长白山温带森林不同演替阶段群落功能性状的空间变化

doi:10.17521/cjpe.2018.0295

摘要/Abstract

摘要：

群落构建机制研究是生态学研究的热点。长白山自然保护区拥有完整的原始阔叶红松林生态系统, 近年来随着物种多样性丧失愈发严重, 对该地区开展群落构建机制研究显得尤为重要。该研究以长白山不同演替阶段的3块5.2 hm ²固定监测样地(次生杨桦林、次生针阔混交林、原始椴树红松林)为研究对象, 通过采集样地内主要树种的6个关键功能性状(叶面积、比叶面积、叶片厚度、叶氮含量、叶磷含量、最大树高), 分析不同空间尺度下(5 m × 5 m, 10 m × 10 m, 20 m × 20 m, 30 m × 30 m, 40 m × 40 m, 50 m × 50 m和60 m × 60 m)及不同演替阶段群落性状空间值的变化, 结合零模型的模拟结果对长白山温带森林演替过程中的群落构建机制进行讨论。结果表明: 种库大小对于研究结果具有重要影响。在较大的种库下, 环境过滤作用影响显著。而在样地水平进行研究时, 演替早期和中期, 群落性状空间值与零模型模拟值无显著差异, 在演替的晚期, 群落性状空间值显著高于零模型模拟值。结合多个群落功能多样性指数分析发现, 环境过滤和竞争作用共同决定该地区顶级群落的物种组成。在演替早期大量物种迁入, 群落内物种间存在强烈的资源竞争, 而随着演替进行, 部分物种逐渐被竞争排除出群落, 群落中的物种呈现明显的生态位分化, 竞争作用是维持物种共存的主要机制。

关键词: 群落构建, 功能性状, 性状空间, 空间尺度

Abstract:

Aims The community assembly mechanisms are among the focal topics in ecological studies. In Changbai Mountains Nature Reserve, there is an intact primary broadleaved-Korean pine forest ecosystem. With increasing loss of species diversity in recent years, study that explores the community assembly mechanisms in this region is particularly important.

Methods This study was conducted in three large permanent plots, each of the size 5.2 hm ², along suessional stages (secondary poplar and birch mixed forest, PBF; secondary mixed conifer and broad-leaved forest, CBF; and primary Tilia amurensis-Pinus koraiensis mixed forest, TKF) in Changbai Mountains. Six functional traits of major tree species were measured, including leaf area, specific leaf area, leaf thickness, leaf nitrogen content, leaf phosphorus content, and maximum tree height. Changes in the spatial values of community traints were analyzed at different spatial scales (5 m × 5 m, 10 m × 10 m, 20 m × 20 m, 30 m × 30 m, 40 m × 40 m, 50 m × 50 m and 60 m × 60 m). By comparing the observed values with expected values of null models, the community assembly mechanisms in temperate forests of Changbai Mountains were explored.

Important findings Results show that the size of species pool has an important impact on the outcome; in a larger species pool, the environmental filtration has a significant impact. At the plot level and for early and intermediate stages of succession, the observed spatial values of community traits do not significantly differ from the expected values. At the late successional stage, the observed spatial values of community traits were greater than the expected values. The analysis of multiple community functional diversity indices shows that the combined processes of habitat filtring and competitive exclusion are the main determinants of the species composition of the climax community in this region. In the early successional stage, large numbers of species are immigrated, and there are strong resource competitions among the species within a community. With progressing succession, some species are excluded, species maintained in the community show significant niche differentiations, and competition is the main mechanism species coexistence.

Key words: community assembly, functional trait, trait space, spatial scale

郝姝珺, 李晓宇, 侯嫚嫚, 赵秀海. 长白山温带森林不同演替阶段群落功能性状的空间变化. 植物生态学报, 2019, 43(3): 208-216. DOI: 10.17521/cjpe.2018.0295

HAO Shu-Jun, LI Xiao-Yu, HOU Man-Man, ZHAO Xiu-Hai. Spatial variations of community functional traits at different successional stages in temperate forests of Changbai Mountains, Northeast China. Chinese Journal of Plant Ecology, 2019, 43(3): 208-216. DOI: 10.17521/cjpe.2018.0295

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2018.0295

https://www.plant-ecology.com/CN/Y2019/V43/I3/208

图/表 6

图1 长白山次生杨桦林(PBF)、次生针阔混交林(CBF)、原始椴树红松林(TKF)研究区位置。

Fig. 1 Location of secondary poplar and birch mixed forest (PBF), secondary mixed conifer and broad-leaved forest (CBF), primary Tilia amurensis-Pinus koraiensis mixed forest (TKF) in Changbai Mountains.

表1 长白山永久监测样地概况

Table 1 Summary of permanent forest plots in Changbai Mountains

林分类型 Forest type	样地面积 Plot area (hm²)	经纬度 Longitude & latitude	平均海拔 Average elevation (m)	物种数量 Species number	总胸高断面积 Total basal area (m²)
次生杨桦林 PBF	5.2 (260 m × 200 m)	42.32° N, 128.13° E	893	69	24.74
次生针阔混交林 CBF	5.2 (260 m × 200 m)	42.35° N, 128.13° E	810	66	32.07
原始椴树红松林 TKF	5.2 (260 m × 200 m)	42.23° N, 128.08° E	1 023	22	56.64

表2 功能性状及生态学意义

Table 2 Functional traits and ecological meaning

功能性状 Functional trait	单位 Unit	生态学意义 Ecological meaning
叶面积 Leaf area	mm2	代表叶片对光的捕获能力, 与环境胁迫和干扰下的生态策略有关 Represents the ability to capture light; linked to ecological strategy with respect to environmental stress and disturbances
比叶面积 Specific leaf area	mm2·mg^-1	与叶片光合能力正相关, 与叶片寿命负相关 Positively related to photosynthetic rate and negatively to leaf longevity
叶片厚度 Leaf thickness	mm	代表植物抗干扰和高投入的能力 Represents the ability to defy changes and high investment
叶氮含量 Leaf nitrogen concentration	mg·g^-1	与光合作用蛋白和最大光合速率有关 Related to the proteins in photosynthetic machinery and maximum photosynthetic rate
叶磷含量 Leaf phosphorus concentration	mg·g^-1	与植物最大光合速率以及营养级质量有关 Related to maximum photosynthetic rate and high nutritional quality in food webs
最大树高 Maximum tree height	m	代表植株在光垂直梯度所处的位置及竞争能力 Associated with position of the species in the vertical light gradient of plants and competitiveness

图2 长白山温带森林不同演替阶段样地单个功能性状的加权平均值(CWM)。PBF, 次生杨桦林; CBF, 次生针阔混交林; TKF, 原始椴树红松林。箱型图为重复抽样20次的结果, 黑色三角为样方水平数值; 不同小写字母为多重比较的结果, 表示差异显著(p ＜ 0.05)。

Fig. 2 The weighted average (CWM) of individual functional traits at different successional stages in temperate forests of Changbai Mountains. PBF, secondary poplar and birch mixed forest; CBF, secondary mixed conifer and broad-leaved forest; TKF, primary Tilia amurensis-Pinus koraiensis mixed forest. The box plots show the results of repeated sampling for 20 times. The black triangle is values at plot level; different lowercase letters designate the results of multiple comparisons, and it indicate significant difference (p ＜ 0.05).

图3 长白山温带森林不同演替阶段功能多样性的变化趋势。PBF, 次生杨桦林; CBF, 次生针阔混交林; TKF, 原始椴树红松林。箱型图为重复抽样20次的结果, 黑色三角为样方水平数值; 不同小写字母为多重比较的结果, 表示差异显著(p ＜ 0.05)。

Fig. 3 Trends in functional diversity at different successional stages in temperate forests of Changbai Mountains. PBF, secondary poplar and birch mixed forest; CBF, secondary mixed conifer and broad-leaved forest; TKF, primary Tilia amurensis-Pinus koraiensis mixed forest. FEve, functional uniformity; FDiv, functional divergence; FDis, functional dispersion; RaoQ, functional entropy index. The box plots show the results of repeated sampling for 20 times. The black triangle is the value at plot level; different lowercase letters show the results of multiple comparisons, and it indicate significant difference (p ＜ 0.05).

图4 长白山温带森林不同演替阶段群落性状空间值随空间尺度的变化。A, D, 次生杨桦林。B, E, 次生针阔混交林。C, F, 原始椴树红松林。A, B, C, 种库一。D, E, F, 种库二。SES, 标准化效应值。***, p ＜ 0.001, 表示性状空间观测值与零模型模拟值存在显著差异。

Fig. 4 Changes in the spatial value of community traits at different succession stages with spatial scale in temperate forests of Changbai Mountains. A, D, Secondary poplar forest. B, E, Secondary coniferous and broad-leaved mixed forest. C, F, Primary Tilia amurensis-Pinus koraiensis mixed forest. A, B, C, Species pool 1. D, E, F, Species pool 2. SES, standardized effect size. ***, p ＜ 0.01, it indicates that there is a significant difference between the eigenspace observations and the zero model simulation values.

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