Interspecific relationship and canonical correspondence analysis within woody plant communities in the karst mountains of Southwest Guangxi, southern China
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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。
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关键词:
- 喀斯特 /
- 生态位 /
- 种间联结性 /
- 双向指示种分析(TWINSPAN) /
- 典范对应分析(CCA)
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. -
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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 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 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 roxburghiana15.44 0.844 3 罗伞树
Ardisia quinquegona14.19 0.917 4 猴耳环
Pithecellobium clypearia9.48 0.702 5 水锦树
Wendlandia uvariifolia8.61 0.944 6 鹅掌柴
Schefflera octophylla7.98 0.888 7 亮叶猴耳环
Archidendron lucidum5.85 0.425 8 广西密花树
Rapanea kwangsiensis3.52 0.859 9 毛果算盘子
Glochidion eriocarpum2.63 0.812 10 九节
Psychotria rubra2.56 0.886 11 烟斗柯
Lithocarpus corneus1.99 0.376 12 山茶
Camellia japonica1.52 0.400 13 枫香
Liquidambar formosana1.50 0.613 14 粗叶榕
Ficus hirta1.43 0.915 15 银柴
Aporosa dioica1.32 0.725 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 cinerascens9.85 0.878 5 杜茎山Maesa japonica 5.37 0.866 6 西南木荷
Schimawallichii Choisy3.95 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 lanceolarium2.00 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 cochinchinense1.25 0.279 17 粗叶榕Ficus hirta 1.22 0.520 18 山黄皮
Clausena excavataBurm1.15 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 uvariifolia1.01 0.510 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 hsienmu74.63 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 cochinchinense1.31 0.333 9 菜豆树Radermachera sinica 1.11 0.664 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 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 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. 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 N,x 2 0.05N) 测度结果
Measurement result麻栎次生林
Quercus acutissima natural secondary forest1.004 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 forest0.905 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 forest1.689 81.072 (33.098,65.171) 显著正关联
Significantly positive correlation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. -
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