植物生态学报 ›› 2013, Vol. 37 ›› Issue (5): 373-383.DOI: 10.3724/SP.J.1258.2013.00039
• 研究论文 • 下一篇
余敏1,周志勇1,康峰峰1,欧阳帅1,米湘成2,孙建新1,*()
发布日期:
2013-05-16
通讯作者:
孙建新
基金资助:
YU Min1,ZHOU Zhi-Yong1,KANG Feng-Feng1,OUYANG Shuai1,MI Xiang-Cheng2,SUN Jian-Xin1,*()
Published:
2013-05-16
Contact:
SUN Jian-Xin
摘要:
通过对山西灵空山小蛇沟集水区的林下草本层植物群落进行调查和多元分析——TWINSPAN分类、典范对应分析(CCA)与生境、生物因素变量分离, 探讨林分水平上草本层物种分布与环境因子之间的关系。结果如下: 1) TWINSPAN将26个调查样方划分为6种群落类型: 以辽东栎(Quercus wutaishanica)为主的辽东栎-油松(Pinus tabulaeformis)林型、辽东栎杂木林型、辽东栎林型、华北落叶松(Larix principis-rupprechtii)林型、油松林和阔叶油松林型、油松-辽东栎均匀混交林型, 体现了该地区地带性植被类型为暖温带森林的特点。2)群落类型的划分与CCA的结果相吻合, 主要反映了CCA排序第一、二轴的环境梯度, CCA排序轴第一轴突出反映了林分类型与土壤养分梯度, 第二排序轴与坡度、坡位显著相关。Monte Carlo检验结果表明, 林分类型、土壤养分和坡度是影响小蛇沟集水区内林下草本物种分异的最主要的环境因子。3)生境因子与生物因子解释了物种格局变化的42.9%, 其中生境因子占31.8%, 生物因子占7.9%, 生境因子与生物因子交互作用解释部分占3.2%。良好的环境解释反映了调查取样和环境因子选取的合理性。对于50%以上未能被解释的变异部分, 可能归咎于未被选取的因子如干扰或者随机过程。4)在海拔梯度较小的山区, 坡向等小地形因子能较好地指示局部生境的小气候条件, 对林下植物的分布有较好的解释力。
余敏,周志勇,康峰峰,欧阳帅,米湘成,孙建新. 山西灵空山小蛇沟林下草本层植物群落梯度分析及环境解释. 植物生态学报, 2013, 37(5): 373-383. DOI: 10.3724/SP.J.1258.2013.00039
YU Min,ZHOU Zhi-Yong,KANG Feng-Feng,OUYANG Shuai,MI Xiang-Cheng,SUN Jian-Xin. Gradient analysis and environmental interpretation of understory herb-layer communities in Xiaoshegou of Lingkong Mountain, Shanxi, China. Chinese Journal of Plant Ecology, 2013, 37(5): 373-383. DOI: 10.3724/SP.J.1258.2013.00039
图1 山西灵空山小蛇沟集水区26个样方双向指示种分析的分类结果。最左侧竖线上的数字表示分类水平。N表示样方总数。图下方长方形内的数字为所调查的26个样方的编号。横线上的物种表示各分类水平的正负指示种(每一个二歧划分左边为负指示种, 右边为正指示种)。adgm, 狭叶沙参; adre, 薄叶荠苨; aram, 东北南星; atla, 苍术; capu, 紫斑风铃草; dini, 穿山龙; epbr, 淫羊霍; irru, 矮紫苞鸢尾; mero, 山罗花; posi, 黄精; ruco, 茜草; thmi, 东亚唐松草; thpe, 瓣蕊唐松草。
Fig. 1 Two-way indicator species analysis (TWINSPAN) classification results of 26 plots in Xiaoshegou catchement of Linkong Mountain, Shanxi. Numbers on the vertical line on the left designates classification level. N represents the total number of plots. Numbers in the rectangles at the bottom of the figure designate the identification code for each of the 26 plots. Species above horizontal lines are indicator species; there are a negative group and a positive group at each level of classification. adgm, Adenophora gmelinii; adre, Adenophora remotiflora; aram, Arisaema amurense; atla, Atractylodes lancea; capu, Campanula punctate; dini, Dioscorea nipponica; epbr, Epimedium brevicornu; irru, Iris ruthenica var. nana; mero, Melampyrum roseum; posi, Polygonatum sibiricum; ruco, Rubia cordifolia; thmi, Thalictrum minus var. hypoleucum; thpe, Thalictrum petaloideum.
变量 Variable | 与典范对应分析的相关性 Correlations with CCA ordination axes | 蒙特卡罗置换检验 Monte Carlo permutation test | ||||
---|---|---|---|---|---|---|
第1轴 Axis 1 | 第2轴 Axis 2 | 第3轴 Axis 3 | F值 F-value | p值 p-value | ||
生境因子 Habitat factors | ||||||
土壤综合变量 Composite soil variable | 0.882 5*** | 0.323 3 | -0.297 4 | 1.95 | 0.014* | |
坡位 Slope position | 0.208 3 | -0.664 1*** | -0.363 4 | 1.66 | 0.028* | |
坡度 Slope | -0.288 3 | 0.700 2*** | -0.489 8* | 1.59 | 0.058 | |
坡向 Slope aspect | 0.356 3 | 0.126 4 | -0.610 6*** | 1.21 | 0.236 | |
生物因子 Biotic factors | ||||||
林分类型 Stand type | -0.629 0*** | -0.058 5 | 0.382 0 | 1.76 | 0.006* | |
林冠孔隙度 Canopy gap fraction | -0.259 7 | 0.460 3* | 0.218 2 | 1.01 | 0.426 | |
特征值 Eigenvalue | 0.212 | 0.136 | 0.095 | |||
对物种数据解释百分比 Percentage explainable of species data | 11.3 | 18.6 | 23.7 | |||
物种环境关系的累积贡献率 Cumulative proportion of species-environment relationships | 33.7 | 55.3 | 70.4 |
表1 草本层典范对应分析(CCA)排序及蒙特卡罗检验结果
Table 1 Results by canonical correspondence analysis (CCA) ordination with the first three axes and Monte Carlo permutation test in herb-layer
变量 Variable | 与典范对应分析的相关性 Correlations with CCA ordination axes | 蒙特卡罗置换检验 Monte Carlo permutation test | ||||
---|---|---|---|---|---|---|
第1轴 Axis 1 | 第2轴 Axis 2 | 第3轴 Axis 3 | F值 F-value | p值 p-value | ||
生境因子 Habitat factors | ||||||
土壤综合变量 Composite soil variable | 0.882 5*** | 0.323 3 | -0.297 4 | 1.95 | 0.014* | |
坡位 Slope position | 0.208 3 | -0.664 1*** | -0.363 4 | 1.66 | 0.028* | |
坡度 Slope | -0.288 3 | 0.700 2*** | -0.489 8* | 1.59 | 0.058 | |
坡向 Slope aspect | 0.356 3 | 0.126 4 | -0.610 6*** | 1.21 | 0.236 | |
生物因子 Biotic factors | ||||||
林分类型 Stand type | -0.629 0*** | -0.058 5 | 0.382 0 | 1.76 | 0.006* | |
林冠孔隙度 Canopy gap fraction | -0.259 7 | 0.460 3* | 0.218 2 | 1.01 | 0.426 | |
特征值 Eigenvalue | 0.212 | 0.136 | 0.095 | |||
对物种数据解释百分比 Percentage explainable of species data | 11.3 | 18.6 | 23.7 | |||
物种环境关系的累积贡献率 Cumulative proportion of species-environment relationships | 33.7 | 55.3 | 70.4 |
图2 26个样地与环境因子的典范对应分析(CCA)排序图。土壤综合变量(SOIL)是利用主成分分析对土壤有机碳、全氮、全磷提取的一个集合变量。阿拉伯字母表示样方编号, 环境变量用带箭头的线段表示, 箭头所指的方向是环境因子变化最大的方向, 箭头连线的长度表示该环境因子与物种分布相关程度的大小, 连线越长, 相关性越大, 能解释物种分布变化的比例也越大。两个箭头之间的夹角表示环境变量间的相关程度, 夹角越小, 表明相关性越大, 夹角呈锐角时环境变量之间正相关, 呈钝角时负相关, 呈直角时表示相关程度很小。箭头连线与排序轴的夹角表示该环境因子与排序轴的相关性大小, 夹角越小, 相关性越高。样方间的距离表示样方的相似程度, 距离越近, 代表样方的物种组成和对环境因子的要求越相似。
Fig. 2 Canonical correspondence analysis (CCA) ordination diagram of 26 plots and environmental factors. SOIL is a combined variable exacted from soil organic carbon, total nitrogen, total phosphorus by principle component analysis. The numerical numbers are the identification code for each of plots, and arrow lines represent environmental variables. Arrows indicate the direction of steepest increase of the corresponded environmental variable. The lengths of the line indicate the correlation between environmental variables and species distribution; longer the arrow line, closer the relationship, and greater the proportion it explains the species distribution of the herb-layer. The correlation of environmental factors was explained by the angle between the two corresponding arrows; smaller angle means tight relationship. An acute angle means positive relationship; whereas an obtuse angle means negative relationship. A right angle indicates a lack of close relationship. The angle between arrow line and ordination axis reflects the relationship between environmental variable and the axis; smaller the angle closer the relationship. The distance between plots indicates the similarity in species composition between the two plots; shorter the distance, more similar the species composition between them.
图3 影响草本物种分布的生境、生物因子的变异分离。a, 生境因子单独解释的部分; b, 生物因子单独解释的变异; c, 生境因子和生物因子交互作用解释的变异; d, 不能由生境和生物因子解释的变异, 用残差表示。
Fig. 3 Variance partitioning of habitat and biotic factors in explaining herb species distribution. a, part explained by habitat factors alone; b, variation explainable by biotic factors alone; c, variation explainable by an interactive effect of habitat and biotic factors; d, variations that are not explainable by habitat and biotic factors and shown as residuals.
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