Biodiversity Science ›› 2019, Vol. 27 ›› Issue (6): 619-629.doi: 10.17520/biods.2019107

• Original Papers • Previous Article     Next Article

Vertical structure and its biodiversity in a subtropical evergreen broad- leaved forest at Dinghushan in Guangdong Province, China

Gui Xujun1, 2, 3, Lian Juyu1, 2, *(), Zhang Ruyun1, 2, 3, Li Yanpeng1, 2, 3, Shen Hao1, 2, Ni Yunlong1, 2, 3, Ye Wanhui1, 2   

  1. 1 Key Laboratory of Vegetation Restoration and Management of Degraded Ecosystems, South China Botanical Garden, Chinese Academy of Sciences, Guangzhou 510650
    2 Guangdong Provincial Key Laboratory of Applied Botany, South China Botanical Garden, Chinese Academy of Sciences, Guangzhou 510650
    3 University of Chinese Academy of Sciences, Beijing 100049
  • Received:2019-03-28 Accepted:2019-05-28 Online:2019-07-08
  • Lian Juyu

Community structure plays a fundamental role in forest ecosystems as one of the basic mechanisms underlying community assembly. Community structure includes both horizontal and vertical structure. We stratified the vertical structure of a lower subtropical evergreen broad-leaved forest at Dinghushan in Guangdong Province, China by surveying each individual with DBH ≥ 1 cm via a canopy crane, and then explored α diversity within each layer, as well as β diversity between layers. We found that: (1) The vertical structure of the forest was stratified into five layers; from bottom to top, the shrub, sub-canopy, lower canopy, middle canopy, and upper canopy correspond to the traditionally defined vertical levels of a forest community. (2) Layer α diversity decreased with height, and the Pielou evenness index was largest in the lower canopy. (3) The β diversity of layers relative to the shrub layer increased with height. Using the POD framework, the differences between layer community compositions were mainly attributed to differences in species richness. However, the lower canopy showed increased species replacement and decreased richness difference relative to the middle canopy layer when compared with other neighbouring layers. (4) Air temperature, light intensity and relative humidity increased with layer height, and the most dramatic change in light occurred in the middle canopy. These microenvironmental features may play an important role in the formation of vertical hierarchy in the forest, with light intensity as the largest factor.

Key words: forest structure, vertical structure, β diversity, β diversity partitioning, biodiversity, stratification, community assembly

Fig. 1

The judgment index partitioning optimal groups. (a) Attribution of the objects and the corresponding ssi (simple structure index) values for different number of groups. The panel on the left shows the groups that each object is categorized with different conditions and different groups were distinguished by different colors; the panel on the right shows the value of ssi for different number of groups. (b) Value of sum of squares within groups for different number of groups."

Table 1

Species abundance and richness of each vertical layer in the crane plot of Dinghushan"

Minimum height (m)
Maximum height (m)
Species richness
优势种例举(多度, 最大胸径)
Dominant species (abundance, maximum DBH)
1.4 3.9 1,966 89 银柴
Aporosa dioica (207, 7.3)
Cryptocarya concinna (251, 4.5)
4.0 6.7 1,173 75 鸭脚木
Schefflera octophylla (161, 16.0)
Aporosa dioica (122, 10.1)
Lower canopy
6.8 11.2 503 56 荷木
Schima superba (54, 39.4)
Schefflera octophylla (51, 18.8)
Middle canopy
11.3 17.2 271 27 荷木
Schima superba (101, 41.9)
Castanopsis chinensis (31, 41.0)
Upper canopy
17.4 27.1 226 17 荷木
Schima superba (111, 66.0)
Pinus massoniana (34, 47.1)
- - 4,140 121 荷木
Schima superba (301,49.0)
Pinus massoniana (66, 47.1)

Table 2

Comparison of species component between shrub and other layers as well as among neighbouring layers in pair in the crane plot of Dinghushan"

Generalist (%)
Specialist (%)
Too rare to clarify (%)
灌木层和亚冠层 Shrub vs sub-canopy 24 (23.08) 4 (3.98) 76 (73.08) 104
灌木层和林冠下层 Shrub vs lower canopy 17 (16.50) 7 (6.79) 79 (76.70) 103
灌木层和林冠中层 Shrub vs middle canopy 8 (8.42) 11 (11.58) 76 (80.00) 95
灌木层和林冠上层 Shrub vs upper canopy 1 (1.06) 14 (14.89) 79 (84.08) 94
亚冠层和林冠下层 Sub-canopy vs lower canopy 19 (20.43) 3 (3.23) 71 (76.34) 93
林冠下层和林冠中层 Lower canopy vs middle canopy 10 (17.24) 4 (6.89) 44 (75.86) 58
林冠中层和林冠上层 Middle canopy vs upper canopy 5 (21.80) 4 (3.13) 23 (75.00) 32

Table 3

Species α diversity of each layer in the crane plot of Dinghushan"

Peliou evenness index
灌木层 Shrub 2.556 ± 0.335a 0.910 ± 0.040a
亚冠层 Sub-canopy 2.300 ± 0.529b 0.917 ± 0.033ab
林冠下层 Lower canopy 1.955 ± 0.334c 0.940 ± 0.027ab
林冠中层 Middle canopy 1.440 ± 0.497d 0.925 ± 0.232ab
林冠上层 Upper canopy 1.172 ± 0.519e 0.896 ± 0.275b

Fig. 2

Results of β diversity (S?rensen dissimilarity) and their partitions based on species abundance data of each layers in the crane plot of Dinghushan. (a) Layers relative to the shrub layer. (b) Neighbouring layers. S, Shrub; SC, Sub-canopy; LC, Lower canopy; MC, Middle canopy; UC, Upper canopy."

Fig. 3

Tree height scatter points of each individual in the crane plot of Dinghushan"

[1] Angeler DG ( 2013) Revealing a conservation challenge through partitioned long-term beta diversity: Increasing turnover and decreasing nestedness of boreal lake metacommunities. Diversity and Distributions, 19, 772-781.
doi: 10.1111/ddi.2013.19.issue-7
[2] Ashton PS, Hall P ( 1992) Comparisons of structure among mixed dipterocarp forests of north-western Borneo. Journal of Ecology, 80, 459-481.
doi: 10.2307/2260691
[3] Baselga A ( 2010) Partitioning the turnover and nestedness components of beta diversity. Global Ecology and Biogeography, 19, 134-143.
doi: 10.1111/j.1466-8238.2009.00490.x
[4] Baselga A, Orme CDL ( 2012) Betapart: An R package for the study of beta diversity. Methods in Ecology and Evolution, 3, 808-812.
doi: 10.1111/j.2041-210X.2012.00224.x
[5] Baselga A ( 2013) Separating the two components of abundance-based dissimilarity: Balanced changes in abundance vs. abundance gradients. Methods in Ecology and Evolution, 4, 552-557.
doi: 10.1111/2041-210X.12029
[6] Brown JH ( 2001) Mammals on mountainsides: Elevational patterns of diversity. Global Ecology and Biogeography, 10, 101-109.
doi: 10.1046/j.1466-822x.2001.00228.x
[7] Chazdon RL, Chao A, Colwell RK, Lin SY, Norden N, Letcher SG, Clark DB, Arroyo JP ( 2011) A novel statistical method for classifying habitat generalists and specialists. Ecology, 92, 1332-1343.
doi: 10.1890/10-1345.1
[8] Chen SB, Ouyang ZY, Xu WH, Xiao Y ( 2010) A review of beta diversity studies. Biodiversity Science, 18, 323-335. (in Chinese with English abstract)
[ 陈圣宾, 欧阳志云, 徐卫华, 肖燚 ( 2010) Beta多样性研究进展. 生物多样性, 18, 323-335.]
[9] Condit R ( 1998) Tropical Forest Census Plots: Methods and Results from Barro Colorado Island, Panama and a Comparison with Other Plots. Springer-Verlag, Berlin.
[10] Dolnicar S, Grabler K, Mazanec JA ( 1999) A tale of three cities: Perceptual charting for analyzing destination images. In: Consumer Psychology of Tourism, Hospitality and Leisure (eds Sakai MY, Woodside AG, Crouch GI, Mazanec JA, Oppermann M), pp. 39-62. CAB International, New York.
[11] Fukami T ( 2015) Historical contingency in community assembly: Integrating niches, species pools, and priority effects. Annual Review of Ecology, Evolution and Systematics, 46, 1-23.
doi: 10.1146/annurev-ecolsys-110411-160340
[12] Gillespie TW, Brock J, Wright CW ( 2014) Prospects for quantifying structure, floristic composition and species richness of tropical forests. International Journal of Remote Sensing, 25, 707-715.
[13] Grubb PJ, Lloyd JR, Pennington TD, Whitmore TC ( 1963) A comparison of montane and lowland rain forest in ecuador. I. The forest structure, physiognomy, and floristics. Journal of Ecology, 51, 567-601.
doi: 10.2307/2257748
[14] Hao ZQ, Zhang J, Song B, Ye J, Li B ( 2007) Vertical structure and spatial associations of dominant tree species in an old- growth temperate forest. Forest Ecology and Management, 252, 1-11.
doi: 10.1016/j.foreco.2007.06.026
[15] Harms KE, Condit R, Hubbell SP, Foster RB ( 2001) Habitat associations of trees and shrubs in a 50-ha neotropical forest plot. Journal of Ecology, 89, 947-959.
doi: 10.1111/jec.2001.89.issue-6
[16] Harrison S, Ross SJ, Lawton JH ( 1992) Beta diversity on geographic gradients in Britain. Journal of Animal Ecology, 61, 151-158.
doi: 10.2307/5518
[17] HilleRisLambers J, Adler PB, Harpole WS, Levine JM, Mayfield MM ( 2012) Rethinking community assembly through the lens of coexistence theory. Annual Review of Ecology, Evolution, and Systematics, 43, 227-248.
[18] Huang ZL, Kong GH, Wei P ( 1998) Plant species diversity dynamics in Dinghu Mountain forests. Chinese Biodiversity, 6, 116-121. (in Chinese with English abstract)
[ 黄忠良, 孔国辉, 魏平 ( 1998) 鼎湖山植物物种多样性动态. 生物多样性, 6, 116-121.]
[19] Latham PA, Zuuring HR, Coble DW ( 1998) A method for quantifying vertical forest structure. Forest Ecology and Management, 104, 157-170.
doi: 10.1016/S0378-1127(97)00254-5
[20] Legendre P ( 2014) Interpreting the replacement and richness difference components of beta diversity. Global Ecology and Biogeography, 23, 1324-1334.
doi: 10.1111/geb.12207
[21] Legendre P, De Cáceres M ( 2013) Beta diversity as the variance of community data: Dissimilarity coefficients and partitioning. Ecology Letters, 16, 951-963.
doi: 10.1111/ele.2013.16.issue-8
[22] Lennon JJ, Koleff P, Greenwood J, Gaston KJ ( 2001) The geographical structure of British bird distributions: Diversity, spatial turnover and scale. Journal of Animal Ecology, 70, 966-979.
doi: 10.1046/j.0021-8790.2001.00563.x
[23] Li DZ, Zang RG ( 2004) The research advances on the structure and function of forest canopy, as well as their temporal and spatial changes. World Forestry Research, 17(3), 12-16. (in Chinese with English abstract)
[ 李德志, 臧润国 ( 2004) 森林冠层结构与功能及其时空变化研究进展. 世界林业研究, 17(3), 12-16.]
[24] Li SP, Cadotte MW, Meiners SJ, Pu Z, Fukami T, Jiang L ( 2016) Convergence and divergence in a long-term old-field succession: The importance of spatial scale and species abundance. Ecology Letters, 19, 1101-1109.
doi: 10.1111/ele.12647
[25] Lowman MD, Rinker HB ( 2004) Forest Canopies, 2nd edn. Elsevier Academic Press, San Diego.
[26] Lowman MD, Wittman PK ( 1996) Forest canopies: Methods, hypotheses, and future directions. Annual Review of Ecology and Systematics, 27, 55-81.
doi: 10.1146/annurev.ecolsys.27.1.55
[27] Ma KP, Liu YM ( 1994) Measurement of biotic community diversity. I. α diversity (Part 2). Chinese Biodiversity, 2, 231-239. (in Chinese)
[ 马克平, 刘玉明 ( 1994) 生物群落多样性的测度方法. I. α多样性的测度方法(下). 生物多样性, 2, 231-239.]
[28] McCain CM, Beck J ( 2016) Species turnover in vertebrate communities along elevational gradients is idiosyncratic and unrelated to species richness. Global Ecology and Biogeography, 25, 299-310.
doi: 10.1111/geb.12410
[29] Neto RB ( 1989) Rain-forest canopy remains elusive. Nature, 340, 586.
[30] Perry DR ( 1978) A method of access into the crowns of emergent and canopy trees. Biotropica, 10, 155-157.
doi: 10.2307/2388019
[31] Pielou EC ( 1975) Ecological Diversity. Wiley, New York.
[32] Podani J, Schmera D ( 2011) A new conceptual and methodological framework for exploring and explaining pattern in presence-absence data. Oikos, 120, 1625-1638.
doi: 10.1111/j.1600-0706.2011.19451.x
[33] Rahbek C ( 2005) The role of spatial scale and the perception of large-scale species-richness patterns. Ecology Letters, 8, 224-239.
[34] Ricklefs RE ( 1987) Community diversity: Relative roles of local and regional processes. Science, 235, 167-171.
doi: 10.1126/science.235.4785.167
[35] Rosenzweig ML ( 1995) Species Diversity in Space and Time. Cambridge University Press, Cambridge.
[36] Shen H, Cai JN, Li MJ, Chen Q, Ye WH, Wang ZF, Lian JY, Song L ( 2017) On Chinese forest canopy biodiversity monitoring. Biodiversity Science, 25, 229-236. (in Chinese with English abstract)
[ 沈浩, 蔡佳宁, 李萌姣, 陈青, 叶万辉, 王峥峰, 练琚愉, 宋亮 ( 2017) 中国森林冠层生物多样性监测. 生物多样性, 25, 229-236.]
[37] Si XF, Baselga A, Leprieur F, Song X, Ding P ( 2016) Selective extinction drives taxonomic and functional alpha and beta diversities in island bird assemblages. Journal of Animal Ecology, 85, 409-418.
doi: 10.1111/1365-2656.12478
[38] Si XF, Zhao YH, Chen CW, Ren P, Zeng Y, Wu LB, Ding P ( 2017) Beta-diversity partitioning: Methods, applications and perspectives. Biodiversity Science, 25, 464-480. (in Chinese with English abstract)
[ 斯幸峰, 赵郁豪, 陈传武, 任鹏, 曾頔, 吴玲兵, 丁平 ( 2017) Beta多样性分解: 方法、应用与展望. 生物多样性, 25, 464-480.]
[39] Smith AP ( 1973) Stratification of temperate and tropical forests. The American Naturalist, 107, 671-683.
doi: 10.1086/282866
[40] Sun RY, Li QF, Niu CJ, Lou AR ( 2007) Foundations in Ecology, 2nd edn. Higher Education Press, Beijing. (in Chinese)
[ 孙儒泳, 李庆芬, 牛翠娟, 娄安如 ( 2007) 基础生态学(第二版). 高等教育出版社, 北京.]
[41] Ulrich W, Almeida-Neto M, Gotelli NJ ( 2009) A consumer’s guide to nestedness analysis. Oikos, 118, 3-17.
doi: 10.1111/oik.2009.118.issue-1
[42] Watt AS ( 1924) On the ecology of British beechwoods with special reference to their regeneration. Part II. The development and structure of beech communities on the Sussex Downs. Journal of Ecology, 12, 145-204.
doi: 10.2307/2255242
[43] Whittaker RH ( 1972) Evolution and measurement of species diversity. Taxon, 21, 213-215.
doi: 10.2307/1218190
[44] Williams PH ( 1996) Mapping variations in the strength and breadth of biogeographic transition zones using species turnover. Proceedings of the Royal Society of London B: Biological Sciences, 263, 579-588.
doi: 10.1098/rspb.1996.0087
[45] Wu Y, Liu WY, Song L, Chen X, Lu HZ, Li S, Shi XM ( 2016) Advances in ecological studies of epiphytes using canopy cranes. Chinese Journal of Plant Ecology, 40, 508-522. (in Chinese with English abstract)
[ 吴毅, 刘文耀, 宋亮, 陈曦, 卢华正, 李苏, 石贤萌 ( 2016) 基于林冠塔吊的附生植物生态学研究进展. 植物生态学报, 40, 508-522.]
[46] Yang L, Sun ZY, Tang GL, Lin ZW, Chen YQ, Li Y, Li Y ( 2016) Identifying canopy species of subtropical forest by lightweight unmanned aerial vehicle remote sensing. Tropical Geography, 36, 833-839. (in Chinese with English abstract)
[ 杨龙, 孙中宇, 唐光良, 林志文, 陈燕乔, 黎喻, 李勇 ( 2016) 基于微型无人机遥感的亚热带林冠物种识别. 热带地理, 36, 833-839.]
[47] Ye WH, Cao HL, Huang ZL, Lian JY, Wang ZG, Li L, Wei SG, Wang ZM ( 2008) Community structure of a 20 hm 2 lower subtropical evergreen broadleaved forest plot in Dinghushan, China . Journal of Plant Ecology (Chinese Version), 32, 274-286. (in Chinese with English abstract)
[ 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.]
[48] Zhao J, Li J, Liu QH ( 2013) Review of forest vertical structure parameter inversion based on remote sensing technology. Journal of Remote Sensing, 17, 697-716. (in Chinese with English abstract)
[ 赵静, 李静, 柳钦火 ( 2013) 森林垂直结构参数遥感反演综述. 遥感学报, 17, 697-716.]
[49] Zhao MF, Xing KX, Wang YH ( 2017) Patterns and determinants of beta diversity in the understory vegetation layers of montane boreal conifer forest in Luya Mountain. Acta Ecologica Sinica, 37, 3327-3334. (in Chinese with English abstract)
[ 赵鸣飞, 邢开雄, 王宇航 ( 2017) 芦芽山寒温性针叶林冠层下植被beta多样性格局及其成因. 生态学报, 37, 3327-3334.]
[50] Zhao ZM, Guo YQ ( 1993) Principles and Methods of Community Ecology. Scientific and Technical Documents Publishing House, Chongqing. (in Chinese)
[ 赵志模, 郭依泉 ( 1993) 群落生态学原理与方法. 科技文献出版社重庆分社, 重庆.]
[51] Zhuang CY, Huang QL, Ma ZB, Luo F, Zhang Y ( 2014) Review on defining methods for canopy stratification. World Forestry Research, 27(6), 34-40. (in Chinese with English abstract)
[ 庄崇洋, 黄清麟, 马志波, 罗芬, 张寅 ( 2014) 林层划分方法综述. 世界林业研究, 27(6), 34-40.]
[1] Lintao Huang Hui Huang Lei Jiang. (2020) A revised taxonomy for Chinese hermatypic corals . Biodiv Sci, 28(4): 515-523.
[2] Xiongwei Yang,Ankang Wu,Qixian Zou,Guangrong Li,Mingming Zhang,Canshi Hu,Haijun Su. (2020) Field monitoring of mammals and birds using infrared cameras in Mayanghe National Nature Reserve, Guizhou, China . Biodiv Sci, 28(2): 219-225.
[3] Haiou Liu,Fengchun Zhang,Fuwei Zhao,Leshan Du,Dayuan Xue. (2020) Biodiversity sensitive issues from changes in the strategic objectives of the financial mechanism for the Convention on Biological Diversity . Biodiv Sci, 28(2): 244-252.
[4] Yisheng Ma,Qingqing Ma,Nianjun He,Dapeng Zhu,Kaihui Zhao,Hongcai Liu,Shuai Li,Liang Sun,Liubin Tang. (2020) Camera-trapping survey of mammals and birds in the Foping National Nature Reserve, China . Biodiv Sci, 28(2): 226-230.
[5] Kai Wang,Jinlong Ren,Hongman Chen,Zhitong Lyu,Xianguang Guo,Ke Jiang,Jinmin Chen,Jiatang Li,Peng Guo,Yingyong Wang,Jing Che. (2020) The updated checklists of amphibians and reptiles of China . Biodiv Sci, 28(2): 189-218.
[6] Yi Li,Zhiyao Tang,Yujing Yan,Ke Wang,Lei Cai,Jinsheng He,Song Gu,Yijian Yao. (2020) Incorporating species distribution model into the red list assessment and conservation of macrofungi: A case study with Ophiocordyceps sinensis . Biodiv Sci, 28(1): 99-106.
[7] Wenying Zhuang,Yi Li,Huandi Zheng,Zhaoqing Zeng,Xincun Wang. (2020) Threat status of non-lichenized macro-ascomycetes in China and its threatening factors . Biodiv Sci, 28(1): 26-40.
[8] WANG Yu-Bing,SUN Yi-Han,DING Wei,ZHANG En-Tao,LI Wen-Huai,CHI Yong-Gang,ZHENG Shu-Xia. (2020) Effects and pathways of long-term nitrogen addition on plant diversity and primary productivity in a typical steppe . Chin J Plant Ecol, 44(1): 22-32.
[9] TANG Li-Li,ZHANG Mei,ZHAO Xiang-Lin,KANG Mu-Yi,LIU Hong-Yan,GAO Xian-Ming,YANG Tong,ZHENG Pu-Fan,SHI Fu-Chen. (2019) Species distribution and community assembly rules of Juglans mandshurica in North China . Chin J Plant Ecol, 43(9): 753-761.
[10] SHI Jing-Jing,ZHAO Ming-Fei,WANG Yu-Hang,XUE Feng,KANG Mu-Yi,JIANG Yuan. (2019) Community assembly of herbaceous layer of the planted forests in the central Loess Plateau, China . Chin J Plant Ecol, 43(9): 834-842.
[11] Shun Li,Liang Zou,Yinan Gong,Haitao Yang,Tianming Wang,Limin Feng,Jianping Ge. (2019) Advances in LiDAR technology in the field of animal ecology . Biodiv Sci, 27(9): 1021-1031.
[12] Rui Yang,Qinyi Peng,Yue Cao,Le Zhong,Shuyu Hou,Zhicong Zhao,Cheng Huang. (2019) Transformative changes and paths toward biodiversity conservation in China . Biodiv Sci, 27(9): 1032-1040.
[13] CHAI Yong-Fu,XU Jin-Shi,LIU Hong-Yan,LIU Quan-Ru,ZHENG Cheng-Yang,KANG Mu-Yi,LIANG Cun-Zhu,WANG Ren-Qing,GAO Xian-Ming,ZHANG Feng,SHI Fu-Chen,LIU Xiao,YUE Ming. (2019) Species composition and phylogenetic structure of major shrublands in North China . Chin J Plant Ecol, 43(9): 793-805.
[14] XU Jin-Shi,CHAI Yong-Fu,LIU Xiao,YUE Ming,GUO Yao-Xin,KANG Mu-Yi,LIU Quan-Ru,ZHENG Cheng-Yang,JI Cheng-Jun,YAN Ming,ZHANG Feng,GAO Xian-Ming,WANG Ren-Qing,SHI Fu-Chen,ZHANG Qin-Di,WANG Mao. (2019) Community assembly, diversity patterns and distributions of broad-leaved forests in North China . Chin J Plant Ecol, 43(9): 732-741.
[15] Yongmin Li,Xiaobing Wu. (2019) A revised species list of amphibians and reptiles in the Anhui Province . Biodiv Sci, 27(9): 1002-1011.
Full text