Effects of topographic heterogeneity on species diversity in a monsoon evergreen broad- leaved forest in Puʼer, Yunnan, China

doi:10.17521/cjpe.2020.0148

Abstract

Abstract:

Aims In subtropical forests, topographic heterogeneity drives ecosystem structure and species composition by regulating the available nutrients, water and light needed for tree growth. A gap in current research is the role of topographic heterogeneity in plant species diversity and distribution in monsoon evergreen broad-leaved forests. Our study aims to fill this gap, providing scientific grounds for the conservation of biodiversity through an exploration of floral structure and ecosystem development in monsoon evergreen broad-leaved forests.
Methods Within a 30 hm² dynamics plot, we established 750 20 m × 20 m sample plots, using fuzzy C-mean clustering to classify different topographic types based on elevations, convexities, slopes, and aspects of each sample plots. We compared community composition and community species diversity between different topographic types, using Torus-translation tests to examine the relationship between floral species and topography.
Important findings We identified five distinct topography types: ridge (8.00 hm²), steep slope (6.04 hm²), less-steep slope (7.68 hm²), high plateau (2.76 hm²), and valley (5.52 hm²). We counted a total of 153 418 woody plant individuals with a diameter at breast height (DBH) ≥ 1 cm, belonging to 271 species, 179 genera and 79 families. Among the topographic types, we noted differences in species richness, plant density, proportion of different diameters, species diversity and the abundance of dominant species. When comparing species-area relationships between the five topographic types, we noticed that the ridge had the lowest number of species, while the valley had the highest. The cumulative species-individual relationships within the topographic types revealed that the ridge possesses the lowest cumulative rate. Of the 123 examined species, 67.5% were significantly related to at least one type of topography. Among species significantly related to topography, those species with significant negative correlation were more abundant in the ridge and steep slope topographic types. The opposite was true in steep slope, high plateau and valley. Based on our data, we determined that topographic heterogeneity contributed 7.8% to the maintenance of species diversity in our study area.

Key words: topographic heterogeneity, monsoon broad-leaved evergreen forest, species composition, community structure, importance value, species diversity, topographic association of woody plants

WANG Yan-Hong, LI Shuai-Feng, LANG Xue-Dong, HUANG Xiao-Bo, LIU Wan-De, XU Chong-Hua, SU Jian-Rong. Effects of topographic heterogeneity on species diversity in a monsoon evergreen broad- leaved forest in Puʼer, Yunnan, China[J]. Chin J Plant Ecol, 2020, 44(10): 1015-1027.

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Figures/Tables 12

References 55

[1]	Anderson MJ, Crist TO, Chase JM, Vellend M, Inouye BD, Freestone AL, Sanders NJ, Cornell HV, Comita LS, Davies KF, Harrison SP, Kraft NJB, Stegen JC, Swenson NG (2011). Navigating the multiple meanings of β diversity: a roadmap for the practicing ecologist. Ecology Letters, 14, 19-28.
[2]	Bell G, Lechowicz MJ, Waterway MJ (2000). Environmental heterogeneity and species diversity of forest sedges. Journal of Ecology, 88, 67-87.
[3]	Bellingham PJ, Tanner EVJ (2000). The influence of topography on tree growth, mortality, and recruitment in a tropical montane forest. Biotropica, 32, 378-384.
[4]	Bezdek JC (1981). Pattern Recognition with Fuzzy Objective Function Algorithms. Plenum Press, New York.
[5]	Bu WS (2013). The Relationships Between Biodiversity and Ecosystem Functioning in Natural Tropical Forests of Hainan Island, China. PhD dissertation, Chinese Academy of Forestry, Beijing. 89-95.
	[ 卜文圣 (2013). 海南岛热带天然林生物多样性与生态系统功能关系的研究. 博士学位论文, 中国林业科学研究院, 北京. 89-95.]
[6]	Chase JM (2003). Community assembly: When should history matter? Oecologia, 136, 489-498.
[7]	Chen L, Mi XC, Comita LS, Zhang LW, Ren HB, Ma KP (2010). Community-level consequences of density dependence and habitat association in a subtropical broad-leaved forest. Ecology Letters, 13, 695-704.
[8]	Chen M, Fang H, Cao M (2008). Sprouting characteristics of sprouted woody plants in the mid-mountain humid evergreen broad-leaved forest on Ailao Mountain, Yunnan Province. Guihaia, 28, 627-632.
	[ 陈沐, 房辉, 曹敏 (2008). 云南哀牢山中山湿性常绿阔叶林树种萌生特征研究. 广西植物, 28, 627-632.]
[9]	Chuyong GB, Kenfack D, Harms KE, Thomas DW, Condit R, Comita LS (2011). Habitat specificity and diversity of tree species in an African wet tropical forest. Plant Ecology, 212, 1363-1374.
[10]	Clark DB, Clark DA, Read JM (1998). Edaphic variation and the mesoscale distribution of tree species in a neotropical rain forest. Journal of Ecology, 86, 101-112.
[11]	Condit R (1998). Tropical Forest Census Plots: Methods and Results from Barro Colorado Island, Panama and a Comparison with Other Plots. Springer-Verlag, Berlin.
[12]	Forrester D, Bauhus J, Connell M (2003). Competition in thinned Silvertop ash ( Eucalyptus sieberi L. Johnson) stands from early coppice growth. Forest Ecology and Management, 174, 459-475.
[13]	Gong GQ, Chen JM, Mi XC, Chen SW, Fang T (2007). Habitat associations of wood species in the Gutianshan subtropical broad-leaved evergreen forest. Science of Soil and Water Conservation, 5(3), 79-83.
	[ 宫贵权, 程积民, 米湘成, 陈声文, 方腾 (2007). 古田山常绿阔叶林木本植物与生境的相关性. 中国水土保持科学, 5(3), 79-83.]
[14]	Guo YL, Wang B, Xiang WS, Ding T, Lu SH, Huang FZ, Wen SJ, Li DX, He YL, Li XK (2016). Responses of spatial pattern of woody plants’ basal area to topographic factors in a tropical karst seasonal rainforest in Nonggang, Guangxi, southern China. Biodiversity Science, 24, 30-39.
	[ 郭屹立, 王斌, 向悟生, 丁涛, 陆树华, 黄甫昭, 文淑均, 李冬兴, 何运林, 李先琨 (2016). 喀斯特季节性雨林木本植物胸高断面积分布格局及其对地形因子的响应. 生物多样性, 24, 30-39.]
[15]	Guo YL, Wang B, Xiang WS, Ding T, Lu SH, Wen SJ, Huang FZ, Li DX, Li XK (2015). Sprouting characteristics of tree species in 15 hm² plot of northern tropical karst seasonal rain forest in Nonggang, Guangxi, southern China. Chinese Journal of Ecology, 34, 955-961.
	[ 郭屹立, 王斌, 向悟生, 丁涛, 陆树华, 文淑均, 黄甫昭, 李冬兴, 李先琨 (2015). 弄岗北热带喀斯特季节性雨林15 hm²样地木本植物萌生特征 . 生态学杂志, 34, 955-961.]
[16]	Hao MH, Zhang ZH, Zhao SS, Zhao XH, Beiketuerhan Y, Zhang CY (2017). Habitat associations of tree growth in a coniferous and broad-leaved mixed forest in Jiaohe, Jilin Province. Acta Ecologica Sinica, 37, 3437-3444.
	[ 郝珉辉, 张忠辉, 赵珊珊, 赵秀海, 叶尔江·拜克吐尔汉, 张春雨 (2017). 吉林蛟河针阔混交林树木生长与生境的关联性. 生态学报, 37, 3437-3444.]
[17]	Harms KE, Condit R, Hubbell SP, Foster RB (2001). Habitat associations of trees and shrubs in a 50 hm² neotropical forest plot . Journal of Ecology, 89, 947-959.
[18]	Hennenberg KJ, Bruelheide H (2003). Ecological investigations on the northern distribution range of Hippocrepis comosa L. in Germany. Plant Ecology, 166, 167-188.
[19]	Hofer G, Wagner HH, Herzog F, Edwards PJ (2008). Effects of topographic variability on the scaling of plant species richness in gradient dominated landscapes. Ecography, 31, 131-139.
[20]	Hubbell S, Foster RB (1986). Commonness and rarity in a neotropical forest: implications for tropical tree conservation//Soule ME. Conservation Biology: Science of Scarcity and Diversity. Sinauer Press, Sunderland, UK.
[21]	Irl SDH, Harter DEV, Steinbauer MJ, Puyol GD, Fernández- Palacios JM, Jentsch A, Beierkuhnlein C (2015). Climate vs. topography-spatial patterns of plant species diversity and endemism on a high-elevation island. Journal of Ecology, 103, 1621-1633.
[22]	Jiang P, Ye J, Wu G (2005). Woody species composition and biomass of main tree species in a 25 hm² plot of broad-leaved and Korean pine mixed forests of Changbai Mountain, northeast China . Journal of Beijing Forestry University, 27(Supp.2), 112-115.
	[ 姜萍, 叶吉, 吴钢 (2005). 长白山阔叶红松林大样地木本植物组成及主要树种的生物量. 北京林业大学学报, 27(增刊2), 112-115.]
[23]	Lai JS, Mi XC, Ren HB, Ma KP (2010). Numerical classification of associations in subtropical evergreen broad- leaved forest based on multivariate regression trees—A case study 24 hm² Gutianshan forest plot in China . Chinese Journal of Plant Ecology, 34, 761-769.
	[ 赖江山, 米湘成, 任海保, 马克平 (2010). 基于多元回归树的常绿阔叶林群丛数量分类——以古田山24公顷森林样地为例. 植物生态学报, 34, 761-769.]
[24]	Lan GY, Hu YH, Cao M, Zhu H (2011). Topography related spatial distribution of dominant tree species in a tropical seasonal rain forest in China. Forest Ecology and Management, 262, 1507-1513.
[25]	Lan GY, Hu YH, Cao M, Zhu H, Wang H, Zhou SS, Deng XB, Cui JY, Huang JG, Liu LY, Xu HL, Song JP, He YC (2008). Establishment of Xishuangbanna tropical forest dynamics plot: species compositions and spatial distribution patterns. Journal of Plant Ecology (Chinese Version), 32, 287-298.
	[ 兰国玉, 胡跃华, 曹敏, 朱华, 王洪, 周仕顺, 邓晓保, 崔景云, 黄建国, 刘林云, 许海龙, 宋军平, 何有才 (2008). 西双版纳热带森林动态监测样地: 树种组成与空间分布格局. 植物生态学报, 32, 287-298.]
[26]	Li SF, Lang XD, Huang XB, Wang YH, Liu WD, Xu CH, Su JR (2020). Association classification of a 30 hm² dynamics plot in the monsoon broad-leaved evergreen forest in Pu’er, Yunnan Province, China. Chinese Journal of Plant Ecology, 44, 248-259.
	[ 李帅锋, 郎学东, 黄小波, 王艳红, 刘万德, 徐崇华, 苏建荣 (2020). 云南普洱30公顷季风常绿阔叶林动态监测样地群丛数量分类. 植物生态学报, 44, 248-259.]
[27]	Li SF, Liu WD, Su JR, Zhang ZJ, Liu QY (2012). Woody seedling regeneration in secondary succession of monsoon broad-leaved evergreen forest in Pu’er, Yunnan, southwest China. Acta Ecologia Sinica, 32, 5653-5662.
	[ 李帅锋, 刘万德, 苏建荣, 张志钧, 刘庆云 (2012). 普洱季风常绿阔叶林次生演替中木本植物幼苗更新特征. 生态学报, 32, 5653-5662.]
[28]	Li X, Nie Y, Song X, Zhang R, Wang G (2011). Patterns of species diversity and functional diversity along the south to north-facing slope gradient in a sub-alpine meadow. Community Ecology, 12, 179-187.
[29]	Liu JJ, Tan YH, Slik JWF (2014). Topography related habitat associations of tree species traits, composition and diversity in a Chinese tropical forest. Forest Ecology and Management, 330, 75-81.
[30]	Liu WP, Cao HL, Liu W, Lian JY, Wu LF (2011). Study on diversity of monsoon evergreen broad leaved forest in diferent kinds of habitat in Dinghushan. Journal of Anhui Agricultural Science, 39, 16159-16163.
	[ 刘文平, 曹洪麟, 刘卫, 练琚愉, 吴林芳 (2011). 鼎湖山季风常绿阔叶林不同生境物种多样性研究. 安徽农业科学, 39, 16159-16163.]
[31]	Lundholm JT (2009). Plant species diversity and environmental heterogeneity: spatial scale and competing hypotheses. Journal of Vegetation Science, 20, 377-391.
[32]	Ma KP (2017). Forest dynamics plot is a crosscutting research platform for biodiversity science. Biodiversity Science, 25, 227-228.
	[ 马克平 (2017). 森林动态大样地是生物多样性科学综合研究平台. 生物多样性, 25, 227-228.]
[33]	Pigott CD (1975). Experimental studies on the influence of climate on the geographical distribution of plants. Weather, 30, 82-90.
[34]	Punchi-Manage R, Getzin S, Wiegand T, Kanagaraj R, Savitri GCV, Nimal GIAU, Wiegand K, Huth A (2012). Effects of topography on structuring local species assemblages in a Sri Lankan mixed dipterocarp forest. Journal of Ecology, 101, 149-160.
[35]	Ru HL, Zhang HD, Jiao F, Xue CY, Guo ML (2015). Impact of micro-landform on grassland plant community structure and function in the hilly Loess Plateau region, China. Chinese Journal of Applied Ecology, 27, 25-32.
	[ 汝海丽, 张海东, 焦峰, 薛超玉, 郭美丽 (2015). 黄土丘陵区微地形对草地植物群落结构组成和功能特征的影响. 应用生态学报, 27, 25-32.]
[36]	Sharma CM, Suyal S, Gairola S, Ghildiyal SK (2009). Species richness and diversity along an altitudinal gradient in moist temperate forest of Garhwal Himalaya. Journal of American Science, 5, 119-128.
[37]	Shen GC, Yu MJ, Hu XS, Mi XC, Ren HB, Sun IF, Ma KP (2009). Species-area relationships explained by the joint effects of dispersal limitation and habitat heterogeneity. Ecology, 90, 3033-3041.
[38]	Shen ZH, Zhang XS, Jin YX (2000). Spatial pattern analysis and topographical interpretation of species diversity in the forests of Dalaoling in the region of the Three Gorges. Acta Botanica Sinica, 42, 620-627.
	[ 沈泽昊, 张新时, 金义兴 (2000). 三峡大老岭森林物种多样性的空间格局分析及其地形解释. 植物学报, 42, 620-627.]
[39]	Song YC, Yan ER, Song K (2015). Synthetic comparison of eight dynamics plots in evergreen broadleaf forests, China. Biodiversity Science, 23, 139-148.
	[ 宋永昌, 阎恩荣, 宋坤 (2015). 中国常绿阔叶林8大动态监测样地植被的综合比较. 生物多样性, 23, 139-148.]
[40]	Su JR, Liu WD, Li SF, Lang XD (2015). Restoration Ecology of Monsoon Broad-leaved Evergreen Forest in West China. Science Press, Beijing. 3-4.
	[ 苏建荣, 刘万德, 李帅锋, 郎学东 (2015). 西部季风常绿阔叶林恢复生态学. 科学出版社, 北京. 3-4.]
[41]	Su JR, Liu WD, Zhang ZJ, Li SF (2012). Sprouting characteristic in restoration ecosystems of monsoon evergreen broadleaved forest in south-central of Yunnan Province. Acta Ecologica Sinica, 32, 805-814.
	[ 苏建荣, 刘万德, 张志钧, 李帅锋 (2012). 云南中南部季风常绿阔叶林恢复生态系统萌生特征. 生态学报, 32, 805-814.]
[42]	Su SJ (2015). Spatial Heterogeneity and Vegetation Maintaining Mechanism of Pinus taiwanensis Forest in Daiyun Mountain High Altitude Area. PhD dissertation, Fujian Agriculture and Forestry University, Fuzhou, 49-65.
	[ 苏松锦 (2015). 戴云山高海拔黄山松林的空间异质性与植被维持机制. 博士学位论文, 福建农林大学, 福州. 49-65.]
[43]	Sun XW, Yang QS, Liu HM, Wang XH (2018). Classification of plant associations based on a 20 hm² dynamics plot of evergreen broad-leaved forest in Mt. Tiantong, Zhejiang, China. Chinese Journal of Plant Ecology, 42, 550-561.
	[ 孙小伟, 杨庆松, 刘何铭, 王希华 (2018). 基于浙江天童20 hm²常绿阔叶林动态监测样地的群丛划分 . 植物生态学报, 42, 550-561.]
[44]	The Editorial Committee of Flora Reipublicae Popularis Sinicae, Chinese Academy of Sciences (2019). Flora Reipublicae Popularis Sinicae. http://www.iplant.cn/frps.
	[ 中国科学院中国植物志编辑委员会 (2019). 中国植物志. http://www.iplant.cn/frps.]
[45]	Tilman D, Pacala S (1993). The Maintenance of Species Richness in Plant Communities. The University of Chicago Press, Chicago.
[46]	Wang JM, Xu H, Li YD, Lin MX, Zhou Z, Luo TS, Chen DX (2018). Effects of topographic heterogeneity on community structure and diversity of woody plants in Jianfengling tropical montane rainforest. Sciental Silvae Sinicae, 54(1), 1-11.
	[ 王家鸣, 许涵, 李意德, 林明献, 周璋, 骆土寿, 陈德祥 (2018). 地形异质性对尖峰岭热带山地雨林木本植物群落结构及多样性的影响. 林业科学, 54(1), 1-11.]
[47]	Wang W, Luo ZR, Zhou RF, Xu DM, Ai JG, Ding BY (2011). Habitat associations of woody plant species in Baishanzu subtropical broad-leaved evergreen forest. Biodiversity Science, 19, 134-142.
	[ 王伟, 骆争荣, 周荣飞, 许大明, 哀建国, 丁炳扬 (2011). 百山祖常绿阔叶林木本植物的生境相关性分析. 生物多样性, 19, 134-142.]
[48]	Wen HD, Lin LX, Yang J, Hu YH, Cao M, Liu YH, Lu ZY, Xie YN (2018). Species composition and community structure of a 20 hm² plot of mid-mountain moist evergreen broad-leaved forest on the Mts. Ailaoshan, Yunnan Province, China . Chinese Journal of Plant Ecology, 42, 419-429.
	[ 温韩东, 林露湘, 杨洁, 胡跃华, 曹敏, 刘玉洪, 鲁志云, 谢有能 (2018). 云南哀牢山中山湿性常绿阔叶林 20 hm² 动态样地的物种组成与群落结构 . 植物生态学报, 42, 419-429.]
[49]	Yang QS, Ma ZP, Xie YB, Zhang ZG, Wang ZH, Liu HM, Li P, Zhang N, Wang DL, Yang HB, Fang XF, Yan ER, Wang XH (2011). Community structure and species composition of an evergreen broadleaved forest in Tiantong’s 20 hm² dynamic plot, Zhejiang Province, eastern China . Biodiversity Science, 19, 215-223.
	[ 杨庆松, 马遵平, 谢玉彬, 张志国, 王樟华, 刘何铭, 李萍, 张娜, 王达力, 杨海波, 方晓峰, 阎恩荣, 王希华 (2011). 浙江天童20 hm²常绿阔叶林动态监测样地的群落特征 . 生物多样性, 19, 215-223.]
[50]	Yang YC, Da LJ, You WH (2005). Vegetation structure in relation to micro-landform in Tiantong National Forest Park, Zhejiang, China. Acta Ecologica Sinica, 25, 38-48.
	[ 杨永川, 达良俊, 由文辉 (2005). 浙江天童国家森林公园微地形与植被结构的关系. 生态学报, 25, 38-48.]
[51]	Ye WH, Cao HL, Huang ZL, Lian JY, Wang ZG, Li L, Wei SG, Wang ZM (2008). Community structure of a 20 hm² lower subtropical evergreen broadleaved forest plot in Dinghushan, China. Journal of Plant Ecology (Chinese Version), 32, 274-286.
	[ 叶万辉, 曹洪麟, 黄忠良, 练琚愉, 王志高, 李林, 魏识广, 王章明 (2008). 鼎湖山南亚热带常绿阔叶林20公顷样地群落特征研究. 植物生态学报, 32, 274-286.]
[52]	Yuan TX, Zhang HP, Ou ZY, Tan YB (2014). Effects of topography on the diversity and distribution pattern of ground plants in karst montane forests in southwest Guangxi, China. Chinese Journal of Applied Ecology, 25, 2803-2810.
	[ 袁铁象, 张合平, 欧芷阳, 谭一波 (2014). 地形对桂西南喀斯特山地森林地表植物多样性及分布格局的影响. 应用生态学报, 25, 2803-2810.]
[53]	Zhang YD, Liu SR, Zhao CM (2005). Spatial pattern of sub- alpine forest restoration in west Sichuan. Chinese Journal of Applied Ecology, 16, 1706-1710.
	[ 张远东, 刘世荣, 赵常明 (2005). 川西亚高山森林恢复的空间格局分析. 应用生态学报, 16, 1706-1710.]
[54]	Zhu H (2019). Floristic divergence of the evergreen broad-leaved forests in Yunnan, southwestern China. Phytotaxa, 393, 1-20.
[55]	Zhu Y, Zhao GF, Zhang LW, Shen GC, Mi XC, Ren HB, Yu MJ, Chen JH, Chen SW, Fang T, Ma KP (2008). Community composition and structure of Gutianshan forest dynamic plot in a mid-subtropical evergreen broad-leaved forest, east China. Journal of Plant Ecology (Chinese Version), 32, 262-273.
	[ 祝燕, 赵谷风, 张俪文, 沈国春, 米湘成, 任海保, 于明坚, 陈建华, 陈声文, 方腾, 马克平 (2008). 古田山中亚热带常绿阔叶林动态监测样地——群落组成与结构. 植物生态学报, 32, 262-273.]

地形 Topography	海拔 Elevation (m)	坡度 Slope (°)	凹凸度 Convexity (m)	坡向 Slope aspect (°)	样地数 Sample plot number
山脊 Ridge	1 547.0 ± 0.96^a	19.6 ± 0.33^a	0.67 ± 0.14^a	164.41 ± 1.56^a	200
陡坡 Steep slope	1 525.7 ± 1.62^b	19.4 ± 0.50^a	-0.17 ± 0.18^b	324.74 ± 2.19^b	151
缓坡 Less-steep slope	1 504.5 ± 1.18^c	18.4 ± 0.34^b	-0.42 ± 0.12^b	185.70 ± 1.27^c	192
高谷 High plateau	1 548.7 ± 3.10^a	19.5 ± 0.56^a	0.41 ± 0.21^a	52.53 ± 4.52^d	69
沟谷 Valley	1 498.3 ± 1.89^d	18.8 ± 0.52^a	-0.91 ± 0.20^bc	294.68 ± 2.23^e	138

地形 Topography	海拔 Elevation (m)	坡度 Slope (°)	凹凸度 Convexity (m)	坡向 Slope aspect (°)	样地数 Sample plot number
山脊 Ridge	1 547.0 ± 0.96^a	19.6 ± 0.33^a	0.67 ± 0.14^a	164.41 ± 1.56^a	200
陡坡 Steep slope	1 525.7 ± 1.62^b	19.4 ± 0.50^a	-0.17 ± 0.18^b	324.74 ± 2.19^b	151
缓坡 Less-steep slope	1 504.5 ± 1.18^c	18.4 ± 0.34^b	-0.42 ± 0.12^b	185.70 ± 1.27^c	192
高谷 High plateau	1 548.7 ± 3.10^a	19.5 ± 0.56^a	0.41 ± 0.21^a	52.53 ± 4.52^d	69
沟谷 Valley	1 498.3 ± 1.89^d	18.8 ± 0.52^a	-0.91 ± 0.20^bc	294.68 ± 2.23^e	138

参数 Parameter	面积 Area (hm²)	个体数 Number of individuals	植株密度 Plant density (individual·hm^-2)	根萌生 Root sprouting	干基萌生 Stem basal sprouting	干萌生 Stem epicormic	枝萌生 Branch epicormic
大样地 Big sample plot	30.00	153 418	5 114	14 691	675	721	-
山脊 Ridge	8.00	44 861	5 608	5 222	98	142	-
陡坡 Steep slope	6.04	29 703	4 918	2 687	118	113	1
缓坡 Less-steep slope	7.68	39 320	5 120	3 295	203	207	2
高谷 High plateau	2.76	13 843	5 016	1 682	62	80	-
沟谷 Valley	5.52	25 691	4 654	1 805	195	179	38

参数 Parameter	面积 Area (hm²)	个体数 Number of individuals	植株密度 Plant density (individual·hm^-2)	根萌生 Root sprouting	干基萌生 Stem basal sprouting	干萌生 Stem epicormic	枝萌生 Branch epicormic
大样地 Big sample plot	30.00	153 418	5 114	14 691	675	721	-
山脊 Ridge	8.00	44 861	5 608	5 222	98	142	-
陡坡 Steep slope	6.04	29 703	4 918	2 687	118	113	1
缓坡 Less-steep slope	7.68	39 320	5 120	3 295	203	207	2
高谷 High plateau	2.76	13 843	5 016	1 682	62	80	-
沟谷 Valley	5.52	25 691	4 654	1 805	195	179	38

地形 Topography	物种 Species	相对频度 Relative frequency	相对多度 Relative abundance	相对胸高断面积 Relative basal area at the breast height	重要值 Importance value
山脊 Ridge	短刺锥 Castanopsis echidnocarpa	3.7	28.5	34.0	22.0
	泥柯 Lithocarpus fenestratus	3.7	14.5	6.5	8.2
	截果柯 Lithocarpus truncatus	3.7	5.3	7.1	5.3
	西南木荷 Schima wallichii	3.6	2.6	9.7	5.3
	茶梨 Anneslea fragrans	3.5	4.4	5.4	4.4
	隐距越桔 Vaccinium exaristatum	3.5	5.4	3.6	4.2
	红梗润楠 Machilus rufipes	3.6	4.6	2.5	3.6
	红叶木姜子 Litsea rubescens	3.6	4.3	2.5	3.5
	红皮水锦树 Wendlandia tinctoria subsp. intermedia	3.3	3.0	2.8	3.0
	深绿山龙眼 Helicia nilagirica	3.5	3.3	2.1	3.0
陡坡 Steep slope	短刺锥 Castanopsis echidnocarpa	2.8	16.4	22.8	14.0
	泥柯 Lithocarpus fenestratus	2.6	9.8	4.2	5.5
	西南木荷 Schima wallichii	2.8	3.4	9.8	5.3
	红梗润楠 Machilus rufipes	2.8	5.5	4.4	4.2
	枹丝锥 Castanopsis calathiformis	1.9	5.6	5.0	4.2
	截果柯 Lithocarpus truncatus	2.6	4.1	5.4	4.0
	红叶木姜子 Litsea rubescens	2.7	4.5	2.6	3.2
	西桦 Betula alnoides	0.9	0.5	7.4	3.0
	茶梨 Anneslea fragrans	2.5	3.4	2.9	2.9
	隐距越桔 Vaccinium exaristatum	2.2	2.8	1.8	2.2
缓坡 Less-steep slope	短刺锥 Castanopsis echidnocarpa	2.8	21.8	24.7	16.5
	西南木荷 Schima wallichii	2.8	3.2	14.3	6.8
	泥柯 Lithocarpus fenestratus	2.8	10.1	6.3	6.4
	枹丝锥 Castanopsis calathiformis	2.5	7.2	4.4	4.7
	红梗润楠 Machilus rufipes	2.9	6.0	4.8	4.6
	截果柯 Lithocarpus truncatus	2.7	4.2	6.3	4.4
	茶梨 Anneslea fragrans	2.6	4.0	3.7	3.5
	红叶木姜子 Litsea rubescens	2.7	3.7	3.8	3.4
	隐距越桔 Vaccinium exaristatum	2.0	2.4	1.7	2.0
	艾胶算盘子 Glochidion lanceolarium	2.7	2.4	0.6	1.9
高谷 High plateau	短刺锥 Castanopsis echidnocarpa	3.1	16.7	24.0	14.6
	泥柯 Lithocarpus fenestratus	3.0	17.2	5.7	8.6
	西南木荷 Schima wallichii	3.2	3.2	13.5	6.7
	红梗润楠 Machilus rufipes	3.2	5.7	5.0	4.7
	截果柯 Lithocarpus truncatus	2.8	3.9	5.7	4.1
	红叶木姜子 Litsea rubescens	3.1	5.4	2.9	3.8
	西桦 Betula alnoides	1.4	0.8	7.7	3.3
	隐距越桔 Vaccinium exaristatum	2.7	4.0	2.4	3.0
	茶梨 Anneslea fragrans	3.1	2.8	2.7	2.9
	枹丝锥 Castanopsis calathiformis	2.0	2.6	2.7	2.5
沟谷 Valley	短刺锥 Castanopsis echidnocarpa	2.5	12.9	15.6	10.4
	西南木荷 Schima wallichii	2.4	3.4	10.9	5.6
	泥柯 Lithocarpus fenestratus	2.3	9.5	4.2	5.4
	红梗润楠 Machilus rufipes	2.6	5.6	6.2	4.8
	截果柯 Lithocarpus truncatus	2.4	3.7	6.3	4.1
	枹丝锥 Castanopsis calathiformis	1.9	5.6	3.1	3.5
	云南瘿椒树 Tapiscia yunnanensis	1.8	1.4	4.7	2.6
	茶梨 Anneslea fragrans	2.2	3.2	2.2	2.6
	红叶木姜子 Litsea rubescens	2.2	3.2	2.2	2.5
	齿叶黄杞 Engelhardtia serrata	1.7	2.4	3.2	2.4