青藏高原“黑土滩”次生毒杂草群落成体植株与幼苗空间异质性及相似性分析

doi:10.3773/j.issn.1005-264x.2008.05.020

摘要/Abstract

摘要：

在样线调查基础上, 用半方差函数、分形维数、空间自相关等方法对青藏高原“黑土滩”次生毒杂草群落地上成体植株、幼苗空间的异质性, 二者物种构成的相似性及其尺度特征进行分析。结果表明, “黑土滩”次生毒杂草群落在较大尺度上地上成体植株物种数的空间依赖性强, 异质性高, 而个体密度则较均匀; 幼苗物种数在小尺度上空间异质性高, 幼苗密度在大尺度上空间异质性较高, 幼苗密度独立于其物种分布, 高密度的幼苗分布在微地形下和群落间隙中, “黑土滩”毒杂草植物幼苗充分利用空余生态位进行群落补充; “黑土滩”群落在不同尺度上地上成体植株与幼苗物种构成相似性变化的规律性不大; “黑土滩”群落幼苗靠其高密度特征完成群落更新, 植物群落幼苗更新力强, 导致“黑土滩”毒杂草群落趋于稳定。因此, 建议加强干扰以减弱“黑土滩”次生毒杂草群落稳定趋势, 这对恢复“黑土滩”具有重要指导意义。

关键词: 青藏高原, “黑土滩”退化草地, 毒杂草, 幼苗, 空间异质性, 群落更新

Abstract:

Aims ‘Black soil land’ grassland on the Tibetan Plateau results from degradation of Kobresia alpine meadow and has many weeds and poisonous plants. This disturbed grassland forms in small bottomland patches. There are no studies of community spatial patterns, relationships and scale patterns of adult plants and seedlings are important to explaining the formation of this secondary weed community.

Methods We selected a typical ‘black soil land’ community of about 30 m × 50 m in the headwaters of the Yellow River and used 100 sample plots (50 cm × 50 cm) to investigate number and density of adults and seedlings (determined by pulling) by species. Spatial heterogeneity of the community and the similarity between adult plants and seedlings were analyzed by semi-variance, fractal dimension, spatial correlation spatial autocorrelation, etc.

Important findings Species number of adult plants is highly spatially heterogeneous and plant density homogeneous at large scales. Species number of seedlings is highly spatially heterogeneous at small scales, and its density is highly spatially heterogeneous at large scales. Seedlings have high density in areas of micro-topography and gaps of adult plants, where seedlings grow and establish in empty ecological niches. The ‘black soil land’ community regenerates and recruits in vegetation gaps. The generation of ‘black soil land’ community depends on high density of seedlings of weeds and poisonous plants, and its generation capability is strong. According to our results, the ‘black soil land’ secondary community becomes more stable without interference. We suggest that human management be used to decrease the stability of the ‘black soil land’ weed community and restore alpine meadow.

Key words: Qinghai-Tibetan Plateau, ‘black soil land’ degraded grassland, weed and poisonous plant, seedling, spatial heterogeneity, community regeneration

尚占环, 龙瑞军, 马玉寿, 丁路明. 青藏高原“黑土滩”次生毒杂草群落成体植株与幼苗空间异质性及相似性分析. 植物生态学报, 2008, 32(5): 1157-1165. DOI: 10.3773/j.issn.1005-264x.2008.05.020

SHANG Zhan-Huan, LONG Rui-Jun, MA Yu-Shou, DING Lu-Ming. SPATIAL HETEROGENEITY AND SIMILARITY OF ADULT PLANTS AND SEEDLINGS IN ‘BLACK SOIL LAND’ SECONDARY WEED COMMUNITY, QINGHAI-TIBETAN PLATEAU. Chinese Journal of Plant Ecology, 2008, 32(5): 1157-1165. DOI: 10.3773/j.issn.1005-264x.2008.05.020

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https://www.plant-ecology.com/CN/Y2008/V32/I5/1157

图/表 11

表1 地上成体植株物种数的半方差图拟合的4个模型统计量

Table 1 Statistics parameters of four models fit semi-variogram about species number of adult plants

参数/模型 Parameters/Models	球状模型 Spherical	指数模型 Exponential	线性模型 Linear	高斯模型 Gaussian
块金值 Nugget Co	0.022 6	0.022 2	0.023 35	0.030 0
台基值 Sill Co+C	0.085 2	0.129 4	0.129 40	0.231 0
变程 Range Ao	51	51	24	51
空间变异比 Proportion C/(Co+C)	0.735	0.828	0.637	0.870
决定系数 R²	0.539	0.530	0.552	0.597
残差平方和 RSS	0.005 577	0.005 678	0.033 200	0.004 877

表2 地上幼苗物种数半方差图拟合的4个模型统计量

Table 2 Statistics parameters of four models fit semi-variogram about species number of seedlings

参数/模型 Parameters/Models	球状模型 Spherical	指数模型 Exponential	线性模型 Linear	高斯模型 Gaussian
块金值 Nugget Co	0.037	0.036 1	0.037 76	0.043 2
台基值 Sill Co+C	0.078	0.109 2	0.065 88	0.160 4
变程 Range Ao	47.89	47.64	24	49.1
空间变异比 Proportion C/(Co+C)	0.526	0.669	0.472	0.731
决定系数 R²	0.359	0.364	0.359	0.321
残差平方和 RSS	0.005 598	0.005 561	0.018 600	0.005 933

图1 双对数坐标下地上成体植株和幼苗的物种数拟合的半方差函数图及分形维数 PSN: 地上成体植株物种数 Adult plant’ s species number SSN: 幼苗物种数 Seedling’s species number

Fig. 1 Semi-variogram and fractal dimension about species number of adult plants and seedlings at double-logarithm coordinate

表3 地上成体植株密度半方差图拟合的4个模型统计量

Table 3 Statistics parameters of four models fit semi-variogram about species density of adult plants

参数/模型 Parameters/Models	球状模型 Spherical	指数模型 Exponential	线性模型 Linear	高斯模型 Gaussian
块金值 Nugget Co	0.023 5	0.021 2	0.024 289	0.041
台基值 Sill Co+C	0.165 0	0.270 4	0.118 935	0.461
变程 Range Ao	51	51	24	49.22
空间变异比 Proportion C/(Co+C)	0.858	0.922	0.796	0.911
决定系数 R²	0.678	0.668	0.685	0.706
残差平方和 RSS	0.016 7	0.017 2	0.016 4	0.015 3

表4 地上幼苗密度半方差图拟合的4个模型统计量

Table 4 Statistics parameters of four models fitting semi-variogram about density of seedlings

参数/模型 Parameters/Models	球状模型 Spherical	指数模型 Exponential	线性模型 Linear	高斯模型 Gaussian
块金值 Nugget Co	0.001	0.001	0.001	0.038
台基值 Sill Co+C	0.356	0.603	0.491	1.240
变程 Range Ao	51	51	48.94	51
空间变异比 Proportion C/(Co+C)	0.997	153	0.998	0.969
决定系数 R²	0.739	0.998	0.757	0.840
残差平方和 RSS	0.082 2	0.722 0	0.076 1	0.050 1

图2 双对数坐标下地上成体植株和幼苗密度的半方差函数图及分形维数 PSD: 地上成体植株密度 Adult plant density SSD: 幼苗密度 Seedling density

Fig. 2 Semi-variogram and fractal dimension about species density of adult plants and seedlings at double-logarithm coordinate

表5 地上成体植株Shannon-Wiener多样性半方差图拟合的4个模型统计量

Table 5 Statistics parameters of four models fitting semi-variogram about Shannon-Wiener of adult plants

参数/模型 Parameters/Models	球状模型 Spherical	指数模型 Exponential	线性模型 Linear	高斯模型 Gaussian
块金值 Nugget Co	0.004 6	0.004 4	0.005 02	0.009 8
台基值 Sill Co+C	0.054 3	0.089 3	0.038 05	0.179 7
变程 Range Ao	51	51	24	51
空间变异比 Proportion C/(Co+C)	0.915	0.951	0.868	0.945
决定系数 R²	0.551	0.535	0.575	0.687
残差平方和 RSS	0.003 384	0.003 506	0.021 300	0.002 362

表6 地上幼苗Shannon-Wiener多样性半方差图拟合的4个模型统计量

Table 6 Statistics parameters of four models fitting semi-variogram about Shannon-Wiener of seedlings

参数/模型 Parameters/Models	球状模型 Spherical	指数模型 Exponential	线性模型 Linear	高斯模型 Gaussian
块金值 Nugget Co	0.004 89	0.004 9	0.005 071	0.007 5
台基值 Sill Co+C	0.030 28	0.047 9	0.022 028	0.095 4
变程 Range Ao	51	51	24	51
空间变异比 Proportion C/(Co+C)	0.839	0.898	0.77	0.921
决定系数 R²	0.467	0.453	0.49	0.601
残差平方和 RSS	0.001 242	0.001 267	0.005 956	0.000 930

图3 双对数坐标下地上成体植株和幼苗Shannon-Wiener多样性的半方差函数图及分形维数 PSWI: 地上成体植株多样性 Adult plant diversity SSWI: 幼苗多样性 Seedling diversity

Fig. 3 Semi-variogram and fractal dimension about Shannon-Wiener index of adult plants and seedlings at double-logarithm coordinate

图4 “黑土滩”幼苗与地上成体植株的相似性在两条样线上的变化

Fig. 4 Variation of Jaccard similarity index between adult plants and seedlings along two sample lines (NW-SE, NE-SW) in ‘Black soil land’

图5 “黑土滩”地上成体植株和幼苗Moran-I相似性系数的自相关系数在两条样线不同尺度上的变化

Fig. 5 Variation of spatial self-relationship coefficients (Moran-I) along two line-samples (NW-SE, NE-SW) in ‘Black soil land’

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