Biodiversity Science ›› 2019, Vol. 27 ›› Issue (9): 970-983.doi: 10.17520/biods.2019133

• Original Papers • Previous Article     Next Article

Effect of environmental factors on understory species diversity in Southwest Guangxi Excentrodendron tonkinense forests

Yibo Tan1, 2, 3, Wenhui Shen1, 2, 3, *(), Zi Fu4, Wei Zheng1, 2, 3, Zhiyang Ou1, 2, 3, Zhangqiang Tan1, 2, 3, Yuhua Peng1, 2, 3, Shilong Pang1, 2, 3, Qinfei He1, 2, 3, Xiaorong Huang1, 2, 3, Feng He1, 2, 3   

  1. 1 Guangxi Zhuang Autonomous Region Forestry Research Institute, Nanning 530002
    2 Guangxi Key Laboratory of Superior Timber Trees Resource Cultivation, Nanning 530002
    3 Guangxi Lijiangyuan Forest Ecosystem Research Station, Guilin, Guangxi 541316
    4 Office of Converting Farmland to Forestry, Guangxi Forestry Bureau, Nanning 530028
  • Received:2019-04-18 Accepted:2019-09-11 Online:2019-09-25
  • Shen Wenhui E-mail:769586022@qq.com

Exploring how distribution patterns of understory vegetation are affected by environmental factors is of great importance for natural forest protection and biodiversity conservation. Natural and mature Excentrodendron tonkinense-dominated forests on karst terrains were studied in eight counties of Southwest Guangxi. One-way ANOVA, Pearson correlation and redundancy analysis (RDA) were used to investigate the variation in species diversity and understory structure along a gradient of environmental factors including soil, topography, canopy structure and light. The results showed that the shrub layer was dominated by Sophora tonkinensis, Schefflera heptaphylla and Ventilago calyculata, as well as seedlings of species from the tree layer, such as Excentrodendron tonkinense, Orophea anceps, Cinnamomum saxatile and Garcinia paucinervis, etc. These species mainly represented the Leguminosae, Araliaceae, Rhamnaceae, Tiliaceae, Annonaceae, Lauraceae, Guttiferae and Euphorbiaceae families. Within the herb layer, the dominant species were Nephrolepis cordifolia, Guihaia argyrata, Pseudodrynaria coronans, Microstegium vimineum, Apluda mutica and Ophiopogon bodinieri, which mainly represent the Nephrolepidaceae, Palmae, Drynariaceae, Gramineae, Liliaceae, Aspleniaceae and Dryopteridaceae families. Soil pH, soil water content (SWC), soil total potassium (TK), soil total phosphorus (TP) and slope (SLO) were the main influencing factors, respectively explaining the variation in understory species diversity by 32.3%, 16.1%, 9.7%, 8.6% and 8.6%. Shrub richness and the shrub diversity index were significantly and negatively correlated with TK, SWC, soil pH and TP, while herb richness and the herb diversity index were significantly and positively correlated with TK. Shrub density and shrub coverage were positively linked to soil pH (P < 0.05), and herb density was positively associated with SWC and TK (P < 0.05). Herb coverage was positively correlated with TP and TK, and negatively correlated with slope (P < 0.05). Soil and topography were the most important factors affecting species diversity in the understory, while canopy structure had little effect on it. Edaphic factors were found to affect understory species diversity more than topographical factors.

Key words: canopy structure, redundancy analysis, soil nutrients, topographical factors, understory species diversity

Table 1

Stand characteristics of eight Excentrodendron tonkinense community sampling plots in karst area of Southwest Guangxi (mean ± SE)"

样地名称
Sample plots
纬度
Latitude
经度
Longitude
海拔
Altitude (m)
坡度
Slope
坡向指数
Transformation of aspect
平均胸径
Mean DBH
(cm)
平均树高
Mean height
(m)
大新县 Daxin County 22°49′ N 106°46′ E 400 43° 0.97 4.93 ± 0.24 4.72 ± 0.12
靖西市 Jingxi City 22o53′ N 106o26′ E 357 40° 0.10 4.89 ± 0.30 5.65 ± 0.19
龙州县 Longzhou County 22o27′ N 106o56′ E 208 46° 0.98 5.30 ± 0.36 5.56 ± 0.19
隆安县 Longan County 23o4′ N 107o24′ E 409 42° 0.39 4.66 ± 0.29 5.48 ± 0.24
那坡县 Napo County 23°0′ N 105°52′ E 823 23° 0.77 12.49 ± 1.73 7.35 ± 0.52
平果县 Pingguo County 23°24′ N 107°32′ E 350 35° 0.97 3.65 ± 0.10 3.72 ± 0.05
天等县 Tiandeng County 23o8′ N 107o16′ E 418 40° 0.43 7.48 ± 0.53 7.77 ± 0.25
武鸣区 Wuming District 23o16′ N 107o51′ E 271 35° 0.98 10.34 ± 0.79 8.38 ± 0.42

Table 2

Information on environment factors of eight Excentrodendron tonkinense community sampling plots in karst area of Southwest Guangxi (mean ± SE). Different letters within the same row meant significant difference among the sampling plots at 0.05 level."

大新县
Daxin County
靖西市
Jingxi City
龙州县
Longzhou County
隆安县
Longan County
那坡县
Napo County
平果县
Pingguo County
天等县
Tiandeng County
武鸣区
Wuming District
土壤pH Soil pH 6.66 ± 0.25e 7.49 ± 0.11abc 7.61 ± 0.10ab 7.11 ± 0.08d 7.30 ± 0.02bcd 6.59 ± 0.02e 7.24 ± 0.09cd 7.76 ± 0.02a
土壤有机质
Soil organic matter (g/kg)
124.17 ± 18.07cd 156.42 ± 21.96bc 97.15 ± 7.06d 144.18 ± 6.59bc 369.20 ± 9.79a 131.96 ± 10.62bcd 174.81 ± 20.34b 172.66 ± 5.77b
土壤全氮
Soil total nitrogen (g/kg)
7.64 ± 1.02b 8.82 ± 1.08b 5.11 ± 0.26c 6.81 ± 0.28bc 22.38 ± 0.40a 7.26 ± 0.63b 8.37 ± 0.77b 7.75 ± 0.29b
土壤全磷
Soil total phosphorus (g/kg)
1.75 ± 0.08bc 1.88 ± 0.12b 0.88 ± 0.05ef 0.62 ± 0.03f 6.47 ± 0.36a 1.90 ± 0.07b 1.26 ± 0.06de 1.37 ± 0.09cd
土壤全钾
Soil total potassium (g/kg)
1.70 ± 0.24d 7.81 ± 0.86a 2.29 ± 0.16cd 3.24 ± 0.49bc 3.34 ± 0.23bc 3.11 ± 0.64bcd 2.38 ± 0.25cd 4.37 ± 0.26b
土壤C : N
Soil C : N
9.42 ± 0.34c 10.24 ± 0.24bc 11.01 ± 0.32b 12.27 ± 0.10a 9.57 ± 0.11c 10.55 ± 0.21b 12.07 ± 0.32a 12.94 ± 0.47a
土壤N : P
Soil N : P
4.33 ± 0.44cd 4.68 ± 0.39cd 5.86 ± 0.68bc 11.04 ± 0.65a 3.48 ± 0.13d 3.85 ± 0.48d 6.74 ± 0.89b 5.68 ± 0.16bc
土壤含水量
Soil water content (g/g)
0.40 ± 0.09b 0.48 ± 0.06b 0.35 ± 0.03b 0.40 ± 0.02b 1.44 ± 0.06a 0.41 ± 0.03b 0.44 ± 0.03b 0.47 ± 0.04b
林冠开阔度
Canopy
openness (%)
17.24 ± 0.41a 14.20 ± 0.31b 13.27 ± 0.88b 13.68 ± 1.42b 15.89 ± 1.78ab 13.46 ± 0.23b 12.79 ± 0.54b 13.83 ± 0.40b
叶面积指数
Leaf area index
2.13 ± 0.02b 2.40 ± 0.05ab 2.50 ± 0.09a 2.54 ± 0.08a 2.34 ± 0.26ab 2.40 ± 0.08ab 2.37 ± 0.02ab 2.31 ± 0.07ab
林内直射光 Transmitted direct in forest (mol·m-2·d-1) 4.42 ± 0.16a 3.08 ± 0.27b 3.20 ± 0.33b 2.96 ± 0.24b 3.28 ± 0.45b 3.16 ± 0.25b 3.47 ± 0.11b 3.55 ± 0.32b
林内散射光 Transmitted diffuse in forest (mol·m-2·d-1) 4.03 ± 0.05a 3.35 ± 0.15b 3.10 ± 0.13b 2.92 ± 0.16b 3.45 ± 0.43b 3.12 ± 0.15b 3.17 ± 0.05b 3.36 ± 0.13b
林内总光照
Transmitted
total in forest (mol·m-2·d-1)
8.45 ± 0.21a 6.43 ± 0.39b 6.30 ± 0.39b 5.88 ± 0.34b 6.73 ± 0.88b 6.28 ± 0.40b 6.64 ± 0.09b 6.91 ± 0.41b
消光系数Extinction coefficient (k) 0.72 ± 0.02a 0.76 ± 0.02a 0.74 ± 0.03a 0.75 ± 0.01a 0.77 ± 0.04a 0.76 ± 0.01a 0.75 ± 0.01a 0.75 ± 0.01a

Table 3

Composition and importance value (IV) of dominant species in eight Excentrodendron tonkinense community sampling plots in karst area of Southwest Guangxi. T, Tree layer; S, Shrub layer; H, Herb layer."

样地
Plot
物种
Species
层次
Layer
重要值
IV (%)
样地
Plot
物种
Species
层次 Layer 重要值 IV (%)
大新县
Daxin County
蚬木 Excentrodendron tonkinense T 58.84 那坡县
Napo
County
蚬木 Excentrodendron tonkinense T 33.10
广西澄广花 Orophea anceps T 7.29 苹婆 Sterculia monosperma T 8.90
岩柿 Diospyros dumetorum T 3.46 樟叶槭 Acer coriaceifolia T 5.59
榔榆 Ulmus parvifolia T 3.36 Toxicodendron vernicifluum T 5.41
蚬木 Excentrodendron tonkinense S 13.89 鹅掌柴 Schefflera heptaphylla S 26.90
红背山麻杆 Alchornea trewioides S 12.52 蚬木 Excentrodendron tonkinense S 19.93
光叶紫玉盘 Uvaria boniana S 7.78 香蒲桃 Syzygium odoratum S 5.71
鱼骨木 Canthium dicoccum S 6.66 香叶树 Lindera communis S 4.49
石山棕 Guihaia argyrata H 27.03 巢蕨 Neottopteris nidus H 14.46
江南卷柏 Selaginella moellendorffii H 13.09 贯众 Cyrtomium fortunei H 10.34
槲蕨 Drynaria roosii H 9.37 肾蕨 Nephrolepis cordifolia H 8.77
铁角蕨 Asplenium trichomanes H 8.92 艾麻 Laportea cuspidata H 8.76
靖西市
Jingxi City
蚬木 Excentrodendron tonkinense T 33.69 平果县Pingguo County 蚬木 Excentrodendron tonkinense T 11.02
岩樟 Cinnamomum saxatile T 17.76 黄荆 Vitex negundo T 8.76
苹婆 Sterculia monosperma T 5.18 清香木 Pistacia weinmannifolia T 6.53
广西澄广花 Orophea anceps T 1.58 岩柿 Diospyros dumetorum T 5.53
苎麻 Boehmeria nivea S 48.41 黄荆 Vitex negundo S 21.05
岩樟 Cinnamomum saxatile S 18.45 蚬木 Excentrodendron tonkinense S 15.33
蚬木 Excentrodendron tonkinense S 13.92 薄皮酒饼簕 Atalantia henryi S 10.25
老虎刺 Pterolobium punctatum S 9.99 山石榴 Catunaregam spinosa S 3.68
肾蕨 Nephrolepis cordifolia H 11.10 柔枝莠竹 Microstegium vimineum H 50.02
蔓生莠竹 Microstegium fasciculatum H 10.26 水蔗草 Apluda mutica H 19.18
苦苣菜 Sonchus oleraceus H 5.91 假鞭叶铁线蕨 Adiantum malesianum H 7.75
艾麻 Laportea cuspidata H 3.78 硬叶兰 Cymbidium bicolor H 4.87
龙州县Longzhou County 蚬木 Excentrodendron tonkinense T 25.22 天等县Tiandeng County 蚬木 Excentrodendron tonkinense T 43.08
广西澄广花 Orophea anceps T 11.38 金丝李 Garcinia paucinervis T 10.74
金丝李 Garcinia pauc T 11.28 异叶榕 Ficus heteromorpha T 4.48
假肥牛树 Cleistanthus petelotii T 4.23 广西澄广花 Orophea anceps T 4.24
广西澄广花 Orophea anceps S 21.26 毛果翼核果 Ventilago calyculata S 17.39
九里香 Murraya exotica S 10.89 异叶榕 Ficus heteromorpha S 7.48
金丝李 Garcinia paucinervis S 7.29 海红豆 Adenanthera pavonina S 7.25
牛筋藤 Malaisia scandens S 6.95 金丝李 Garcinia paucinervis S 4.73
崖姜 Pseudodrynaria coronans H 9.10 肾蕨 Nephrolepis cordifolia H 20.28
鞭叶蕨 Cyrtomidictyum lepidocaulon H 5.56 狭叶沿阶草 Ophiopogon stenophyllus H 9.57
竹叶草 Oplismenus compositus H 5.31 槲蕨 Drynaria roosii H 8.98
鞭叶铁线蕨 Adiantum caudatum H 4.37 石韦 Pyrrosia lingua H 8.64
隆安县Longan County 蚬木 Excentrodendron tonkinense T 46.81 武鸣区Wuming District 蚬木 Excentrodendron tonkinense T 69.88
九里香 Murraya exotica T 5.98 苹婆 Sterculia monosperma T 8.42
肥牛树 Cephalomappa sinensis T 4.65 菜豆树 Radermachera sinica T 5.07
青檀 Pteroceltis tatarinowii T 4.58 黄荆 Vitex negundo T 3.29
越南槐 Sophora tonkinensis S 23.65 越南槐 Sophora tonkinensis S 34.65
九里香 Murraya exotica S 10.05 红背山麻杆 Alchornea trewioides S 18.87
肥牛树 Cephalomappa sinensis S 5.55 潺槁木姜子 Litsea glutinosa S 5.45
红鳞蒲桃 Syzygium hancei S 4.05 黄荆 Vitex negundo S 3.47
沿阶草 Ophiopogon bodinieri H 47.05 肾蕨 Nephrolepis cordifolia H 48.21
褐果薹草 Carex brunnea H 14.07 艾麻 Laportea cuspidata H 6.70
肾蕨 Nephrolepis cordifolia H 6.57 鞭叶铁线蕨 Adiantum caudatum H 5.46
广西前胡 Peucedanum guangxiense H 4.35 硬叶兰 Cymbidium bicolor H 1.99

Table 4

Variance analysis of understory species diversity in eight Excentrodendron tonkinense community sampling plots in karst area of Southwest Guangxi. * P < 0.05, ** P < 0.01."

林下植被物种多样性 Understory species diversity 平方和 Sum of squares 均方 Mean squares F
灌木丰富度 Shrub richness (S_shrub) 502.625 71.804 4.97**
灌木多样性 Shrub diversity (SW_shrub) 10.455 1.494 6.08**
灌木密度 Shrub density (D_shrub) 3,728,711.958 532,673.137 11.19**
灌木盖度 Shrub coverage (C_shrub) 2,969.312 424.187 4.93**
草本丰富度 Herb richness (S_herb) 256.000 36.571 3.46*
草本多样性 Herb diversity (SW_herb) 7.880 1.126 2.46
草本密度 Herb density (D_herb) 167,753,218.958 23,964,745.565 10.51**
草本盖度 Herb coverage (C_herb) 11,636.273 1,662.325 20.44**

Table 5

Correlation coefficients of understory species diversity in eight Excentrodendron tonkinense community sampling plots in karst area of Southwest Guangxi. * P < 0.05, ** P < 0.01."

灌木多样性
Shrub diversity
灌木密度
Shrub density
灌木盖度
Shrub coverage
草本丰富度
Herb richness
草本多样性
Herb diversity
草本密度
Herb density
草本盖度
Herb coverage
灌木丰富度
Shrub richness
0.867** 0.063 0.297 -0.327 -0.268 -0.619** -0.706**
灌木多样性
Shrub diversity
0.103 0.293 -0.25 -0.049 -0.576** -0.660**
灌木密度
Shrub density
0.738** -0.314 -0.111 -0.076 -0.205
灌木盖度
Shrub coverage
-0.350 -0.171 0.05 -0.169
草本丰富度
Herb richness
0.783** 0.431* 0.634**
草本多样性
Herb diversity
0.275 0.430*
草本密度
Herb density
0.861**

Table 6

Summary of redundancy analysis (RDA) ordination of understory species diversity and environmental factors"

排序概要
Summary of ordination
第1轴
Axis 1
第2轴
Axis 2
第3轴
Axis 3
第4轴
Axis 4
总方差
Total variance
特征值 Eigenvalues 0.611 0.254 0.035 0.017 1
林下植被物种多样性-环境因子相关
Understory species diversity-environmental factors correlations
0.980 0.969 0.886 0.892
解释的林下植被物种多样性组成变异的累积百分比
Cumulative percentage variance of understory species diversity data (%)
61.1 86.5 90.0 91.7
林下植被物种多样性-环境因子关系方差的累积百分比
Cumulative percentage variance of understory species diversity-environmental factors relation (%)
65.3 92.5 96.2 98.1
特征值总和 Sum of eigenvalues 1
典型特征值总和 Sum of canonical eigenvalues 0.935
第一典范轴的显著性测验 Significant test of the first canonical axis F = 10.975, P = 0.002
所有典范轴的显著性测验 Significant test of all canonical axes F = 6.248, P= 0.002

Table 7

Contribution of significant environmental factors affecting understory species diversity in redundancy analysis (RDA). * P < 0.05, ** P < 0.01."

环境因子 Environmental factors 解释率
Contribution (%)
F
土壤pH Soil pH 32.3 9.4**
土壤含水量 Soil water content (SWC) 16.1 6.0**
土壤全钾 Soil total potassium (TK) 9.7 4.7**
土壤全磷 Soil total phosphorus (TP) 8.6 4.9**
坡度 Slope (SLO) 8.6 3.4*
海拔 Elevation (ELE) 3.6 2.2
土壤有机质 Soil organic matter (SOM) 3.6 2.2
林内散射光 Transmitted diffuse in forest (Trans Dif) 3.1 1.9
林冠开阔度 Canopy openness (CO) 2.5 1.5
坡向 Slope aspect (SLOA) 2.4 1.4
林内总光照 Transmitted total in forest (Trans Tot) 2.3 1.3
土壤全氮 Soil total nitrogen (TN) 2 1.2
林内直射光 Transmitted direct in forest (Trans Dir) 1.6 0.9
坡位 Slope position 1.5 0.8
叶面积指数 Leaf area index (LAI) 1 0.5
消光系数 Extinction coefficient (k) 0.6 0.3

Fig. 1

RDA ordination diagram of understory species diversity and environmental factors. The arabic numerals in the figure represent 24 sample plots. The red arrow lines represent environmental factors (the factors with significant effects are display only, P < 0.05). The blue arrows lines represent diversity factors of understory plants. pH, SWC, SLO, TK, TP, SW_shrub, D_shrub, C_shrub, S_herb, SW_herb, D_herb and C_herb are the same as in Table 4 and Table 7."

Table 8

Stepwise regression analysis of understory species diversity and environmental factors. SWC, SLO, TK and TP are the same as in Table 7."

林下植被物种多样性
Understory species diversity
逐步回归方程
Stepwise regression equation
调整后决定系数
Adjusted R square (R2adj)
P
灌木丰富度 Shrub richness (S_shrub) S_shrub = 57.560 - 1.184TK - 6.416SWC - 5.410pH 0.599 0.015
灌木多样性 Shrub diversity (SW_shrub) SW_shrub = 3.938 - 0.290TK - 0.173TP 0.661 0.004
灌木密度 Shrub density (D_shrub) D_shrub = - 2888.420 + 482.654pH 0.189 0.019
灌木盖度 Shrub coverage (C_shrub) C_shrub = 5.855 + 0.023D_shrub 0.524 < 0.001
草本丰富度 Herb richness (S_herb) S_herb = 4.752 + 1.156TK 0.239 0.009
草本多样性 Herb diversity (SW_herb) SW_herb = 0.849 + 0.148S_herb 0.595 < 0.001
草本密度 Herb density (D_herb) D_herb = - 3061.719 + 6073.551SWC + 579.980TK 0.663 0.005
草本盖度 Herb coverage (C_herb) C_herb = - 90.729 + 7.579TK + 12.862TP - 1.653SLO 0.725 0.007
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