Plant beta diversity and its influence factors in the Liangshui National Nature Reserve in the central region of the Xiaoxing’an Mountains

doi:10.17520/biods.2021274

Abstract

Abstract:

Aims In biological conservation sciences, implementing nature reserves to protect biological diversity is a hot topic. Specifying plant beta diversity, structural components, and influence factors is the critical basis for creating nature reserves.

Methods In this paper, a national nature reserve named Liangshui, located in the central region of the Xiaoxing’an Mountains, was used to survey 80 quadrats in the size of 20 m × 20 m in the core zone, buffer zone, experimental zone, and outside the reserve. We recorded topographic information (Longitude and latitude, altitude, slope, slope aspect, and slope position), community structure (canopy density, tree age, tree height, tree diameter at breast height (DBH), shrub height, and shrub ground diameter), and species name for each tree, shrub, and herb. We also sampled 0-20 cm soils for soil property determination (soil organic carbon, total nitrogen, pH, electroconductibility, soil water content, and bulk density). Abundance data was used for beta diversity calculation as well as computing species abundance differences and species turnover. The Mantel test, redundancy ordination, and variation partitioning were used to examine the effects of abiotic factors (geography and terrain, protection intensity, soil matrix) and biological factors (community structure) on beta diversity and its partitioning components.

Results (1) Species turnover contributed most to the overall beta diversity (Sørensen dissimilarity) for all layers with a range from 65% (for shrub) to 73% (for herb). (2) Mantel correlation analysis indicated that beta diversity, their two components of abundance differences and species turnover, had close associations with the geography and terrain in the arbor, shrub, and herb layer. In contrast, soil properties expressed more association with beta diversity from the arbor and shrub layer. The slope position, slope degree, tree height, and protection intensity were significantly associated with beta diversity in 3 layers (P< 0.05). (3) Variation partitioning and redundancy analysis ordination indicated that geography and terrain affected the beta diversity of whole forests of the reserve significantly, but differences existed between arbor, shrub and herb layers. The arbor beta diversity was most influenced by biological factors. In contrast, the shrub layer was mainly affected by soil factors, twice more than the geography and terrain and biological factors. In the herb layer, the geography and terrain contributed the most to variation, at a rate of respectively 26-fold and 3-fold higher than that from soil factors and biological factors. Tree DBH had the most significant explanatory effect on the beta diversity variations in the reserve.

Conclusion Our findings highlights that nature reserve settlements in the future should consider combining assemblage of the proper community structure of the arbor, shrub, and herb layers, and soil factors, rather than only emphasis on protection intensity. Such steps will improve the conservation efficiency for plant diversity protection.

Key words: beta diversity, species turnover, species abundance difference, protection intensity, Northeast China forest belt

Danqi She, Xiting Zhang, Lu Xiao, Zhaoliang Zhong, Huimei Wang, Wenjie Wang. Plant beta diversity and its influence factors in the Liangshui National Nature Reserve in the central region of the Xiaoxing’an Mountains[J]. Biodiv Sci, 2022, 30(3): 21274.

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URL: https://www.biodiversity-science.net/EN/10.17520/biods.2021274

https://www.biodiversity-science.net/EN/Y2022/V30/I3/21274

Figures/Tables 5

Fig. 1 The relationship of taxonomic and functional trait beta diversity and its components (turnover and nestedness) of macrobenthic communities with difference in geographic distance

Fig. 2 Triangle figures for arbor (a), shrub (b), and herb (c) of plant beta diversity (Btotal), and its components of species turnover (Repl) and abundance differences (AbDiff). Each black dot represents a pair of sites. Their positions were determined by a triplet of values from the species composition similarity (S = 1 - Btotal), Repl, and AbDiff; each triplet sums to 1. The larger grey dots represent the mean values.

Table 1 Mantel correlations analysis between beta diversity (Btotal), its components of species turnover (Repl) and abundance differences (AbDiff) and geography and terrain, protection intensity, soil factors and biological factors

		乔木层 Arbor layer			灌木层 Shrub layer			草本层 Herb layer
		Sørensen index	Ab_Diff	R_epl	Sørensen index	Ab_Diff	R_epl	Sørensen index	Ab_Diff	R_epl
地理地形 Geography and terrain	经纬度 Longitude and latitude	0.079^**	0.01	0.06	0.05	0.02	0.02	0.06	-0.06	0.1^**
	海拔 Altitude	0.21^***	0.21^*	0.01	0.07	0.07	<0.001	0.18^***	0.06	0.07
	坡度 Slope	0.2^***	-0.01	0.174^**	0.18^**	0.03	0.12^*	0.3^***	0.05	0.16^***
	坡位 Slope position	0.35^***	0.14^***	0.182^***	0.18^***	0.09^*	0.08^**	0.13^***	0.05	0.05*
	坡向 Aspect	-0.08	0.02	-0.08	-0.07	-0.08	0.01	-0.01	-0.04	0.03
保护强度 Protection intensity		0.14^***	0.12^***	0.02	0.1^**	0.06	0.03	0.08^**	-0.01	0.07^**
土壤因子 Soil factors	有机碳 Soil organic carbon	0.03	-0.04	0.06	0.05	0.09	-0.03	-0.01	-0.01	<0.001
	全氮 Total nitrogen	0.08	0.04	0.04	0.16^**	0.06	0.08	0.06	0.02	0.03
	C : N	-0.06	0.004	-0.05	0.01	-0.06	0.06	-0.02	<0.001	-0.01
	pH	0.05	0.13^*	-0.06	0.09	0.03	0.05	-0.04	<0.001	-0.03
	电导率 Electroconductibility	0.05	-0.08	0.02	-0.003	0.11	-0.09	-0.01	-0.01	<0.001
	容重 Soil bulk density	0.04	0.04	0.003	0.13^*	0.11	0.02	-0.003	-0.01	0.01
	含水量 Soil water content	0.09^*	0.07	0.16	0.21^**	-0.01	0.18^*	0.07	0.02	0.03
生物因子 Biological factors	郁闭度 Canopy density	0.06^*	0.01	0.05	0.008	-0.04	0.04	0.06	0.03	0.01
	林龄 Forest age	0.28^***	0.42^***	-0.1	0.08	0.07	0.004	0.08	0.02	0.03
	乔木树高 Tree height	0.24^***	-0.01	0.21^***	0.14^**	0.07	0.05	0.11^*	-0.06	0.13^***
	乔木胸径 Tree DBH	0.18^***	0.45^***	-0.21	0.05	0.02	0.03	0.09^*	0.03	0.03
	灌木树高 Shrub height	0.05	0.04	-0.01	0.12^*	0.14^*	-0.02	-0.03	0.01	-0.03
	灌木地径 Shrub ground diameter	0.02	-0.02	0.03	0.16^**	0.17^*	0.01	-0.08	<0.001	-0.06

		乔木层 Arbor layer			灌木层 Shrub layer			草本层 Herb layer
		Sørensen index	Ab_Diff	R_epl	Sørensen index	Ab_Diff	R_epl	Sørensen index	Ab_Diff	R_epl
地理地形 Geography and terrain	经纬度 Longitude and latitude	0.079^**	0.01	0.06	0.05	0.02	0.02	0.06	-0.06	0.1^**
	海拔 Altitude	0.21^***	0.21^*	0.01	0.07	0.07	<0.001	0.18^***	0.06	0.07
	坡度 Slope	0.2^***	-0.01	0.174^**	0.18^**	0.03	0.12^*	0.3^***	0.05	0.16^***
	坡位 Slope position	0.35^***	0.14^***	0.182^***	0.18^***	0.09^*	0.08^**	0.13^***	0.05	0.05*
	坡向 Aspect	-0.08	0.02	-0.08	-0.07	-0.08	0.01	-0.01	-0.04	0.03
保护强度 Protection intensity		0.14^***	0.12^***	0.02	0.1^**	0.06	0.03	0.08^**	-0.01	0.07^**
土壤因子 Soil factors	有机碳 Soil organic carbon	0.03	-0.04	0.06	0.05	0.09	-0.03	-0.01	-0.01	<0.001
	全氮 Total nitrogen	0.08	0.04	0.04	0.16^**	0.06	0.08	0.06	0.02	0.03
	C : N	-0.06	0.004	-0.05	0.01	-0.06	0.06	-0.02	<0.001	-0.01
	pH	0.05	0.13^*	-0.06	0.09	0.03	0.05	-0.04	<0.001	-0.03
	电导率 Electroconductibility	0.05	-0.08	0.02	-0.003	0.11	-0.09	-0.01	-0.01	<0.001
	容重 Soil bulk density	0.04	0.04	0.003	0.13^*	0.11	0.02	-0.003	-0.01	0.01
	含水量 Soil water content	0.09^*	0.07	0.16	0.21^**	-0.01	0.18^*	0.07	0.02	0.03
生物因子 Biological factors	郁闭度 Canopy density	0.06^*	0.01	0.05	0.008	-0.04	0.04	0.06	0.03	0.01
	林龄 Forest age	0.28^***	0.42^***	-0.1	0.08	0.07	0.004	0.08	0.02	0.03
	乔木树高 Tree height	0.24^***	-0.01	0.21^***	0.14^**	0.07	0.05	0.11^*	-0.06	0.13^***
	乔木胸径 Tree DBH	0.18^***	0.45^***	-0.21	0.05	0.02	0.03	0.09^*	0.03	0.03
	灌木树高 Shrub height	0.05	0.04	-0.01	0.12^*	0.14^*	-0.02	-0.03	0.01	-0.03
	灌木地径 Shrub ground diameter	0.02	-0.02	0.03	0.16^**	0.17^*	0.01	-0.08	<0.001	-0.06

Fig. 3 Using variation partitioning to analysis the effect of geography and terrain, protection intensity, community characteristics and soil factor on beta diversity characteristics. (a) Arbor-shrub-herb; (b) Arbor; (c) Shrub; (d) Herb.

Fig. 4 Redundancy analysis (RDA) ordination between geography and terrain, protection intensity, soil matrix, and community characteristics and beta diversity indices (left) and the most significant explaining factors (right). Btotal, Beta diversity; AbDiff, component of abundance differences; Repl, component of species turnover; T-D, Arbor Btotal; T-Ab, Arbor AbDiff; T-Rep, Arbor Repl; S-D, Shrub Btotal; S-Ab, Shrub AbDiff; S-Rep, Shrub Repl; H-D, Herb Btotal; H-Ab, Herb AbDiff; H-Rep, Herb Repl; Lon. And Lat., Longitude and Latitude; SP, Slope position; PI, Protection intensity; CD, Crown density; FA, Forest age; TDBH, Tree diameter; TH, Tree height; SH, Shrub height; SGD, Shrub ground diameter; SOC, Soil organic carbon; TN, Total nitrogen; CN, C : N; EC, Electroconductibility; BD, Soil bulk density; SWC, Soil water content.

References 42

[1]	Bao SD (2000) Soil Agro-chemistry Analysis, 3rd edn. China Agriculture Press, Beijing. (in Chinese)
	[ 鲍士旦 (2000) 土壤农化分析(第3版). 中国农业出版社, 北京.]
[2]	Carvalho JC, Cardoso P, Gomes P (2012) Determining the relative roles of species replacement and species richness differences in generating beta-diversity patterns. Global Ecology and Biogeography, 21, 760-771. DOI URL
[3]	Chen C, Wang Y, Wang JM, Yang H, Wang YC, Xu C, Li JW, Chu JM (2020) Species diversity of plant communities and its main influencing factors in Horqin Sandy Land, Inner Mongolia of northern China. Journal of Beijing Forestry University, 42(5), 106-114. (in Chinese with English abstract)
	[ 陈晨, 王寅, 王健铭, 杨欢, 王雨辰, 徐超, 李景文, 褚建民 (2020) 科尔沁沙地植物群落物种多样性及其主要影响因素. 北京林业大学学报, 42(5), 106-114.]
[4]	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) DOI
	[ 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.] DOI
[5]	Chen XB, Zhu DQ, Zhao CC, Zhang LL, Chen LX, Duan WB (2019) Community composition and diversity of fungi in soils under different types of Pinus koraiensis forests. Acta Pedologica Sinica, 56, 1221-1234. (in Chinese with English abstract)
	[ 陈秀波, 朱德全, 赵晨晨, 张路路, 陈立新, 段文标 (2019) 不同林型红松林土壤真菌群落组成和多样性. 土壤学报, 56, 1221-1234.]
[6]	Condit R, Pitman N, Leigh EG Jr, Chave J, Terborgh J, Foster RB, Núñez P, Aguilar S, Valencia R, Villa G, Muller-Landau HC, Losos E, Hubbell SP (2002) Beta-diversity in tropical forest trees. Science, 295, 666-669. DOI URL
[7]	Dray S, Blanchet FG, Legendre P, Madi N, Wagner H (2017) adespatial: Multivariate Multiscale Spatital Analysis. R package version 0.2-0. https://cran.r-project.org/web/packages/adespatital/ . (accessed on 2018-05-23)
[8]	Fernandez-Going BM, Harrison SP, Anacker BL, Safford HD (2013) Climate interacts with soil to produce beta diversity in Californian plant communities. Ecology, 94, 2007-2018. PMID
[9]	Gutiérrez-Cánovas C, Millán A, Velasco J, Vaughan IP, Ormerod SJ (2013) Contrasting effects of natural and anthropogenic stressors on beta diversity in river organisms. Global Ecology and Biogeography, 22, 796-805. DOI URL
[10]	Hájek M, Hájková P, Kočí M, Jiroušek M, Mikulášková E, Kintrová K (2012) Do we need soil moisture measurements in the vegetation-environment studies in wetlands? Journal of Vegetation Science, 24, 127-137. DOI URL
[11]	Harrison S, Ross SJ, Lawton JH (1992) Beta diversity on geographic gradients in Britain. Journal of Animal Ecology, 61, 151-158. DOI URL
[12]	Hubbell SP (2001) The Unified Neutral Theory of Biodiversity and Biogeography. Princeton University Press, Princeton.
[13]	Kong LW, Chen XW, Lu SW, Li SN, Chen B, Gao C, Shi Y, Yang XY (2014) Relationships among growth of Larix principis-rupprechtii, herbaceous plants diversity and landform. Bulletin of Soil and Water Conservation, 34(5), 60-66. (in Chinese with English abstract)
	[ 孔令伟, 陈祥伟, 鲁绍伟, 李少宁, 陈波, 高琛, 石媛, 杨小燕 (2014) 华北落叶松林木生长、草本植物多样性及地形因子之间的关系. 水土保持通报, 34(5), 60-66.]
[14]	Kraft NJB, Valencia R, Ackerly DD (2008) Functional traits and niche-based tree community assembly in an Amazonian forest. Science, 322, 580-582. DOI URL
[15]	Legendre P (2014) Interpreting the replacement and richness difference components of beta diversity. Global Ecology and Biogeography, 23, 1324-1334. DOI URL
[16]	Lennon JJ, Koleff P, GreenwooD JJD, Gaston KJ (2001) The geographical structure of British bird distributions: Diversity, spatial turnover and scale. Journal of Animal Ecology, 70, 966-979. DOI URL
[17]	Leprieur F, Tedesco PA, Hugueny B, Beauchard O, Dürr HH, Brosse S, Oberdorff T (2011) Partitioning global patterns of freshwater fish beta diversity reveals contrasting signatures of past climate changes. Ecology Letters, 14, 325-334. DOI PMID
[18]	Li XH, Liu YH, Liu Y, Xu Y, Yang Y, Shen ZH (2016) Impacts of geographical distances and environmental differences on the beta diversity of plant communities in the dry-hot valley of the Yuanjiang River. Biodiversity Science, 24, 399-406. (in Chinese with English abstract) DOI URL
	[ 李新辉, 刘延虹, 刘晔, 许玥, 杨阳, 沈泽昊 (2016) 地理距离及环境差异对云南元江干热河谷植物群落beta多样性的影响. 生物多样性, 24, 399-406.] DOI
[19]	Liu HW, Chen LX, Ma HJ, Qiao L, Zhang J (2010) Seasonal dynamics of soil phosphorus for main forest types in Liangshui Nature Reserve and its availability. Journal of Northeast Forestry University, 38(4), 62-65. (in Chinese with English abstract)
	[ 刘宏伟, 陈立新, 马海娟, 乔璐, 张杰 (2010) 凉水自然保护区主要森林类型土壤磷季节动态及有效性. 东北林业大学学报, 38(4), 62-65.]
[20]	Liu SC, Duan WB, Feng J, Han SZ (2011) Effects of forest gap on tree species regeneration and diversity of mixed broadleaved Korean pine forest in Xiaoxing’an Mountains. Chinese Journal of Applied Ecology, 22, 1381-1388. (in Chinese with English abstract)
	[ 刘少冲, 段文标, 冯静, 韩生忠 (2011) 林隙对小兴安岭阔叶红松林树种更新及物种多样性的影响. 应用生态学报, 22, 1381-1388.]
[21]	Lu ZS, Guo JP, Hu SC (1993) Use the growth cone-like wood method to determine the number of stands at maturity. East China Forest Management, 7(2), 29-33. (in Chinese)
	[ 陆兆苏, 郭晋平, 胡世才 (1993) 用生长锥样木法确定林分数量成熟龄. 华东森林经理, 7(2), 29-33.]
[22]	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 URL
[23]	Si XF, Zhao YH, Chen CW, Ren P, Zeng D, Wu LB, Ding P (2017) Beta-diversity partitioning: Methods, applications and perspectives. Biodiversity Science, 25, 464-480. (in Chinese with English abstract) DOI URL
	[ 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.] DOI
[24]	Socolar JB, Gilroy JJ, Kunin WE, Edwards DP (2016) How should beta-diversity inform biodiversity conservation? Trends in Ecology and Evolution, 31, 67-80. DOI PMID
[25]	Soininen J, Heino J, Wang JJ (2018) A meta-analysis of nestedness and turnover components of beta diversity across organisms and ecosystems. Global Ecology and Biogeography, 27, 96-109. DOI URL
[26]	Tan SS, Ye ZL, Yuan LB, Zhou RF, Hu G, Jin XF, Yu MJ (2013) Beta diversity of plant communities in Baishanzu Nature Reserve. Acta Ecologica Sinica, 33, 6944-6956. (in Chinese with English abstract) DOI URL
	[ 谭珊珊, 叶珍林, 袁留斌, 周荣飞, 胡广, 金孝锋, 于明坚 (2013) 百山祖自然保护区植物群落beta多样性. 生态学报, 33, 6944-6956.]
[27]	Tang ZY, Fang JY, Zhang L (2004) Patterns of woody plant species diversity along environmental gradients on Mt. Taibai, Qinling Mountains. Biodiversity Science, 12, 115-122. (in Chinese with English abstract) DOI URL
	[ 唐志尧, 方精云, 张玲 (2004) 秦岭太白山木本植物物种多样性的梯度格局及环境解释. 生物多样性, 12, 115-122.] DOI
[28]	Tisseuil C, Leprieur F, Grenouillet G, Vrac M, Lek S (2012) Projected impacts of climate change on spatio-temporal patterns of freshwater fish beta diversity: A deconstructing approach. Global Ecology and Biogeography, 21, 1213-1222. DOI URL
[29]	Tuomisto H, Ruokolainen K, Yli-Halla M (2003) Dispersal, environment, and floristic variation of western Amazonian forests. Science, 299, 241-244. PMID
[30]	Wang SX, Xia TT, Wang XA (2021) Understory β diversity differences and influential factors between Pinus tabuliformis plantation and natural Quercus wutaishanica forest on the Loess Plateau. Guilhaia, 41, 362-371. (in Chinese with English abstract)
	[ 王世雄, 夏婷婷, 王孝安 (2021) 黄土高原油松和辽东栎林下植物β多样性差异及影响因素. 广西植物, 41, 362-371.]
[31]	Wang WJ, Du HJ, Xiao L, Zhang JY, Zhong ZL, Zhou W, Zhang B, Wang HY (2019) Differences in plant composition and forest structure among of 3 forest types in Liangshui National Nature Reserve. Scientia Silvae Sinicae, 55, 166-176. (in Chinese with English abstract)
	[ 王文杰, 杜红居, 肖路, 张建宇, 仲召亮, 周伟, 张波, 王洪元 (2019) 凉水自然保护区3种森林类型的植物组成和林分结构特征. 林业科学, 55, 166-176.]
[32]	Wang WJ, Sun JX, Zhong ZL, Xiao L, Wang YY, Wang HM (2021) Relating macrofungal diversity and forest characteristics in boreal forests in China: Conservation effects, inter-forest-type variations, and association decoupling. Ecology and Evolution, 11, 13268-13282. DOI URL
[33]	Wang XG, Wiegand T, Anderson-Teixeira KJ, Bourg NA, Myers JA (2018) Ecological drivers of spatial community dissimilarity, species replacement and species nestedness across temperate forests. Global Ecology and Biogeography, 27, 581-592. DOI URL
[34]	Whittaker RH (1972) Evolution and measurement of species diversity. Taxon, 21, 213-251. DOI URL
[35]	Williams PH (1996) Mapping variations in the strength and breadth of biogeographic transition zones using species turnover. Proceedings of the Royal Society B: Biological Sciences, 263, 579-588.
[36]	Yao XY, Hu YS, Liu YH (2014) Functional diversity of typical broad-leaved Korean pine forest communities in Changbai Mountains and its relationship with topographical factors. Journal of Northwest A&F University (Natural Science Edition), 42(10), 95-102. (in Chinese with English abstract)
	[ 么旭阳, 胡耀升, 刘艳红 (2014) 长白山阔叶红松林典型森林群落功能多样性及其与地形因子的关系. 西北农林科技大学学报(自然科学版), 42(10), 95-102.]
[37]	Yao ZL, Wen HD, Deng Y, Cao M, Lin LX (2020) Driving forces underlying the beta diversity of tree species in subtropical mid-mountain moist evergreen broad-leaved forests in Ailao Mountains. Biodiversity Science, 28, 445- 454. (in Chinese with English abstract) DOI URL
	[ 姚志良, 温韩东, 邓云, 曹敏, 林露湘 (2020) 哀牢山亚热带中山湿性常绿阔叶林树种beta多样性格局形成的驱动力. 生物多样性, 28, 445-454.] DOI
[38]	Zhang JY, Wang WJ, Du HJ, Zhong ZL, Xiao L, Zhou W, Zhang B, Wang HY (2018) Differences in community characteristics, species diversity, and their coupling associations among three forest types in the Huzhong area, Daxinganling Mountains. Acta Ecologica Sinica, 38, 4684-4693. (in Chinese with English abstract)
	[ 张建宇, 王文杰, 杜红居, 仲召亮, 肖路, 周伟, 张波, 王洪元 (2018) 大兴安岭呼中地区3种林分的群落特征、物种多样性差异及其耦合关系. 生态学报, 38, 4684-4693.]
[39]	Zhang XT, Zhang JY, Li SW, Zhong ZL, Hu XG, Wang WJ (2021) Characteristics of plant diversity and community structure in Shuanghe Nature Reserve in Daxing’anling area of northeastern China. Journal of Beijing Forestry University, 43(7), 79-87. (in Chinese with English abstract)
	[ 张喜亭, 张建宇, 李斯雯, 仲召亮, 胡晓光, 王文杰 (2021) 大兴安岭双河保护区植物多样性和群落结构特征分析. 北京林业大学学报, 43(7), 79-87.]
[40]	Zhen Z, Guo ZY, Zhao YH, Li FR, Wei QB (2016) Spatial distribution of soil total nitrogen in Liangshui National Nature Reserve based on local model. Chinese Journal of Applied Ecology, 27, 549-558. (in Chinese with English abstract) PMID
	[ 甄贞, 郭志英, 赵颖慧, 李凤日, 魏庆彬 (2016) 基于局域模型的凉水国家自然保护区土壤全氮空间分布. 应用生态学报, 27, 549-558.] PMID
[41]	Zhu B, Chen AP, Liu ZL, Fang JY (2004) Plant community composition and tree species diversity on eastern and western Nanling Mountains, China. Biodiversity Science, 12, 53-62. (in Chinese with English abstract) DOI URL
	[ 朱彪, 陈安平, 刘增力, 方精云 (2004) 南岭东西段植物群落物种组成及其树种多样性垂直格局的比较. 生物多样性, 12, 53-62.] DOI
[42]	Zhu LJ, Jin GZ, Du WX, Wang XC (2016) Characteristics of canopy disturbance for a typical broadleaf-Korean pine mixed forest in Xiaoxing’an Mountains, Liangshui, northeastern China. Journal of Beijing Forestry University, 38(6), 17-27. (in Chinese with English abstract)
	[ 朱良军, 金光泽, 杜文先, 王晓春 (2016) 小兴安岭凉水典型阔叶红松林林冠干扰特征分析. 北京林业大学学报, 38(6), 17-27.]