种群内I、II、III、IV径级呈现聚集分布, 聚集程度随径级的增加减弱; V和VI径级小尺度上均匀分布, 大尺度上以随机分布为主。幼树与中龄树空间正关联; 幼树与成年树空间负关联; 成年树与中龄树在较小的尺度上负关联, 大尺度上微弱正关联。但是不同生活史阶段的个体之间彼此分离, 个体间无直接的促进作用。地形因子中, 坡度、海拔、凹凸度对幼树的分布影响显著; 坡度、凹凸度对中龄树的分布影响显著; 仅坡度对成年树的分布影响显著。从现有的空间格局可以推断出环境异质性和密度依赖死亡对格局形成起作用, 但是种子的扩散限制对空间格局的影响没有明确地表现出来。

关键词: 白颜树, 点格局分析, 径级, 种群分布格局, 地形因子

Abstract:

Aims Dominant species play a major role in the formation of community. Our objectives were to detect and assess the spatial pattern of Gironniera subaequalis, which is a dominant species with the second largest importance value in the tropical montane rainforest of Jianfengling, and to explore the effects of environmental heterogeneity, density-dependent mortality and dispersal limitation on the formation of its current spatial pattern.

Methods 10022 individuals were categorized into six size classes and three life-history stages, i.e. young trees, middle-aged trees and adult trees, respectively. The spatial pattern of each age class was analyzed by pair-correlation function. The spatial associations between paris of life-history stages were analyzed by bivariate pair-correlation function and mark-connection function. The impacts of topographic factors on the distributions of young trees, middle-aged trees and adult trees were analyzed by Berman test.

Important findings Results showed that the size classes I, II, III, and IV of G. subaequalis occurred in aggregated distributions. The aggregation intensities declined with increasing size classes. Size classes V and VI exhibited a uniform distribution at the smaller scales and a random distribution mainly at the larger scales. Young trees and middle-aged trees had a positive association; whereas young trees and adult trees had a negative association. Middle-aged trees and adult trees had a negative association at smaller scales and a weakly positive association at larger scales. The individuals of different life-history stages were separated from each other. There were no direct facilitative interactions between pairs of individuals. The tree topographic factors, slope degree, altitude and concave-convex, had significant impacts on the distribution of small trees; slope degree and concave-convex had significant impacts on the distribution of the middle-aged trees; whilst only slope degree had significant impacts on the adults trees. We inferred that the environmental heterogeneity and density-dependent mortality contributed to current spatial pattern of G. subaequalis, while the effects of dispersal limitation were not obvious.

Key words: Gironniera subaequalis, point pattern analysis, size class, spatial pattern of population, topographic factor

梁爽, 许涵, 林家怡, 李意德, 林明献. 尖峰岭热带山地雨林优势树种白颜树空间分布格局. 植物生态学报, 2014, 38(12): 1273-1282. DOI: 10.3724/SP.J.1258.2014.00122

LIANG Shuang, XU Han, LIN Jia-Yi, LI Yi-De, LIN Ming-Xian. Spatial distribution pattern of the dominant species Gironniera subaequalis in tropical montane rainforest of Jianfengling, Hainan Island, China. Chinese Journal of Plant Ecology, 2014, 38(12): 1273-1282. DOI: 10.3724/SP.J.1258.2014.00122

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链接本文: https://www.plant-ecology.com/CN/10.3724/SP.J.1258.2014.00122

https://www.plant-ecology.com/CN/Y2014/V38/I12/1273

图/表 7

表1 白颜树在群落中的数量指标

Table 1 Quantitative indices of Gironniera subaequalis in community

	重要值 Importance value	相对胸高断面积 Relative cross-section area at the breast height	相对密度 Relative density	相对频度 Relative frequency
数值 Value	3.01	5.50	2.28	1.24
在样地内的排名 Relative position of all species	2	2	9	4

表2 生活史的划分

Table 2 Classification of life-history

	幼树 Young trees (44%)		中龄树 Middle-aged trees (34.8%)		成年树 Adult trees (21.2%)
	径级I Class I	径级II Class II	径级III Class III	径级IV Class IV	径级V Class V	径级VI Class VI
胸径 DBH	1 cm ≤ DBH < 5 cm	5 cm ≤ DBH < 10 cm	10 cm ≤ DBH < 15 cm	15 cm ≤ DBH < 20 cm	20 cm ≤ DBH < 25 cm	DBH ≥ 25 cm
株数 Number of stems	2 503	1 948	1 830	1 655	1 194	932

图1 白颜树所有个体的空间分布。三角形代表幼树; 黑色圆点代表中龄树; 正方形代表成年树。

Fig. 1 Spatial distribution of all stems of Gironniera subaequalis. Young trees are marked by the triangles; middle-aged trees are marked by the black dots; and adult trees are marked by the squares.

图2 白颜树各径级的成对相关函数。实线表示成对相关函数, 虚线表示99%包迹线。零模型为完全空间随机模型(CSR)和sigma = 15 m的异质泊松模型(HP)。

Fig. 2 Univariate pair-correlation functions of different age classes of Gironniera subaequalis. Solid lines are pair-correlation functions, and dotted lines are 99% confidence envelopes. Null models: complete spatial randomness (CSR) and Heterogeneous Poisson (HP) with sigma = 15 m.

图3 白颜树各个生活史阶段之间的双变量成对相关函数。实线表示双变量成对相关函数; 虚线表示99%包迹线。A, 包迹线采用完全空间随机零模型的幼树与中龄树的关系。B, 包迹线采用先决条件零模型的幼树与中龄树的关系。C, 包迹线采用完全空间随机零模型的幼树与成年树的关系。D, 包迹线采用先决条件零模型的幼树与成年树的关系。E, 包迹线采用完全空间随机零模型的中龄树与成年树的关系。F, 包迹线采用先决条件零模型的中龄树与成年树的关系。

Fig. 3 Bivariate pair-correlation functions between pairs of life-history stages of Gironniera subaequalis. Solid lines are bivariate pair-correlation functions and dotted lines are 99% confidence envelopes. A, Relationship between young trees and middle-aged trees with envelopes calculated by complete spatial randomness null model. B, Relationship between young trees and middle-aged trees with envelops calculated by antecedent condition null model. C, Relationship between young trees and adult trees with envelopes calculated by complete spatial randomness null model. D, Relationship between young trees and adult trees with envelops calculated by antecedent condition null model. E, Relationship between middle-aged trees and adult trees with envelopes calculated by complete spatial randomness null model. F, Relationship between middle-aged trees and adult trees with envelops calculated by antecedent condition null model.

图4 标签关联函数P(r)。A, 从所有幼树和中龄树中任选两个距离为r的个体, 两者均为幼树的概率。B, 从所有幼树和中龄树中任选两个距离为r的个体, 其中一个为幼树, 另一个为中龄树的概率。C, 从所有幼树和中龄树中任选两个距离为r的个体, 两者均为中龄树的概率。D, 从所有幼树和成年树中任选两个距离为r的个体, 两者均为幼树的概率。E, 从所有幼树和成年树中任选两个距离为r的个体, 其中一个为幼树, 另一个为成年树的概率。F, 从所有幼树和成年树中任选两个距离为r的个体, 两者均为成年树的概率。G, 从所有中龄树和成年树中任选两个距离为r的个体, 两者均为中龄树的概率。H, 从所有中龄树和成年树中任选两个距离为r的个体, 其中一个为中龄树, 另一个为成年树的概率。I, 从所有中龄树和成年树中任选两个距离为r的个体, 两者均为成年树的概率。实线表示标签关联函数; 虚线表示99%包迹线。

Fig. 4 The mark-connection functions P(r). A, Probabilities by randomly choosing two individuals (separated by distance r) from all young trees and middle-aged trees, when both are young trees. B, Probabilities by randomly choosing two individuals (separated by distance r) from all young trees and middle-aged trees, when one is a young tree and the other is a middle-aged tree. C, Probabilities by randomly choosing two individuals (separated by distance r) from all young trees and middle-aged trees, when both are middle-aged trees. D, Probabilities by randomly choosing two individuals (separated by distance r) from all young trees and adult trees, when both are young trees. E, Probabilities by randomly choosing two individuals (separated by distance r) from all young trees and adult trees, when one is a young tree and the other is an adult tree. F, Probabilities by randomly choosing two individuals (separated by distance r) from all young trees and adult trees, when both are adult trees. G, Probabilities by randomly choosing two individuals (separated by distance r) from all middle-aged trees and adult trees, when both are middle-aged trees. H, Probabilities by randomly choosing two individuals (separated by distance r) from all middle-aged trees and adult trees, when one is a young tree and the other is an adult tree. I, Probabilities by randomly choosing two individuals (separated by distance r) from all middle-aged trees and adult trees, when both are adult trees. Solid lines are mark-connection functions, and dotted lines are 99% confidence envelopes.

表3 地形因子对白颜树各生活史阶段个体分布的影响

Table 3 Influences of topographic factors on the distribution of the individuals in each life history stage of Gironniera subaequalis

	幼树 Young tree	中龄树 Middle-aged tree	成年树 Adult tree
坡度 Slope	+	+	+
海拔 Elevation	+	-	-
凹凸度 Convex	+	+	-

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