Soil faunal diversity characteristics and influencing factors across the altitude gradient in the poplar-birch forest of Guandi Mountains

doi:10.17520/biods.2024070

Abstract

Abstract:

Aim: The purpose of this study was to clarify the changes of a soil fauna community in the poplar-birch forest of Guandi Mountains along the altitude gradient and the resulting effects of environmental factors on its diversity.
Methods: Four sampling plots were set up along an elevation gradient in the poplar-birch forest (1,600 m, 1,800 m, 2,000 m, 2,200 m). Soil fauna in different soil layers under the forest were investigated in May, July and October of 2022 and 2023. The different characteristics of soil fauna community structure and diversity index in the poplar-birch forest at different elevations were analyzed by the one-way univariate analysis, and using Jaccard similarity index and Cody index to compare the similarity of the soil fauna community at different elevations. Redundancy analysis was used to identify the environmental factors affecting the diversity of soil fauna at different elevations.
Results: The results showed that a total of 24,434 soil fauna of 127 species were captured in the forest plots across different elevations at Guandi Mountains. There were obvious differences in the differentiation characteristics of community composition along the elevation gradient. With the exception of the Simpson index, the α diversity indices of soil fauna communities at different elevations were significantly different (P < 0.05). The Jaccard index was very similar between the two adjacent elevations, and the species richness similarity was higher. Shannon-Wiener, Pielou and Margalef indices of soil fauna at different elevations all decreased with the increase of soil depth, suggesting super-aggregation. In terms of seasonal dynamics, diversity, richness and evenness indices were highest in the summer. Furthermore, the results of the redundancy analysis suggest that temperature, humidity and pH were the main environmental factors affecting soil fauna diversity at different elevations.
Conclusion: The diversity of the soil fauna community is different across elevation gradients, soil layers and seasonal dynamics. Environmental temperature, humidity and pH are the main environmental factors affecting the diversity of soil fauna community.

Key words: elevation gradient, seasonal dynamics, poplar-birch forest, biodiversity

Rongjiao Li, Jianghai Dong, Wenfang Zheng, Ruyuan Liu, Lijuan Zhao, Ruihe Gao. Soil faunal diversity characteristics and influencing factors across the altitude gradient in the poplar-birch forest of Guandi Mountains[J]. Biodiv Sci, 2024, 32(9): 24070.

Add to citation manager EndNote|Ris|BibTeX

URL: https://www.biodiversity-science.net/EN/10.17520/biods.2024070

https://www.biodiversity-science.net/EN/Y2024/V32/I9/24070

Figures/Tables 10

References 50

[1]	Blankinship JC, Niklaus PA, Hungate BA (2011) A meta-analysis of responses of soil biota to global change. Oecologia, 165, 553-565. DOI PMID
[2]	Chen JX, Ma ZC, Yan HJ, Zhang F (2007) Roles of springtails in soil ecosystem. Biodiversity Science, 15, 154-161. (in Chinese with English abstract) DOI
	[陈建秀, 麻智春, 严海娟, 张峰 (2007) 跳虫在土壤生态系统中的作用. 生物多样性, 15, 154-161.] DOI
[3]	Chen YN, Liang C, Chen J (2022) The composition of the community structure of oribatid mites in subtropical forests of different tree species: A case study of Xingangshan, Jiangxi Province. Biodiversity Science, 30, 22334. (in Chinese with English abstract) DOI
	[陈燕南, 梁铖, 陈军 (2022) 亚热带不同树种组成森林中土壤甲螨群落结构特征: 以江西新岗山为例. 生物多样性, 30, 22334.] DOI
[4]	Chen YQ, Sun T, Zang JC, Hong DW (2021) Relationship between diversity of surface arthropods at different altitudes and seed damage of Sophora moorcroftiana shrubs. Southwest China Journal of Agricultural Sciences, 34, 1540-1547. (in Chinese with English abstract)
	[陈韵秋, 孙涛, 臧建成, 洪大伟 (2021) 砂生槐灌丛不同海拔地表节肢动物多样性与砂生槐种子危害的关系. 西南农业学报, 34, 1540-1547.]
[5]	Han YR, Xue QQ, Song HJ, Qi JY, Gao RH, Cui SP, Men LN, Zhang ZW (2022) Diversity and influencing factors of flower-visiting insects in the Yanshan area. Biodiversity Science, 30, 21448. (in Chinese with English abstract) DOI
	[韩艺茹, 薛琪琪, 宋厚娟, 祁靖宇, 高瑞贺, 崔绍朋, 门丽娜, 张志伟 (2022) 燕山地区访花昆虫多样性及其影响因子. 生物多样性, 30, 21448.] DOI
[6]	Hsieh TC, Ma KH, Chao A (2016) iNEXT: An R package for rarefaction and extrapolation of species diversity (Hill numbers). Methods in Ecology and Evolution, 7, 1451-1456.
[7]	Huang HY, Liu JH, Yang JS, Hu JD, Wang JY, Zhu SH (2018) Community structure and characteristics of seasonal change in number of soil animals under Saccharum arundinaceum root in different remediation years. Journal of Shaoguan University, 39(3), 58-63. (in Chinese with English abstract)
	[黄红英, 刘静华, 杨家绅, 胡金德, 王佳颖, 祝三华 (2018) 不同修复年限斑茅根系下土壤动物群落结构及季节特征. 韶关学院学报, 39(3), 58-63.]
[8]	Kagawa S, Suh S, Hubacek K, Wiedmann T, Nansai K, Minx J (2015) CO₂ emission clusters within global supply chain networks: Implications for climate change mitigation. Global Environmental Change, 35, 486-496.
[9]	Keylock CJ (2005) Simpson diversity and the Shannon-Wiener index as special cases of a generalized entropy. Oikos, 109, 203-207.
[10]	Koehler H (1998) Secondary succession of soil mesofauna: A thirteen year study. Applied Soil Ecology, 9, 81-86.
[11]	Körner C (2007) The use of ‘altitude’ in ecological research. Trends in Ecology & Evolution, 22, 569-574.
[12]	Laiolo P, Pato J, Obeso JR (2018) Ecological and evolutionary drivers of the elevational gradient of diversity. Ecology Letters, 21, 1022-1032. DOI PMID
[13]	Lee KE, Foster RC (1991) Soil fauna and soil structure. Australian Journal of Soil Research, 29, 745-775.
[14]	Li M, Wu PF, Wang Y (2015) Vertical distributions of soil fauna communities on the eastern slope of Gongga Mountain. Acta Ecologica Sinica, 35, 2295-2307. (in Chinese with English abstract)
	[李萌, 吴鹏飞, 王永 (2015) 贡嘎山东坡典型植被类型土壤动物群落特征. 生态学报, 35, 2295-2307.]
[15]	Li SZ, Wang SJ, Liu JA, Wu Y, He YH, Li H (2015) Research on soil nematode community structure and diversity of different Cunninghamia lanceolata plantations. Journal of Central South University of Forestry & Technology, 35(10), 101-108. (in Chinese with English abstract)
	[李树战, 王圣洁, 刘君昂, 吴毅, 何苑皞, 李河 (2015) 不同杉木人工林土壤线虫群落结构及多样性研究. 中南林业科技大学学报, 35(10), 101-108.]
[16]	Li W, Cui LJ, Zhao XS, Zhang MY, Gao CJ, Zhang Y, Wang YF (2015) Community structure and diversity of soil animals in the Lake Taihu lakeshore wetland. Acta Ecologica Sinica, 35, 944-955. (in Chinese with English abstract)
	[李伟, 崔丽娟, 赵欣胜, 张曼胤, 高常军, 张岩, 王义飞 (2015) 太湖岸带湿地土壤动物群落结构与多样性. 生态学报, 35, 944-955.]
[17]	Li XH, Ma J, Gao B, Wang RY, Chen SL (2018) The hatching of cereal cyst nematode (Heteroderaavenae) at different soil depth in the field. Journal of Hebei Agricultural University, 41(1), 70-75. (in Chinese with English abstract)
	[李秀花, 马娟, 高波, 王容燕, 陈书龙 (2018) 不同土壤深度禾谷孢囊线虫的孵化特点. 河北农业大学学报, 41(1), 70-75.] DOI
[18]	Lin LW, Zhong JH, Tan J, Liang GZ (2012) Influence of different land use types on the soil fauna diversity. Ecology and Environmental Sciences, 21, 1678-1682. (in Chinese with English abstract)
	[林兰稳, 钟继洪, 谭军, 梁广灶 (2012) 土地利用方式对土壤动物多样性的影响. 生态环境学报, 21, 1678-1682.] DOI
[19]	Liu RT, Zhu F, Zhao HL (2013) Effects of land use/cover changes on soil macro-faunal community in agro-pastoral transitional zone, Northern China. Acta Agrestia Sinica, 21, 643-649. (in Chinese with English abstract)
	[刘任涛, 朱凡, 赵哈林 (2013) 北方农牧交错区土地利用覆盖变化对大型土壤动物群落结构的影响. 草地学报, 21, 643-649.] DOI
[20]	Luo DH, Li X, Luo BJ, Xu MS, Tuo B, Yan ER, You WH (2020) Soil meso- and micro-fauna characteristics in evergreen broad-leaved forest and deciduous broad-leaved forest in Dajinshan Island, China. Journal of Ecology and Rural Environment, 36, 349-357. (in Chinese with English abstract)
	[罗鼎晖, 李翔, 骆蓓菁, 许洺山, 妥彬, 阎恩荣, 由文辉 (2020) 大金山岛常绿阔叶林和落叶阔叶林中小型土壤动物群落特征. 生态与农村环境学报, 36, 349-357.]
[21]	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.]
[22]	Margalef DR (1958) Information theory in ecology. General Systematics, 3, 36-71.
[23]	Nedved O (1998) Modelling the relationship between cold injury and accumulated degree days in terrestrial arthropods. Cryo Letters, 19, 267-274.
[24]	Niu YJ, Zhou JW, Yang SW, Wang GZ, Liu L, Du GZ, Hua LM (2017) Relationships between soil moisture and temperature, plant species diversity, and primary productivity in an alpine meadow considering topographic factors. Acta Ecologica Sinica, 37, 8314-8325. (in Chinese with English abstract)
	[牛钰杰, 周建伟, 杨思维, 王贵珍, 刘丽, 杜国祯, 花立民 (2017) 基于地形因素的高寒草甸土壤温湿度和物种多样性与初级生产力关系研究. 生态学报, 37, 8314-8325.]
[25]	Pielou EC (1975) Ecological Diversity. John Wiley & Sons, New Jersey.
[26]	Qian FS, Wang ZY (1995) Relationship between soil fauna and soil environment in jujube garden of Shuidong, Anhui Province. Chinese Journal of Applied Ecology, 6, 44-50. (in Chinese with English abstract)
	[钱复生, 王宗英 (1995) 水东枣园土壤动物与土壤环境的关系. 应用生态学报, 6, 44-50.]
[27]	Sileshi G, Mafongoya PL (2006) Variation in macrofaunal communities under contrasting land use systems in eastern Zambia. Applied Soil Ecology, 33, 49-60.
[28]	Stiegel S, Entling MH, Mantilla-Contreras J (2017) Reading the leaves’ palm: Leaf traits and herbivory along the microclimatic gradient of forest layers. PLoS ONE, 12, e0169741.
[29]	Sun X, Li Q, Yao HF, Liu MQ, Wu DH, Zhu D, Zhu YG (2021) Soil fauna and soil health. Acta Pedologica Sinica, 58, 1073-1083. (in Chinese with English abstract)
	[孙新, 李琪, 姚海凤, 刘满强, 吴东辉, 朱冬, 朱永官 (2021) 土壤动物与土壤健康. 土壤学报, 58, 1073-1083.]
[30]	Tan B, Yin R, Zhang J, Xu ZF, Liu Y, He SQ, Zhang L, Li H, Wang LX, Liu SN, You CM, Peng CH (2021) Temperature and moisture modulate the contribution of soil fauna to litter decomposition via different pathways. Ecosystems, 24, 1142-1156. DOI
[31]	Torstensdson L, Pell M, Stenberg B (1998) Soil quality assessment of cultivated land requires a strategy. Ambio, 27, 28-30.
[32]	Wang CE, Huang M, Wang WY, Li ZH, Zhang T, Ma L, Bai YF, Wang YL, Shi JJ, Long RJ, Liu Y, Wang XL, Ma YS, Shang ZH (2022) Variation characteristics of plant community diversity and above-ground biomass in alpine degraded slopes along altitude gradients in the headwaters region of three-river on Tibetan Plateau. Acta Ecologica Sinica, 42, 3640-3655. (in Chinese with English abstract)
	[王采娥, 黄梅, 王文银, 李子好, 张涛, 马林, 白彦福, 王彦龙, 施建军, 龙瑞军, 刘玉, 王晓丽, 马玉寿, 尚占环 (2022) 三江源区高寒坡地退化植物群落多样性和地上生物量沿海拔梯度的变化特征. 生态学报, 42, 3640-3655.]
[33]	Wang HY, Zhu YH (2022) The study of the characteristics of soil fauna community and soil humus characteristics at different elevations in Lushan. Hubei Agricultural Sciences, 61(7), 25-30. (in Chinese with English abstract)
	[王海燕, 朱永恒 (2022) 庐山不同海拔土壤动物群落特征与土壤腐殖质特性研究. 湖北农业科学, 61(7), 25-30.]
[34]	Wang XF, Mi XC, Wang XH, Jiang MX, Yang T, Zhang J, Shen ZH (2023) Comparison of woody plant diversity in the evergreen broad-leaved forest in mid-subtropical China. Biodiversity Science, 31, 23296. (in Chinese with English abstract) DOI
	[王晓凤, 米湘成, 王希华, 江明喜, 杨涛, 张健, 沈泽昊 (2023) 中国中亚热带常绿阔叶林群落木本植物多样性比较. 生物多样性, 31, 23296.] DOI
[35]	Wang XG, Hao ZQ, Ye J, Zhang J, Li BH, Yao XL (2008) Spatial variation of species diversity across scales in an old-growth temperate forest of China. Ecological Research, 23, 709-717.
[36]	Whittaker RH (1972) Evolution and measurement of species diversity. Taxon, 21, 213-251.
[37]	Wu Q, Wu HT, Sun X, Lin YL, Liu DD, Kang YJ, Liu JP (2023) Community composition and surface aggregation characteristics of soil Collembola in Changbai Mountain Tundra. Environmental Ecology, 5, 39-46. (in Chinese with English abstract)
	[吴琼, 武海涛, 孙新, 林奕伶, 刘丹丹, 康玉娟, 刘吉平 (2023) 长白山苔原带土壤跳虫群落组成和表聚性特征研究. 环境生态学, 5, 39-46.]
[38]	Wu TH, Ayres E, Bardgett RD, Wall DH, Garey JR (2011) Molecular study of worldwide distribution and diversity of soil animals. Proceedings of the National Academy of Sciences, USA, 108, 17720-17725.
[39]	Xiao NW, Liu XH, Ge F, Ouyang ZY (2009) Research on soil faunal community composition and structure in the Gaoligong Mountains National Nature Reserve. Acta Ecologica Sinica, 29, 3576-3584. (in Chinese with English abstract)
	[肖能文, 刘向辉, 戈峰, 欧阳志云 (2009) 高黎贡山自然保护区大型土壤动物群落特征. 生态学报, 29, 3576-3584.]
[40]	Yan HB, Han YZ, Yang XQ, Wang LY, Xiang XY (2010) Spatial distribution patterns and associations of tree species in typical natural secondary mountain forest communities of Northern China. Acta Ecologica Sinica, 30, 2311-2321. (in Chinese with English abstract)
	[闫海冰, 韩有志, 杨秀清, 王丽艳, 项小英 (2010) 华北山地典型天然次生林群落的树种空间分布格局及其关联性. 生态学报, 30, 2311-2321.]
[41]	Yan JC, Cui D, Jiang ZC, Lü LQ, Liu X, Wu YX, Li JG (2023) Research progress of soil fauna in arid and semi-arid farmland ecosystems in China. Journal of Xinjiang Normal University (Natural Sciences Edition), 42, 34-41. (in Chinese with English abstract)
	[闫江超, 崔东, 江智诚, 吕立琴, 刘璇, 武玉鑫, 李金哥 (2023) 干旱半干旱地区农田生态系统中土壤动物研究进展. 新疆师范大学学报(自然科学版), 42, 34-41.]
[42]	Ye Y, Jiang YX, Chen H (2019) Responses of functional groups of large soil fauna to niche environmental factors. Jiangsu Agricultural Sciences, 47, 253-257. (in Chinese with English abstract)
	[叶岳, 姜玉霞, 陈华 (2019) 大型土壤动物功能类群对小生境环境因子的响应. 江苏农业科学, 47, 253-257.]
[43]	Yin WY (1992) Soil Fauna in the Subtropical of China. Science Press, Beijing. (in Chinese)
	[尹文英 (1992) 中国亚热带土壤动物. 科学出版社, 北京.]
[44]	Yin WY (1998) Pictorial Keys to Soil Fauna of China. Science Press, Beijing. (in Chinese)
	[尹文英 (1998) 中国土壤动物检索图鉴. 科学出版社, 北京.]
[45]	Yin XQ, Cui D, Xu H (2019) Characteristics of soil faunal communities under different disturbance levels in the forests of Changbai Mountains. Journal of Northeast Normal University (Natural Science Edition), 51(4), 118-123. (in Chinese with English abstract)
	[殷秀琴, 崔东, 许还 (2019) 长白山地森林不同干扰程度土壤动物的群落特征. 东北师大学报(自然科学版), 51(4), 118-123.]
[46]	Yin XQ, Zhang GR (1993) Correlation between forest litter and soil macroanimals. Chinese Journal of Applied Ecology, 4, 167-173. (in Chinese with English abstract)
	[殷秀琴, 张桂荣 (1993) 森林凋落物与大型土壤动物相关关系的研究. 应用生态学报, 4, 167-173.]
[47]	Zang JC, Song MC, Huang Z, Zhang PP (2023) Spatial and temporal distribution of soil animal community diversity at different altitudes in Sejila Mountain (sunny slope), Tibet. Journal of China Agricultural University, 28(5), 189-199. (in Chinese with English abstract)
	[臧建成, 宋美成, 黄智, 张盼盼 (2023) 西藏色季拉山(阳坡)不同海拔土壤动物群落多样性时空分布特征. 中国农业大学学报, 28(5), 189-199.]
[48]	Zhang WR, Ye B, Li YK, Yuan KN, Shi PQ, Xu BT, Zhang GZ, Zhou FD (1987) Methods for Analyzing Forest Soils. Standards Press of China, Beijing. (in Chinese)
	[张万儒, 叶炳, 李酉开, 袁可能, 施培青, 许本彤, 张国珠, 周斐德 (1987) 森林土壤分析方法. 中国标准出版社, 北京.]
[49]	Zhao SK, Guo JP, Zhang YX (2023) Responses of root respiration to woodland warming during growing season in succession series of natural secondary forest in warm-temperate mountain. Scientia Silvae Sinicae, 59(2), 10-21. (in Chinese with English abstract)
	[赵世魁, 郭晋平, 张芸香 (2023) 暖温带山地天然次生林演替序列根系呼吸速率对林地增温的响应. 林业科学, 59(2), 10-21.]
[50]	Zhong JM (1990) Taxonomy of Larvae. China Agriculture Press, Beijing. (in Chinese)
	[钟觉民 (1990) 幼虫分类学. 中国农业出版社, 北京.]

海拔 Elevation (m)	纬度 Latitude (N)	经度 Longitude (E)	坡度 Slope	坡向 Slope aspect	郁闭度 Canopy density	凋落物厚度 Litter thickness (cm)	胸径 DBH (cm)	平均树高 Mean tree height (m)
1,600	37°49′	111°29′	19°	半阳 Semi-sunny	0.78	6.3	17.5 ± 0.7	13.4 ± 1.6
1,800	37°49′	111°29′	21°	半阳 Semi-sunny	0.75	5.9	16.7 ± 2.7	12.9 ± 3.9
2,000	37°52′	111°26′	18°	半阳 Semi-sunny	0.76	6.5	17.2 ± 1.2	12.6 ± 1.5
2,200	37°53′	111°25′	20°	半阳 Semi-sunny	0.73	6.1	15.6 ± 2.5	11.9 ± 2.2

海拔 Elevation (m)	纬度 Latitude (N)	经度 Longitude (E)	坡度 Slope	坡向 Slope aspect	郁闭度 Canopy density	凋落物厚度 Litter thickness (cm)	胸径 DBH (cm)	平均树高 Mean tree height (m)
1,600	37°49′	111°29′	19°	半阳 Semi-sunny	0.78	6.3	17.5 ± 0.7	13.4 ± 1.6
1,800	37°49′	111°29′	21°	半阳 Semi-sunny	0.75	5.9	16.7 ± 2.7	12.9 ± 3.9
2,000	37°52′	111°26′	18°	半阳 Semi-sunny	0.76	6.5	17.2 ± 1.2	12.6 ± 1.5
2,200	37°53′	111°25′	20°	半阳 Semi-sunny	0.73	6.1	15.6 ± 2.5	11.9 ± 2.2

海拔 Elevation (m)	春季 Spring (5月 May)		夏季 Summer (7月 July)		秋季 Autumn (10月 October)
海拔 Elevation (m)	物种数 Ns (%)	个体数 Ni (%)	物种数 Ns (%)	个体数 Ni (%)	物种数 Ns (%)	个体数 Ni (%)
1,600	78 (75.00%)	5,004 (36.94%)	87 (76.99%)	2,998 (35.20%)	64 (69.57%)	746 (31.45%)
1,800	79 (75.96%)	3,863 (28.52%)	87 (76.99%)	2,205 (25.89%)	72 (78.26%)	706 (29.76%)
2,000	70 (67.31%)	2,777 (20.50%)	83 (73.45%)	1,915 (22.48%)	53 (57.61%)	542 (22.85%)
2,200	67 (64.42%)	1,901 (14.03%)	75 (66.37%)	1,399 (16.43%)	51 (55.43%)	378 (15.94%)
总计 Total	104	13,545	113	8,517	92	2,372

海拔 Elevation (m)	春季 Spring (5月 May)		夏季 Summer (7月 July)		秋季 Autumn (10月 October)
海拔 Elevation (m)	物种数 Ns (%)	个体数 Ni (%)	物种数 Ns (%)	个体数 Ni (%)	物种数 Ns (%)	个体数 Ni (%)
1,600	78 (75.00%)	5,004 (36.94%)	87 (76.99%)	2,998 (35.20%)	64 (69.57%)	746 (31.45%)
1,800	79 (75.96%)	3,863 (28.52%)	87 (76.99%)	2,205 (25.89%)	72 (78.26%)	706 (29.76%)
2,000	70 (67.31%)	2,777 (20.50%)	83 (73.45%)	1,915 (22.48%)	53 (57.61%)	542 (22.85%)
2,200	67 (64.42%)	1,901 (14.03%)	75 (66.37%)	1,399 (16.43%)	51 (55.43%)	378 (15.94%)
总计 Total	104	13,545	113	8,517	92	2,372

指数 Index	海拔 Elevation (m)	枯枝落叶层 Litter layer	0-10 cm	10-20 cm	20-30 cm	F	P
Shannon-Wiener多样性指数 Shannon-Wiener diversity index (H')	1,600	4.34 ± 0.15^Ba	4.15 ± 0.10^Ba	4.10 ± 0.24^Aa	3.91 ± 0.25^Aa	1.56	0.273
	1,800	4.53 ± 0.12^Aa	4.31 ± 0.14^Bab	4.20 ± 0.03^Ac	4.00 ± 0.27^Ac	5.53	0.024^*
	2,000	4.24 ± 0.15^ABa	4.21 ± 0.16^Ba	4.19 ± 0.21^Aa	4.11 ± 0.21^Aa	0.90	0.482
	2,200	4.52 ± 0.06^Aa	4.45 ± 0.02^Aa	4.36 ± 0.16^Aa	3.92 ± 0.14^Ab	20.81	0.00039^**
Margalef丰富度指数 Margalef richness index (D)	1,600	6.23 ± 0.19^Ba	5.00 ± 0.31^ABb	4.30 ± 0.27^Abc	3.53 ± 0.70^Ac	20.72	0.000396^**
	1,800	6.93 ± 0.45^Aa	5.41 ± 0.12^Ab	4.57 ± 0.46^Ac	3.72 ± 0.10^Ad	51.19	0.000014
	2,000	6.26 ± 0.34^ABa	4.71 ± 0.26^Bb	4.04 ± 0.21^Abc	3.56 ± 0.40^Ac	42.44	0.000029^**
	2,200	6.13 ± 0.47^ABa	4.36 ± 0.15^Ab	4.03 ± 0.09^Ac	3.41 ± 0.24^Ad	59.34	0.000008^**
Pielou均匀度指数 Pielou evenness index (E)	1,600	0.84 ± 0.01^Ba	0.79 ± 0.03^Cab	0.74 ± 0.03^Cbc	0.70 ± 0.03^Ac	17.06	0.001^**
	1,800	0.87 ± 0.05^ABa	0.81 ± 0.02^BCb	0.76 ± 0.02^BCbc	0.73 ± 0.01^ABc	10.61	0.004^**
	2,000	0.91 ± 0.01^Aa	0.85 ± 0.03^ABb	0.80 ± 0.04^ABb	0.72 ± 0.02^Bc	25.72	0.000184^**
	2,200	0.90 ± 0.01^Aa	0.88 ± 0.03^Aa	0.83 ± 0.01^Ab	0.76 ± 0.01^Ac	36.69	0.00005^**
Simpson优势度指数 Simpson dominance index (C)	1,600	0.08 ± 0.01^ABa	0.08 ± 0.01^Aa	0.09 ± 0.02^Aa	0.09 ± 0.02^Aa	0.45	0.725
	1,800	0.07 ± 0.00^Ba	0.08 ± 0.01^Aa	0.08 ± 0.03^Aa	0.08 ± 0.01^Aa	0.39	0.764
	2,000	0.06 ± 0.01^Aa	0.08 ± 0.02^Aa	0.08 ± 0.01^Aa	0.09 ± 0.01^Ba	2.89	0.103
	2,200	0.06 ± 0.02^Ba	0.06 ± 0.00^Aa	0.07 ± 0.02^Aa	0.08 ± 0.00^ABa	1.45	0.298