Effects of stand density on community structure and species diversity of Cupressus funebris plantation in Yunding Mountain, southwestern China
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摘要:
目的探索林分密度对柏木人工林群落结构和林下物种多样性及林下物种多样性与林分因子相关性的影响，为云顶山柏木人工林经营管理提供依据。 方法以四川云顶山5种不同林分密度（A ~ E：500、650、800、950、1 100株/hm2）60 年生柏木人工林为研究对象，采用典型样地法进行植被调查，综合分析其群落结构、物种组成和物种多样性指数（Pielou均匀度指数Jsw、Simpson优势度指数H′、Shannon-Wiener多样性指数H、和物种丰富度指数D）。 结果（1）研究区内共记录植物170种，隶属于68科136属。不同密度下灌木层或草本层优势种多属阴性、耐阴性或适应性强的植物。（2）随着林分密度的减小，灌木层物种多样性指数均呈先增后减的单峰变化，基本在密度B达到最大，除D外均无显著差异（P > 0.05）；草本层D、H、H′则呈先增后减再增再减的双峰变化，在密度B和D出现峰值，不同密度间各指数差异性显著。林分密度和郁闭度与灌木层D呈极显著负相关，与草本层H 、H′、Jsw呈显著或极显著正相关。（3）密度A群落径级和高度级结构均呈波动状不规则分布，其余密度下均呈单峰型分布。密度B群落内中、大乔木个体占比相对较多，不同大小个体数在群落中分布趋于合理，群落稳定性较好。 结论650株/hm2为云顶山柏木人工林的相对最适林分密度，利于维持群落结构稳定并提高林下物种多样性。 Abstract:ObjectiveThe effects of stand density on the community structure and species diversity of Cupressus funebris plantation and the correlation between species diversity and stand factors were explored, providing a basis for the management of Cupressus funebris plantation in Yunding Mountains, southwestern China. MethodThe research objects were five different stand densities (A−E: 500, 650, 800, 950, 1 100 tree/ha) of 60 years Cupressus funebris in Yunding Mountain, Sichuan Province, southwestern China. The vegetation was investigated by typical plot method. The community structure, species composition and species diversity index (Pielou evenness index Jsw, Simpson dominance index H', Shannon-Wiener diversity index H and species richness index D) were comprehensively analyzed. Result(1) A total of 170 species of plants were recorded in the study area, belonging to 136 genera and 68 families. The dominant species in shrub layer or herb layer under different densities were mostly shady, shade-tolerant or adaptable plants. (2) With the decrease of stand density, the trend of species diversity index in shrub layer increased first and then decreased. It reached the maximum at density B, with no significant difference except D (P > 0.05). The trend of D, H and H' in herbaceous layer increased first, then decreased, then increased and then decreased. The peak values appeared in density B and D, and there were significant differences among different densities. Stand density and canopy density were negatively correlated with shrub layer D, and significantly positively correlated with herb layer H, H' and Jsw . (3) Distribution between diameter and height structure of density A community was irregular and fluctuated, while distribution of other densities was unimodal. In density B community, medium and large individual trees accounted for a relatively large proportion of individuals, and the stability of the community was better. ConclusionThe relative optimum stand density of cypress plantation in Yunding Mountain is 650 tree/ha, which is very helpful to maintain the stability of community structure and improve the species diversity under the forest. -
Key words:
- stand density /
- Cupressus funebris plantation /
- community structure /
- species diversity
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图 1 不同密度柏木林乔木层径级和高度级结构
同径级（高度级）不同字母表示不同密度在该径级（高度级）个体数差异显著（P < 0.05）。Different letters appearing at the same diameter class (height class) indicate that the individual numbers with different densities at this diameter class (height class) are significantly different (P < 0.05).
Figure 1. Diameter and height structure of different densities of C. funebris plantation
图 2 不同密度柏木林灌草层物种组成
F为科，G为属，S为种。F，family；G，genera；S，species.
Figure 2. Species composition of shrub and grass layers of C. funebris plantation with different densities
表 1 样地基本情况
Table 1. Basic situation of the survey plots
样地号
Sample plot No.密度/（株·hm−2）
Density/(tree·ha− 1)密度划分
Density class海拔
Altitude/m坡向
Slope aspect坡度
Slope degree/(°)9 ~ 12 500 A 827 ~ 867 正南 Due south 25 ~ 28 1 ~ 4 650 B 858 ~ 861 东北48° ~ 60° NE48°−60° 19 ~ 26 5 ~ 8 800 C 835 ~ 841 正东 ~ 东北66° Due east−NE66° 20 ~ 25 17 ~ 20 950 D 833 ~ 835 正南 Due south 21 ~ 24 13 ~ 16 1 100 E 830 ~ 833 正南 Due south 25 ~ 27 
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表 2 不同密度柏木林灌草层优势种重要值
Table 2. Important values of dominant species of shrub and grass layers in different density C. funebris plantations
层次
Layer密度
Density优势种
Dominant species重要值
Important value灌木层
Shrub layerA 黄荆 + 石海椒 + 豆腐柴 + 六道木 + 荚蒾
Vitex negundo + Reinwardtia indica + Premna microphylla + Abelia biflora + Viburnum dilatatum0.226 1 + 0.195 6 + 0.070 7 + 0.067 4 + 0.062 9 B 石海椒 + 珊瑚朴 + 荚蒾 + 黄荆 + 豆腐柴
Reinwardtia indica + Celtis julianae + Viburnum dilatatum + Vitex negundo + Premna microphylla0.192 8 + 0.105 6 + 0.084 1 + 0.075 0 + 0.066 4 C 黄荆 + 石海椒 + 六道木 + 荚蒾 + 豆腐柴
Vitex negundo + Reinwardtia indica + Abelia biflora + Viburnum dilatatum + Premna microphylla0.262 4 + 0.135 6 + 0.083 5 + 0.080 4 + 0.069 5 D 黄荆 + 石海椒 + 荚蒾 + 蕊帽忍冬 + 珊瑚朴
Vitex negundo + Reinwardtia indica + Viburnum dilatatum + Lonicera pileata + Celtis julianae0.360 2 + 0.099 7 + 0.097 3 + 0.096 0 + 0.058 8 E 石海椒 + 六道木 + 烟管荚蒾 + 黄荆 + 豆腐柴
Reinwardtia indica + Abelia biflora + Viburnum utile + Vitex negundo + Premna microphylla0.212 7 + 0.197 7 + 0.122 5 + 0.108 5 + 0.052 6 草本层
Herb layerA 莩草 + 求米草 + 韩信草 + 过路黄 + 白接骨
Setaria chondrachne + Oplismenus undulatifolius + Scutellaria indica + Lysimachia christinae + Asystasiella neesiana0.226 7 + 0.136 8 + 0.120 3 + 0.098 3 + 0.082 1 B 过路黄 + 白接骨 + 莩草 + 麦冬 + 求米草
Lysimachia christinae + Asystasiella neesiana + Setaria chondrachne + Ophiopogon japonicus + Oplismenus undulatifolius0.136 1 + 0.133 1 + 0.119 4 + 0.112 3 + 0.094 4 C 莩草 + 韩信草 + 白接骨 + 过路黄 + 爵床
Setaria chondrachne + Scutellaria indica+ Asystasiella neesiana + Lysimachia christinae + Rostellularia procumbens 0.372 8 + 0.142 7 + 0.133 6 + 0.111 9 + 0.082 6 D 莩草 + 白接骨 + 三花莸 + 麦冬 + 求米草
Setaria chondrachne + Asystasiella neesiana + Caryopteris terniflora + Ophiopogon japonicus + Oplismenus undulatifolius0.177 9 + 0.116 5 + 0.059 6 + 0.045 6 + 0.043 3 E 沿阶草 + 莩草 + 过路黄 + 求米草 + 韩信草
Ophiopogon bodinieri + Setaria chondrachne + Lysimachia christinae + Oplismenus undulatifolius + Scutellaria indica0.132 2 + 0.121 4 + 0.114 3 + 0.110 9 + 0.093 4
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表 3 不同密度柏木林灌草层物种多样性指数
Table 3. Species diversity indices of shrub and grass layers in C. funebris plantation under different densities
层次
Layer密度
DensityPielou均匀度指数
Pielou eveness index (Jsw)Simpson优势度指数
Simpson dominance index (H')Shannon-Wiener指数
Shannon-Wiener index (H)物种丰富度指数
Species richness index (D)灌木层
Shrub layerA 0.615 3 ± 0.021 5a 0.758 5 ± 0.029 8a 1.989 0 ± 0.050 3a 25.500 0 ± 1.040 8ab B 0.671 1 ± 0.064 4a 0.764 3 ± 0.057 3a 2.253 6 ± 0.196 8a 29.000 0 ± 0.912 9a C 0.681 5 ± 0.076 5a 0.778 3 ± 0.081 8a 2.158 6 ± 0.274 8a 23.500 0 ± 1.707 8b D 0.657 3 ± 0.037 9a 0.767 5 ± 0.023 7a 1.878 4 ± 0.112 7a 17.500 0 ± 0.866 0c E 0.600 8 ± 0.063 3a 0.670 6 ± 0.074 6a 1.715 2 ± 0.194 4a 17.250 0 ± 0.478 7c 草本层
Herb layerA 0.748 7 ± 0.047 7c 0.849 9 ± 0.025 7b 2.192 3 ± 0.146 8b 18.750 0 ± 1.030 8c B 0.752 4 ± 0.036 1c 0.878 4 ± 0.015 1ab 2.536 6 ± 0.114 6b 29.250 0 ± 1.108 7a C 0.768 6 ± 0.029 1bc 0.868 5 ± 0.019 5b 2.384 2 ± 0.107 5b 22.250 0 ± 0.853 9bc D 0.943 7 ± 0.004 8a 0.942 8 ± 0.002 8a 3.049 9 ± 0.059 3a 25.500 0 ± 1.707 8ab E 0.882 3 ± 0.012 3ab 0.904 1 ± 0.002 6ab 2.547 8 ± 0.024 1b 18.000 0 ± 0.577 4c 注：同列含有不同字母表示该列指数不同密度间差异显著（P < 0.05）。Note: same column containing different letters indicate that the index of this column is significantly different between different densities (P < 0.05).
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表 4 灌草层物种多样性指数与林分因子的相关性
Table 4. Correlation coefficients between understory species diversity index and stand factors
林分因子
Stand factor密度 Density 郁闭度 Canopy density 平均树高 Mean tree height 平均胸径 Mean DBH 平均冠幅 Mean crown width 草本层 Herb layer 灌木层 Shrub layer D H H' Jsw D H H' Jsw 密度
Density1.00 郁闭度
Canopy density0.97** 1.00 平均树高
Mean tree height−0.31 0.29 1.00 平均胸径
Mean DBH−0.17 −0.18 0.65** 1.00 平均冠幅
Mean crown width−0.05 −0.11 0.01 0.15 1.00 HL-D −0.16 −0.13 0.20 0.31 −0.22 1.00 HL-H 0.52* 0.53* 0.16 0.13 −0.04 0.44* 1.00 HL-H' 0.58** 0.58** 0.07 0.00 −0.07 0.20 0.93** 1.00 HL-J 0.69** 0.67** 0.07 −0.03 0.04 −0.05 0.87** 0.94** 1.00 SH-D −0.80** −0.78** 0.07 0.31 −0.17 0.36 −0.49* 0.60** 0.74** 1.00 SH-H −0.35 −0.35 −0.12 −0.11 −0.22 0.11 −0.31 −0.33 −0.42 0.52* 1.00 SH-H’ −0.23 −0.23 −0.06 −0.18 −0.17 −0.01 −0.17 −0.18 −0.21 0.26 0.91** 1.00 SH-J −0.06- −0.07 −0.13 −0.22 −0.18 0.00 −0.12 −0.10 −0.14 0.17 0.93** 0.94** 1.00 注：*表示相关性显著（P < 0.05），**表示相关性极显著（P < 0.01）。HL表示草本层，SH表示灌木层。Notes: * indicates significant correlation (P < 0.05), ** indicates that the correlation is extremely significant (P < 0.01). HL means herb layer, SH means shrub layer.
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