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桂西南喀斯特山地木本植物群落种间关系及CCA排序

马丰丰, 潘高, 李锡泉, 韩云娟

马丰丰, 潘高, 李锡泉, 韩云娟. 桂西南喀斯特山地木本植物群落种间关系及CCA排序[J]. 北京林业大学学报, 2017, 39(6): 32-44. DOI: 10.13332/j.1000-1522.20160379
引用本文: 马丰丰, 潘高, 李锡泉, 韩云娟. 桂西南喀斯特山地木本植物群落种间关系及CCA排序[J]. 北京林业大学学报, 2017, 39(6): 32-44. DOI: 10.13332/j.1000-1522.20160379
MA Feng-feng, PAN Gao, LI Xi-quan, HAN Yun-juan. Interspecific relationship and canonical correspondence analysis within woody plant communities in the karst mountains of Southwest Guangxi, southern China[J]. Journal of Beijing Forestry University, 2017, 39(6): 32-44. DOI: 10.13332/j.1000-1522.20160379
Citation: MA Feng-feng, PAN Gao, LI Xi-quan, HAN Yun-juan. Interspecific relationship and canonical correspondence analysis within woody plant communities in the karst mountains of Southwest Guangxi, southern China[J]. Journal of Beijing Forestry University, 2017, 39(6): 32-44. DOI: 10.13332/j.1000-1522.20160379

桂西南喀斯特山地木本植物群落种间关系及CCA排序

基金项目: 

中南林业科技大学研究生科技创新基金项目 CX2015A03

湖南省研究生创新项目 CX2015B287

世界银行贷款项目 8215-CN-KT-2

详细信息
    作者简介:

    马丰丰,助理研究员。主要研究方向:森林生态学。Email:mafengfeng0403@126.com   地址: 410004 湖南省长沙市韶山南路658号湖南省林业科学院

    责任作者:

    李锡泉,研究员。主要研究方向:生态学。Email:Li5578761@126.com   地址;同上

  • 中图分类号: S718.5

Interspecific relationship and canonical correspondence analysis within woody plant communities in the karst mountains of Southwest Guangxi, southern China

  • 摘要: 基于桂西南喀斯特山地木本植物群落11个样方(20 m×20 m)取样调查,运用生态位理论、种间关系原理、双向指示种分析(TWINSPAN)和典范对应分析(CCA)方法,探讨主要木本植物间的种间联结性和相关性及其与环境因子之间的关系。结果表明:桂西南喀斯特山地木本植物群落中麻栎、枫香、蚬木占优势地位,罗伞树-鹅掌柴、九节-琴叶榕、澄广花-鱼骨木的生态位重叠度分别在3种木本植物群落中最大。麻栎次生林群落总体种间关系表现为不显著正关联,χ2检验统计显示正关联种对数有50对,负关联种对数有52对,无关联种对数有3对,正负关联比为0.96,检验显著率为6.67%。枫香次生林群落总体种间关系表现为不显著负关联,χ2检验统计显示正关联种对数有96对,负关联种对数有113对,无关联种对数有1对,正负关联比为0.85,检验显著率为8.10%。蚬木次生林群落总体种间关系表现为显著正关联,正关联种对数有19对,负关联种对数有17对,正负关联比为1.12,检验显著率为8.33%。CCA前瞻选择显示土壤pH值、全磷、坡向、土壤全钾、海拔是影响桂西南喀斯特山地木本植物群落物种组成与分布最重要的因子,总体上可以对76.95%的环境因子进行解释。物种间的正联结性越显著,其生态位重叠程度越高,反之,物种间的负联结性越显著,其生态位重叠程度则越低。
    Abstract: The interspectific associations of main woody plant species and the relationships between environmental factors and plant communities were investigated in the karst mountains of Southwest Guangxi, southern China. Eleven sampling plots (20 m×20 m in size each) were established in three forest communities (Quercus acutissima forest, Liquidambar formosana forest, and Excentrodendron hsienmu forest) in the study site, and statistic methods including niche theory, interspecific relation principle, two-way indicator species analysis and canonical correspondence analysis were used. The results showed that Quercus acutissima, Liquidambar formosana, and Excentrodendron hsienmu were dominant species in the study area. The largest niche overlap was observed between Ardisia quinquegona and Schefflera octophylla, between Psychotria rubra and Ficus pandurata, and between Orophea hainanensis and Canthium dicoccum for the Quercus acutissima natural secondary forest, Liquidambar formosana natural secondary forest and Excentrodendron hsienmu natural secondary forest, respectively. No significant positive correlations were found in terms of the overall interspecific associations among dominant woody plant species in Quercus acutissima natural secondary forest. The χ2 test showed that the number of species pairs in positive association was 50, the number of species pairs in negative association was 52, and the number of species pairs in non-association was 3, the association ratio was 0.96, and the significance rate of test was 6.67%. The overall interspecific associations were negative correlations among dominant woody plant species in Liquidambar formosana natural secondary forest, but the correlations were not significant. The χ2 test showed that the number of species pairs in positive association was 96, the number of species pairs in negative association was 113, the number of species pairs in non-association was 1, the association ratio was 0.85, and the significance rate of test was 8.10%. The overall interspecific associations were significantly positive correlations among dominant woody plant species in Excentrodendron hsienmu natural secondary forests. The χ2 test showed that the number of species pairs in positive association was 19, the number of species pairs in negative association was 17, the association ratio was 1.12, and the significance rate of test was 8.33%. By CCA analysis, soil pH, total phosphorus, slope aspect, soil total potassium and elevation were the most important factors affecting species composition and distribution in plant communities in the study area, which accounted for about 76.95% of the variation. Our study indicates that the more significantly positive the associations between plant species are, the higher the degree of niche overlap is. In contrast, the more significantly negative the associations between plant species are, the lower the degree of niche overlap is.
  • 图  1   桂西南喀斯特山地木本植物群落11个样方的TWINSPAN分类树状图

    Figure  1.   Dendrogram of the TWINSPAN classification of 11 plots within woody plant communities in the karst mountains of Southwest Guangxi

    图  2   麻栎次生林主要木本植物种间联结性半矩阵图

    △0.5≤AC; □0.1≤AC<0.5;○-0.1≤AC<0.1;☆-0.5≤AC<-0.1;◇AC<-0.5;AC为联结系数。+正关联; -负关联; ★极显著正关联; ◆极显著负关联; ■显著正关联; ▼显著负关联; ●无关联。下同。种号同表 2

    Figure  2.   Semi-matrix of interspecific association for dominant woody plants in the Quercus acutissima natural secondary forest

    AC is association coefficient; + Positive correlation; -Negative correlation; ★Positive correlation with extreme significance; ◆ Negative correlation with extreme significance; ■Significantly positive correlation; ▼Significantly negative correlation; ●Non-correlation. The same below. Species No. was the same as Tab. 2.

    图  3   枫香次生林主要木本植物种间联结性半矩阵图

    物种序号同表 3

    Figure  3.   Semi-matrix of interspecific association for dominant woody plants in the Liquidambar formosana natural secondary forest

    Species No. was the same as Tab. 3.

    图  4   蚬木次生林主要木本植物种间联结性半矩阵图

    物种序号同表 4

    Figure  4.   Semi-matrix of interspecific association for dominant woody plants in the Excentrodendron hsienmu natural secondary forest

    Species No. was the same as Tab. 4.

    图  5   物种和环境因子的典范对应分析(CCA)排序

    图中物种序号同表 1。Elv.海拔; Slo.坡度; Asp.坡向; Pos.坡位; pH. pH值; SOM.土壤有机质; TN.全氮; TP.全磷; TK.全钾。

    Figure  5.   Canonical correspondence analysis(CCA)ordination diagram of species and environmental factors

    Species were the same as Tab. 1. Elv, elevation; Slo, slope degree; Asp, slope aspect; Pos, slope postion; pH, pH value; SOM, soil organic matter; TN, total nitrogen; TP, total phosphorus; TK, total potassium.

    表  1   43种木本植物

    Table  1   43 kinds of woody plants

    编号No. 物种名Species
    S1 猴耳环Pithecellobium clypearia
    S2 山茶Camellia japonica
    S3 麻栎Quercus acutissima
    S4 广西密花树Rapanea kwangsiensis
    S5 黄杞Engelhardtia roxburghiana
    S6 亮叶猴耳环Archidendron lucidum
    S7 罗伞树Ardisia quinquegona
    S8 鹅掌柴Schefflera octophylla
    S9 水锦树Wendlandia uvariifolia
    S10 银柴Aporosa dioica
    S11 九节Psychotria rubra
    S12 龙眼Dimocarpus longan
    S13 枫香Liquidambar formosana
    S14 毛果算盘子Glochidion eriocarpum
    S15 粗叶榕Ficus hirta
    S16 艾胶算盘子Glochidion lanceolarium
    S17 苎麻Boehmeria nivea
    S18 烟斗柯Lithocarpus corneus
    S19 杜茎山Maesa japonica
    S20 粗糠柴Mallotus philippensis
    S21 西南木荷Schima wallichii
    S22 南方荚蒾Viburnum fordiae
    S23 水东哥Saurauia tristyla
    S24 灰毛浆果楝Cipadessa cinerascens
    S25 扁担杆Grewia biloba
    S26 孱槁树Litsea glutinosa
    S27 黄皮Clausena lansium
    S28 三叉苦Evodia lepta
    S29 化香树Platycarya strobilacea
    S30 琴叶榕Ficus pandurata
    S31 蚬木Excentrodendron hsienmu
    S32 榔榆Ulmus parvifolia
    S33 黄牛木Cratoxylum cochinchinense
    S34 岩柿Diospyros dumetorum
    S35 鱼骨木Canthium dicoccum
    S36 澄广花Orophea hainanensis
    S37 九里香Murraya exotica
    S38 菜豆树Radermachera sinica
    S39 山黄皮Clausena excavata
    S40 山石榴Catunaregam spinosa
    S41 扣匹Uvaria tonkinensis
    S42 海南大风子Hydnocarpus hainanensis
    S43 三角榄Canarium bengalense
    下载: 导出CSV

    表  2   麻栎次生林主要木本植物的重要值及生态位宽度

    Table  2   Importance value and niche breadth of dominant woody plants in the Quercus acutissima natural secondary forests

    排序
    Rank
    物种
    Species
    重要值
    Importance value(Ⅳ)
    生态位宽度
    Niche breadth(Bi)
    1 麻栎Quercus acutissima 19.26 0.940
    2 黄杞
    Engelhardtia roxburghiana
    15.44 0.844
    3 罗伞树
    Ardisia quinquegona
    14.19 0.917
    4 猴耳环
    Pithecellobium clypearia
    9.48 0.702
    5 水锦树
    Wendlandia uvariifolia
    8.61 0.944
    6 鹅掌柴
    Schefflera octophylla
    7.98 0.888
    7 亮叶猴耳环
    Archidendron lucidum
    5.85 0.425
    8 广西密花树
    Rapanea kwangsiensis
    3.52 0.859
    9 毛果算盘子
    Glochidion eriocarpum
    2.63 0.812
    10 九节
    Psychotria rubra
    2.56 0.886
    11 烟斗柯
    Lithocarpus corneus
    1.99 0.376
    12 山茶
    Camellia japonica
    1.52 0.400
    13 枫香
    Liquidambar formosana
    1.50 0.613
    14 粗叶榕
    Ficus hirta
    1.43 0.915
    15 银柴
    Aporosa dioica
    1.32 0.725
    下载: 导出CSV

    表  3   枫香次生林主要木本植物的重要值及生态位宽度

    Table  3   Importance value and niche breadth of dominant woody plants in the Liquidambar formosana natural secondary forest

    排序
    Rank
    物种
    Species
    重要值
    Importance value(Ⅳ)
    生态位宽度
    Niche breadth(Bi)
    1 枫香Liquidambar formosana 26.67 0.839
    2 鹅掌柴Schefflera octophylla 12.07 0.440
    3 罗伞树Ardisia quinquegona 10.70 0.557
    4 灰毛浆果楝
    Cipadessa cinerascens
    9.85 0.878
    5 杜茎山Maesa japonica 5.37 0.866
    6 西南木荷
    Schimawallichii Choisy
    3.95 0.539
    7 菜豆树Radermachera sinica 3.69 0.287
    8 粗糠柴Mallotus philippensis 3.08 0.559
    9 黄皮Clausena lansium 2.73 0.594
    10 艾胶算盘子
    Glochidion lanceolarium
    2.00 0.459
    11 九里香Murraya exotica 1.75 0.400
    12 九节Psychotria rubra 1.59 0.442
    13 南方荚蒾Viburnum fordiae 1.34 0.372
    14 水东哥Saurauia tristyla 1.28 0.580
    15 琴叶榕Ficus pandurata 1.25 0.370
    16 黄牛木
    Cratoxylum cochinchinense
    1.25 0.279
    17 粗叶榕Ficus hirta 1.22 0.520
    18 山黄皮
    Clausena excavataBurm
    1.15 0.358
    19 毛果算盘子
    (G.eriocarpum)
    1.14 0.590
    20 山石榴Catunaregam spinosa 1.08 0.329
    21 水锦树
    Wendlandia uvariifolia
    1.01 0.510
    下载: 导出CSV

    表  4   蚬木次生林主要木本植物的重要值及生态位宽度

    Table  4   Importance value and niche breadth of dominant woody plants in the Excentrodendron hsienmu natural secondary forest

    排序
    Rank
    物种
    Species
    重要值
    Importance vlue(Ⅳ)
    生态位宽度
    Niche breadth(Bi)
    1 蚬木
    Excentrodendron hsienmu
    74.63 0.986
    2 澄广花Orophea hainanensis 8.77 0.677
    3 鱼骨木Canthium dicoccum 3.35 0.660
    4 岩柿Diospyros dumetorum 2.62 0.333
    5 榔榆Ulmus parvifolia 2.57 0.333
    6 九里香Murraya exotica 2.08 0.507
    7 鹅掌柴Schefflera octophylla 1.44 0.333
    8 黄牛木
    Cratoxylum cochinchinense
    1.31 0.333
    9 菜豆树Radermachera sinica 1.11 0.664
    下载: 导出CSV

    表  5   麻栎次生林主要木本植物生态位重叠

    Table  5   Niche overlap of dominant woody plants in the Quercus acutissima natural secondary forest

    物种序号
    Species No.
    1 2 3 4 5 6 7 8 9 10 11 12 13 14 15
    1 1.000
    2 0.811 1.000
    3 0.981 0.853 1.000
    4 0.873 0.580 0.762 1.000
    5 0.996 0.802 0.961 0.911 1.000
    6 0.962 0.862 0.997 0.708 0.936 1.000
    7 0.447 0.849 0.465 0.350 0.466 0.469 1.000
    8 0.947 0.859 0.991 0.670 0.916 0.999 0.461 1.000
    9 0.830 0.797 0.748 0.903 0.871 0.708 0.716 0.673 1.000
    10 0.910 0.944 0.964 0.621 0.883 0.976 0.628 0.978 0.716 1.000
    11 0.691 0.282 0.540 0.945 0.742 0.472 0.090 0.427 0.753 0.341 1.000
    12 0.424 0.839 0.445 0.324 0.442 0.451 1.000 0.443 0.697 0.613 0.064 1.000
    13 0.760 0.579 0.631 0.959 0.816 0.571 0.498 0.527 0.951 0.527 0.907 0.476 1.000
    14 0.882 0.990 0.919 0.646 0.869 0.925 0.768 0.922 0.806 0.978 0.360 0.755 0.609 1.000
    15 0.704 0.910 0.678 0.665 0.731 0.661 0.932 0.641 0.918 0.754 0.432 0.921 0.763 0.873 1.000
    注:物种序号同表 2。Note:species No. was the same as Tab. 2
    下载: 导出CSV

    表  6   枫香次生林主要木本植物生态位重叠

    Table  6   Niche overlap of dominant woody plants in the Liquidambar formosana natural secondary forest

    物种序号
    SpeciesNo.
    1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21
    1 1.000
    2 0.439 1.000
    3 0.604 0.221 1.000
    4 0.927 0.453 0.778 1.000
    5 0.806 0.727 0.612 0.753 1.000
    6 0.549 0.973 0.237 0.507 0.834 1.000
    7 0.385 0.000 0.960 0.597 0.405 0.000 1.000
    8 0.626 0.283 0.722 0.714 0.750 0.409 0.645 1.000
    9 0.919 0.283 0.474 0.901 0.558 0.360 0.277 0.515 1.000
    10 0.468 0.234 0.891 0.676 0.619 0.281 0.876 0.906 0.360 1.000
    11 0.486 0.850 0.211 0.344 0.805 0.897 0.000 0.248 0.180 0.142 1.000
    12 0.615 0.658 0.206 0.400 0.868 0.799 0.000 0.459 0.313 0.219 0.901 1.000
    13 0.652 0.392 0.173 0.430 0.789 0.594 0.000 0.630 0.437 0.293 0.623 0.896 1.000
    14 0.636 0.282 0.764 0.695 0.793 0.415 0.690 0.984 0.465 0.906 0.329 0.534 0.664 1.000
    15 0.581 0.574 0.189 0.341 0.826 0.722 0.000 0.414 0.261 0.178 0.878 0.992 0.890 0.505 1.000
    16 0.715 0.000 0.527 0.721 0.242 0.000 0.416 0.134 0.790 0.182 0.000 0.000 0.000 0.144 0.000 1.000
    17 0.671 0.754 0.291 0.489 0.833 0.826 0.059 0.268 0.386 0.136 0.956 0.910 0.674 0.356 0.899 0.257 1.000
    18 0.514 0.000 0.967 0.699 0.413 0.000 0.974 0.595 0.439 0.808 0.000 0.000 0.000 0.636 0.000 0.612 0.115 1.000
    19 0.633 0.948 0.314 0.592 0.784 0.939 0.063 0.260 0.463 0.200 0.885 0.716 0.443 0.283 0.650 0.275 0.878 0.123 1.000
    20 0.378 0.087 0.922 0.561 0.535 0.130 0.952 0.798 0.214 0.954 0.116 0.178 0.212 0.843 0.172 0.198 0.125 0.877 0.092 1.000
    21 0.805 0.365 0.263 0.705 0.701 0.528 0.066 0.715 0.796 0.392 0.335 0.607 0.819 0.662 0.551 0.290 0.441 0.130 0.423 0.196 1.000
    注:物种序号同表 3。Note:species No. was the same as Tab. 3
    下载: 导出CSV

    表  7   蚬木次生林主要木本植物生态位重叠

    Table  7   Niche overlap of dominant woody plants in the Excentrodendron hsienmu natural secondary forest

    物种序号
    Species No.
    1 2 3 4 5 6 7 8 9
    1 1.000
    2 0.783 1.000
    3 0.810 0.948 1.000
    4 0.493 0.438 0.181 1.000
    5 0.493 0.438 0.181 1.000 1.000
    6 0.770 0.209 0.324 0.271 0.271 1.000
    7 0.493 0.438 0.181 1.000 1.000 0.271 1.000
    8 0.493 0.438 0.181 1.000 1.000 0.271 1.000 1.000
    9 0.870 0.664 0.838 0.708 0.000 0.720 0.000 0.000 1.000
    注:物种序号同表 4。Note: species No. was the same as Tab. 4.
    下载: 导出CSV

    表  8   桂西南喀斯特山地木本植物群落的总体关联性

    Table  8   Overall interspecific associations within woody plant communities in the karst mountains of Southwest Guangxi

    林分类型
    Forest type
    方差比率
    Variance ratio(VR)
    检验统计量Test
    statistics
    (χ 0.952 Nx 2 0.05N) 测度结果
    Measurement result
    麻栎次生林
    Quercus acutissima natural secondary forest
    1.004 48.172 (33.098,65.171) 不显著正关联No significantly positive correlation
    枫香次生林
    Liquidambar formosana natural secondary forest
    0.905 72.370 (60.391,101.879) 不显著负关联
    No significantly negative correlation
    蚬木次生林
    Excentrodendron hsienmu natural secondary forest
    1.689 81.072 (33.098,65.171) 显著正关联
    Significantly positive correlation
    下载: 导出CSV

    表  9   环境因子与CCA排序轴的相关系数

    Table  9   Correlation coefficients between environmental factors and CCA ordination axes

    项目Item 第1轴Axis 1 第2轴Axis 2 第3轴Axis 3 第4轴Axis 4
    海拔Elevation, Elv -0.294 9 -0.702 9* -0.518 5 -0.293 4
    坡度Slope degree, Slo 0.117 4 -0.641 3* 0.626 0* -0.308 0
    坡向Slope aspect, Asp -0.004 4 -0.904 4 *** -0.338 3 -0.104 6
    坡位Slope position, Pos 0.336 3 0.108 2 -0.268 5 -0.011 3
    pH值pH value, pH 0.486 2 -0.717 5 ** 0.377 4 0.032 2
    土壤有机质Soil organic matter, SOM 0.344 2 -0.355 2 0.654 9* -0.324 5
    全氮Total nitrogen, TN 0.362 2 -0.420 5 0.619 3* -0.311 5
    全磷Total phosphorus, TP 0.598 8* -0.475 3 0.575 3 -0.059 6
    全钾Total potassium, TK -0.603 4* 0.445 2 -0.576 7* 0.091 7
    特征值Eigenvalue 0.763 0.557 0.293 0.256
    物种-环境相关Species-environment correlation 0.995 0.996 0.999 0.997
    物种环境关系的方差累积比例/% Cumulative percentage variance of species-environment 31.43 54.36 66.42 76.95
    所有典范轴的显著性检验Test of significance of all canonical axes 0.002 0
    注:*表示在P<0.05水平上显著相关, **表示在P<0.01水平上极显著相关, ***表示在P<0.001水平上极显著相关。Notes: * means correlation is significant at P<0.05 level; **means correlation is significant at P<0.01 level; ***means correlation is significant at P<0.001 level.
    下载: 导出CSV
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