Biodiversity Science ›› 2017, Vol. 25 ›› Issue (1): 23-33.doi: 10.17520/biods.2016199

• Orginal Article • Previous Article     Next Article

Spatial heterogeneity and its causes in evergreen broad-leaved forests in the Ailao Mountains, Yunnan Province

Yuanjie Xu1, Dunmei Lin2, Ming Shi3, Yanjie Xie3, Yizhi Wang1, Zhenhua Guan1, *(), Jianying Xiang1   

  1. 1 Yunnan Academy of Biodiversity, Southwest Forestry University, Kunming 650224
    2 Key Laboratory of the Three Gorges Reservoir Region’s Eco-Environment, Ministry of Education, Chongqing University, Chongqing 400044
    3 The Faculty of Forestry, Southwest Forestry University, Kunming 650224
  • Received:2016-07-20 Accepted:2017-01-05 Online:2017-02-08
  • Guan Zhenhua

Understanding the variation of species richness over spatial scales, and elucidating the response of plant community composition to habitat heterogeneity has been one of the major topics in the study of community ecology. Niche differentiation related to habitat heterogeneity plays an important role in shaping diversity levels and species distribution patterns of plant communities. Based on a survey of 42 plots of four types of evergreen broad-leaved forests in the Ailao Mountains of Yunnan Province, this paper explored how community distribution and tree species richness of such forests changed along topographic and edaphic gradients. We found significant differences in community composition among the four types of evergreen broad-leaved forests. Tree species richness of monsoon evergreen broad-leaved forests was significantly higher than that of the other forest types. Tree species richness of mid-montane moist evergreen broad-leaved forests was significantly higher than that of semi-humid evergreen broad-leaved forests, whereas no significant difference was observed among the other forest types. Non-metric multidimensional scaling (NMDS) revealed that nine environmental gradients, including elevation, slope, soil water content, soil organic matter, were the determinants of forest community composition and tree distribution patterns. As the best predictors of tree species richness, soil available boron, elevation, soil total potassium and convexity together explained 63.2% of the variance of tree species richness. The combination of topographic and edaphic factors explained most of the variance of community composition and tree species richness of the evergreen broad leaved forests in the Ailao Mountains.

Key words: habitat heterogeneity, community composition, tree species richness, evergreen broad-leaved forests, the Ailao Mountains

Fig. 1

Location of the Ailao Mountains and overview of the study area"

Table 1

Relative dominance (%) of tree species and their contributions (%) to within-group similarities. The contributions of different tree species are shown in brackets."

Species name
倒卵叶石栎 Lithocarpus crassifolius 25.54(23.40)
云南桤叶树 Clethra delavayi 12.81(17.81)
露珠杜鹃 Rhododendron irroratum 19.47(17.51)
云南越桔 Vaccinium duclouxii 8.37(13.47) 8.68(21.18)
珍珠花 Lyonia ovalifolia 10.45(12.52) 9.99(10.38)
珊瑚冬青 Ilex corallina 5.94(6.35) 8.50(13.34)
木果石栎 Lithocarpus xylocarpus 16.88 (20.10)
腾冲栲 Castanopsis wattii 7.50(7.53)
硬斗柯 Lithocarpus hancei 8.96(6.42)
薄叶马银花 Rhododendron leptothrium 3.30(4.70)
舟柄茶 Stewartia pteropetiolata 3.94(3.51)
滇木荷 Schima noronhae 5.13(2.59)
四川冬青 Ilex szechwanensis 2.72(2.01)
旱冬瓜 Alnus nepalensis 19.39(21.73) 9.71(18.22)
高山栲 Castanopsis delavayi 9.56(21.37)
母猪果 Helicia nilagirica 4.29(10.23)
茶梨 Anneslea fragrans 2.51(8.21)
截头石栎 Lithocarpus truncatus 10.20 (7.47)
小果栲 Castanopsis fleuryi 8.93(7.35)
滇南木姜子 Litsea garrettii 4.37(6.19)
西南木荷 Schima wallichii 6.13(4.77)
红梗润楠 Machilus rufipes 4.96(2.97)
滇青冈 Cyclobalanopsis glaucoides 35.04(27.95)
白穗石栎 Lithocarpus craibianus 28.38(27.93)
多穗石栎 Lithocarpus polystachyus 20.47(24.96)

Table 2

Correlation coefficients of the NMDS ordination axes and the environmental factors"

Environmental factors
海拔 Elevation -0.779 -0.627 0.897 0.001
凹凸度 Convexity 0.992 -0.128 0.098 0.134
坡向 Aspect -0.025 -1.000 0.035 0.490
坡度 Slope -0.352 -0.936 0.241 0.007
土壤含水率 Soil water content (SW) -0.994 -0.107 0.635 0.001
土壤有机质 Soil organic matter (SOM) -0.825 -0.566 0.293 0.003
土壤pH值 pH 0.998 0.067 0.597 0.001
全氮 Total nitrogen (TN) -0.367 -0.930 0.060 0.312
全磷 Total phosphorus (TP) -0.543 0.840 0.108 0.121
全钾 Total potassium (TK) 0.150 -0.989 0.374 0.002
有效氮 Available nitrogen (AN) -0.582 0.813 0.079 0.210
有效磷 Available phosphorus (AP) -0.918 0.396 0.432 0.001
有效钾 Available potassium (AK) 0.112 0.994 0.316 0.001
有效硼 Available boron (AB) -0.272 0.962 0.503 0.001

Fig. 2

Non-metric multidimensional scaling ordination of forest plots (4 vegetation sub-types) in the Ailao Mountains. Colored circles, symbols “+” and arrows indicate plots, tree species and environmental factors respectively. Notes see Fig. 1 and Table 2."

Table 3

Model selection with the “dredge” function"

pH 全钾
AICc ΔAICc 权重值
1 1.796 -0.128 -0.001 -0.345 5 90.0 0.00 0.261
2 2.082 -0.001 -0.250 4 90.4 0.39 0.214
3 2.374 -0.001 4 91.0 0.95 0.162
4 2.001 -0.005 -0.124 -0.002 -0.409 6 91.3 1.26 0.139
5 2.290 -0.005 -0.002 -0.320 6 91.5 1.44 0.127
6 2.544 0.676 -0.302 5 92.0 2.00 0.096
相对重要性Importance 1.00 0.27 0.40 0.90 0.10 0.84

Fig. 3

Tree species richness of four vegetation sub-types in the Ailao Mountains. Forest types see Fig. 1."

Fig. 4

Responds of tree species richness to environmental factors. The solid line shows the estimations of tree richness, and the dashed lines indicate 95% confidence interval."

1 Baldeck CA, Harms KE, Yavitt JB, John R, Turner BL, Valencia R, Navarrete H, Davies SJ, Chuyong GB, Kenfack D, Thomas DW, Madawala S, Gunatilleke N, Gunatilleke S, Bunyavejchewin S, Kiratiprayoon S, Yaacob A, Supardi MNN, Dalling JW (2013) Soil resources and topography shape local tree community structure in tropical forests. Proceedings of the Royal Society of London B: Biological Sciences, 280, 20122532.
2 Balvanera P, Aguirre E (2006) Tree diversity, environmental heterogeneity, and productivity in a Mexican tropical dry forest. Biotropica, 38, 479-491.
3 Bao SD (2005) Agricultural Chemistry Analysis of Soil. China Agriculture Press, Beijing. (in Chinese)
[鲍士旦 (2005)土壤农化分析. 中国农业出版社, 北京.]
4 Bartoń K (2015) MuMIn: Multi-model Inference. R package version, 1.15.6.
5 Bohlman SA, Laurance WF, Laurance SG, Nascimento HE, Fearnside PM, Andrade A (2008) Importance of soils, topography and geographic distance in structuring central Amazonian tree communities. Journal of Vegetation Science, 19, 863-874.
6 Budke JC, Jarenkow JA, de Oliveira-Filho AT (2007) Relationships between tree component structure, topography and soils of a riverside forest, Rio Botucaraí, Southern Brazil. Plant Ecology, 189, 187-200.
7 Christensen M, Heilmann-Clausen J (2009) Forest biodiversity gradients and the human impact in Annapurna Conservation Area, Nepal. Biodiversity and Conservation, 18, 2205-2221.
8 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.
9 Clark DB, Palmer MW, Clark DA (1999) Edaphic factors and the landscape-scale distributions of tropical rain forest trees. Ecology, 80, 2662-2675.
10 Comprehensive Investigation Group of the Ailao Mountain National Nature Reserve(1988) Reports on a Comprehen- sive Investigation of the Ailao Mountain National Nature Reserve. The Nationalities Publishing House of Yunnan, Kunming. (in Chinese)
[哀牢山自然保护区综合考察团(1988) 哀牢山自然保护区综合考察报告集. 云南民族出版社, 昆明.]
11 de Toledo JJ, Magnusson WE, Castilho CV, Nascimento HE (2011) How much variation in tree mortality is predicted by soil and topography in Central Amazonia? Forest Ecology and Management, 262, 331-338.
12 Dinno A (2015) Nonparametric pairwise multiple comparisons in independent groups using Dunn’s test. The Stata Journal, 15, 292-300.
13 Dinno A (2016) Dunn’s Test of Multiple Comparisons Using Rank Sums, R package version, 132. https://CRAN.R
14 Editorial Committee of the Vegetation of Yunnan(1987) The Vegetation of Yunnan. Science Press, Beijing. (in Chinese)
[云南植被编写组(1987) 云南植被. 科学出版社, 北京.]
15 Engelbrecht BM, Comita LS, Condit R, Kursar TA, Tyree MT, Turner BL, Hubbell SP (2007) Drought sensitivity shapes species distribution patterns in tropical forests. Nature, 447, 80-82.
16 Ferreira-Júnior WG, Silva AF, Schaefer CEGR, Neto JAAM, Dias AS, Ignácio M, Medeiros MCMP (2007) Influence of soils and topographic gradients on tree species distribution in a Brazilian Atlantic tropical semideciduous forest. Edinburgh Journal of Botany, 64, 137-157.
17 Gong HD, Yang GP, Lu ZY, Liu YH, Cao M (2011) Composition and spatio-temporal distribution of tree seedlings in an evergreen broad-leaved forest in the Ailao Mountains, Yunnan. Biodiversity Science, 19, 151-157. (in Chinese with English abstract)
[巩合德, 杨国平, 鲁志云, 刘玉洪, 曹敏 (2011) 哀牢山常绿阔叶林乔木树种的幼苗组成及时空分布特征. 生物多样性, 19, 151-157.]
18 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. (in Chinese with English abstract)
[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.]
19 Guo YL, Wang B, Mallik AU, Huang FZ, Xiang WS, Ding T, Wen SJ, Lu SH, Li DX, He YL, Li XK (2016) Topographic species-habitat associations of tree species in a heterogeneous tropical karst seasonal rain forest, China. Journal of Plant Ecology, 9, 1-11.
20 Homeier J, Breckle SW, Günter S, Rollenbeck RT, Leuschner C (2010) Tree diversity, forest structure and productivity along altitudinal and topographical gradients in a species-rich ecuadorian montane rain forest. Biotropica, 42, 140-148.
21 Itoh A, Yamakura T, Ohkubo T, Kanzaki M, Palmiotto PA, LaFrankie JV, Ashton PS, Lee HS (2003) Importance of topography and soil texture in the spatial distribution of two sympatric dipterocarp trees in a Bornean rainforest. Ecological Research, 18, 307-320.
22 John R, Dalling JW, Harms KE, Yavitt JB, Stallard RF, Mirabello M, Hubbell SP, Valencia R, Navarrete H, Vallejo M, Foster RB (2007) Soil nutrients influence spatial distributions of tropical tree species. Proceedings of the National Academy of Sciences, USA, 104, 864-869.
23 Jones MM, Tuomisto H, Borcard D, Legendre P, Clark DB, Olivas PC (2008) Explaining variation in tropical plant community composition: influence of environmental and spatial data quality. Oecologia, 155, 593-604.
24 Lai JS, Mi XC, Ren HB, Ma KP (2009) Species-habitat associations change in a subtropical forest of China. Journal of Vegetation Science, 20, 415-423.
25 Laurance SG, Laurance WF, Andrade A, Fearnside PM, Harms KE, Vicentini A, Luizão RC (2010) Influence of soils and topography on Amazonian tree diversity: a landscape-scale study. Journal of Vegetation Science, 21, 96-106.
26 Lomolino MV (2001) Elevation gradients of species-density: historical and perspective views. Global Ecology and Biogeography, 10, 3-13.
27 Lundholm JT (2009) Plant species diversity and environmental heterogeneity: spatial scale and competing hypotheses. Journal of Vegetation Science, 20, 377-391.
28 Magurran AE (2004) Measuring Biological Diversity. Blackwell Science Ltd., Blackwell Publishing Company, Victoria, Australia.
29 Oksanen J, Blanchet FG, Kindt R, Legendre P, Minchin PR, O’Hara RB, Simpson GL, Solymos P, Stevens MHH, Wagner H (2016) Vegan: Community Ecology Package, Version 2.3-5. (accessed on 2017-2-3).
30 Oliveira-Filho AT, Vilela EA, Carvalho DA, Gavilanes ML (1994) Effects of soils and topography on the distribution of tree species in a tropical riverine forest in south-eastern Brazil. Journal of Tropical Ecology, 10, 483-508.
31 Oliveira-Filho AD, Curi N, Vilela EA, Carvalho DA (2001) Variation in tree community composition and structure with changes in soil properties within a fragment of semi-deciduous forest in south-eastern Brazil. Edinburgh Journal of Botany, 58, 139-158.
32 Palmiotto PA, Davies SJ, Vogt KA, Ashton MS, Vogt DJ, Ashton PS (2004) Soil-related habitat specialization in dipterocarp rain forest tree species in Borneo. Journal of Ecology, 92, 609-623.
33 Pereira JAA, Oliveira-Filho AT, Lemos-Filho JP (2007) Environmental heterogeneity and disturbance by humans control much of the tree species diversity of Atlantic montane forest fragments in SE Brazil. Plant Conservation and Biodiversity, 16, 1761-1784.
34 Peruggia M (2003) Model selection and multimodel inference: a practical information-theoretic approach. Journal of the American Statistical Association, 98, 778-779.
35 Pinto JRR, Oliveira-Filho AT, Hay JDV (2005) Influence of soil and topography on the composition of a tree community in a central Brazilian valley forest. Edinburgh Journal of Botany, 62, 69-90.
36 Potts MD, Ashton PS, Plotkin JB, Kaufmann LS (2002) Habitat patterns in tropical rainforests: a comparison of 105 plots in northwest Borneo. Ecology, 83, 2782-2797.
37 Poulos HM, Camp AE (2010) Topographic influences on vegetation mosaics and tree diversity in the Chihuahuan Desert Borderlands. Ecology, 91, 1140-1151.
38 Russo SE, Davies SJ, King DA, Tan S (2005) Soil-related performance variation and distributions of tree species in a Bornean rain forest. Journal of Ecology, 93, 879-889.
39 Steidinger B (2015) Qualitative differences in tree species distributions along soil chemical gradients give clues to the mechanisms of specialization: why boron may be the most important soil nutrient at Barro Colorado Island. New Phytologist, 206, 895-899.
40 Takyu M, Aiba S, Kitayama K (2002) Effects of topography on tropical lower montane forests under different geological conditions on Mount Kinabalu, Borneo. Plant Ecology, 159, 35-49.
41 Tateno R, Takeda H (2003) Forest structure and tree species distribution in relation to topography-mediated heterogeneity of soil nitrogen and light at the forest floor. Ecological Research, 18, 559-571.
42 Tripler CE, Kaushal SS, Likens GE, Walter TM (2006) Patterns in potassium dynamics in forest ecosystems. Ecology Letters, 9, 451-466.
43 Tsui CC, Chen ZS, Hsieh CF (2004) Relationships between soil properties and slope position in a lowland rain forest of southern Taiwan. Geoderma, 123, 131-142.
44 Urban DL, Miller C, Halpin PN, Stephenson NL (2000) Forest gradient response in Sierran landscapes: the physical template. Landscape Ecology, 15, 603-620.
45 Wood S (2016) Mixed GAM Computation Vehicle with GCV/AIC/REML Smoothness Estimation, Version 1.8-12.(accessed on 2017-2-3).
46 Wright SJ, Yavitt JB, Wurzburger N, Turner BL, Tanner EV, Sayer EJ, Santiago LS, Kaspari M, Hedin LO, Harms KE, Garcia MN, Corre MD (2011) Potassium, phosphorus, or nitrogen limit root allocation, tree growth, or litter production in a lowland tropical forest. Ecology, 92, 1616-1625.
47 Xu YJ, Chen YN, Li WH, Fu AH, Ma XD, Gui DW, Chen YP (2011) Distribution pattern of plant species diversity in the mountainous region of Ili River Valley, Xinjiang. Environmental Monitoring and Assessment, 177, 681-694.
48 Yamakura T, Kanzaki A, Itoh T, Ohkubo K, Ogino EOK, Chai HS, Ashton PS (1995) Topography of a large-scale research plot established within a tropical rain forest at Lambir, Sarawak. Tropics, 5, 41-56.
49 Yasuhiro K, Hirofumi M, Kihachiro K (2004) Effects of topographic heterogeneity on tree species richness and stand dynamics in a subtropical forest in Okinawa Island, southern Japan. Journal of Ecology, 92, 230-240.
50 Yimer F, Ledin S, Abdelkadir A (2006) Soil property variations in relation to topographic aspect and vegetation community in the south-eastern highlands of Ethiopia. Forest Ecology and Management, 232, 90-99.
[1] Yu Zhang, Gang Feng. Distribution pattern and mechanism of insect species diversity in Inner Mongolia [J]. Biodiv Sci, 2018, 26(7): 701-706.
[2] MENG Ling-Jun, YAO Jie, QIN Jiang-Huan, FAN Chun-Yu, ZHANG Chun-Yu, ZHAO Xiu-Hai. Drivers of composition and density pattern of tree seedlings in a secondary mixed conifer and broad-leaved forest, Jiaohe, Jilin, China [J]. Chin J Plan Ecolo, 2018, 42(6): 653-662.
[3] Qian SUN, Zi-Ke XUE, Lin-Lin XIE, Xue-Li HE, Li-Li ZHAO. Diversity of dark septate endophyte in the roots of Ammopiptanthus mongolicus and its companion plants [J]. Chin J Plan Ecolo, 2017, 41(7): 729-737.
[4] Mingfei Zhao,Guoyi Wang,Kaixiong Xing,Yuhang Wang,Feng Xue,Muyi Kang,Kai Luo. Patterns and determinants of species similarity decay of forest communities in the western Qinling Mountains [J]. Biodiv Sci, 2017, 25(1): 3-10.
[5] Shuai LIU, Jia-Xing LIAO, Cui XIAO, Xiu-Hua FAN. Effects of biotic neighbors and habitat heterogeneity on tree seedling survival in a secondary mixed conifer and broad-leaved forest in Changbai Mountain [J]. Chin J Plan Ecolo, 2016, 40(7): 711-722.
[6] Gexi Xu,Zuomin Shi,Jingchao Tang,Han Xu,Huai Yang,Shirong Liu,Yide Li,Mingxian Lin. Effects of species abundance and size classes on assessing community phylogenetic structure: a case study in Jianfengling tropical montane rainforest [J]. Biodiv Sci, 2016, 24(6): 617-628.
[7] Yun CHEN, Ting WANG, Pei-Kun LI, Cheng-Liang YAO, Zhi-Liang YUAN, Yong-Zhong YE. Community characteristics and spatial distribution of dominant tree species in a deciduous broad-leaved forest of Muzhaling, Henan, China [J]. Chin J Plan Ecolo, 2016, 40(11): 1179-1188.
[8] Yuanjie Xu,Dunmei Lin,Xiangcheng Mi,Haibao Ren,Keping Ma. Recovery dynamics of secondary forests with different disturbance intensity in the Gutianshan National Nature Reserve [J]. Biodiv Sci, 2014, 22(3): 358-365.
[9] Jun Gong,Yanjing Song,Xiaoli Zhang. Phylogenetic and functional diversity of nitrogen cycling microbes in coastal sediments [J]. Biodiv Sci, 2013, 21(4): 433-444.
[10] Shengbin Chen, Zhiyun Ouyang, Yu Fang, Zhenji Li. Geographic patterns of endemic seed plant genera diversity in China [J]. Biodiv Sci, 2011, 19(4): 414-423.
[11] Kai Song, Xiangcheng Mi, Qi Jia, Haibao Ren, Dan Bebber, Keping Ma. Variation in phylogenetic structure of forest communities along a human disturbance gradient in Gutianshan forest, China [J]. Biodiv Sci, 2011, 19(2): 190-196.
[12] Hede Gong, Guoping Yang, Zhiyun Lu, Yuhong Liu. Diversity and spatial distribution patterns of trees in an evergreen broad-leaved forest in the Ailao Mountains, Yunnan [J]. Biodiv Sci, 2011, 19(2): 143-150.
[13] Haifeng Liu, Liang Li, Weiguo Sang. Species composition and community structure of the Donglingshan forest dynamic plot in a warm temperate deciduous broad-leaved secondary forest, China [J]. Biodiv Sci, 2011, 19(2): 232-242.
[14] GUO Ke, LIU Chang-Cheng, and DONG Ming. Ecological adaptation of plants and control of rocky-desertification on karst region of Southwest China [J]. Chin J Plan Ecolo, 2011, 35(10): 991-999.
[15] Xinxin Sun, Huirong Liu, Fuying Feng, Jianyu Meng, Heng Li, Malina. Diversity and phylogenetic analysis of planktonic bacteria in eutrophic zone of Lake Wuliangsuhai [J]. Biodiv Sci, 2009, 17(5): 490-498.
Full text



[1] WANG Hong-Xing, CHEN Xin, TANG Jian-Jun, SHIMIZU Katsuyoshi. Influence of released transgenic pest and disease resistant crops on plant associated microorganisms in soil[J]. Biodiv Sci, 2002, 10(2): 232 -237 .
[2] . [J]. Biodiv Sci, 2013, 21(1): 136 .
[3] Qin SUN, Xiao-Rong WANG, Shi-Ming DING,Xin-Fang YUAN. Effects of Interaction Between Cadmium and Plumbum on Phytochelatins and Glutathione Production in Wheat (Triticum aestivum L.)[J]. J Integr Plant Biol, 2005, 47(4): .
[4] Cao Jiashu, Miao Ying. Evolutionary principle and conservation strategies of biodiversity[J]. Biodiv Sci, 1997, 05(3): 220 -223 .
[5] . [J]. Chin J Plan Ecolo, 2015, 39(11): 1123 .
[6] CUI Xiao-Yong, Du Zhan-Chi, Wang Yan-Fen. Photosynthetic Characteristics of a Semi-arid Sandy Grassland Community in Inner Mongolia[J]. Chin J Plan Ecolo, 2000, 24(5): 541 -546 .
[7] Kao Tso-Ching, Cheng Ching-Yung. New Taxa of the Chinese Celastraceae (1)[J]. J Syst Evol, 1988, 26(4): 310 -314 .
[8] CHANG Chen-Hui,WU Fu-Zhong,YANG Wan-Qin,TAN Bo,XIAO Sa,LI Jun,GOU Xiao-Lin. Changes in log quality at different decay stages in an alpine forest[J]. Chin J Plan Ecolo, 2015, 39(1): 14 -22 .
[9] ZHAN Zha_Jun, SUN Han_Dong, WU Hou_Ming and YUE Jian_Min. Chemical Components from the Fungus Englero myces goetzei[J]. J Integr Plant Biol, 2003, 45(2): 248 -252 .
[10] Li Zhong-Ming. Palaeosmunda Emended and Two New Species[J]. J Syst Evol, 1983, 21(2): 153 -160 .