王朗国家级自然保护区滑坡体兰科植物分布格局及其保护策略

doi:10.17520/biods.2021313

摘要/Abstract

摘要：

滑坡体是大熊猫国家公园岷山片区中广泛存在的一种生境, 但其在生物多样性研究和保护中的作用却没有得到足够的重视。王朗国家级自然保护区的滑坡体具有丰富的兰科植物资源, 其中白沙沟滑坡体最为丰富。本文以白沙沟滑坡体为例, 探究岷山地区滑坡体生境中兰科植物的空间分布格局, 拟为王朗国家级自然保护区及周边区域的兰科植物保护提供指导。本研究设置4条样带共186个样方, 利用除趋势对应分析(DCA)和双向指示种分析(TWINSPAN)研究滑坡体兰科植物的空间分布格局, 同时对兰科植物做了种间关联和相关性分析。结果表明, 白沙沟滑坡体的草本群落受环境因素的影响, 物种间竞争较弱, 处于自然演替的初始阶段。郁闭度和水流冲刷对滑坡体兰科植物的分布具有重要影响。受郁闭度和水流干扰因素的影响, 滑坡体可分成滑坡体-林缘生境(类型1)、滑坡体生境(类型2)和高郁闭度滑坡体生境(类型3) 3种类型。兰科植物在类型1和类型3两种生境中产生了适应性分化。建议在一些兰科物种集中分布的区域设立保护点并进行监测。此外还应对其他不同时间、不同土壤基质形成的滑坡体进行详细的调查, 摸清王朗国家级自然保护区甚至岷山地区兰科植物分布的整体情况。

关键词: 滑坡体, 兰科植物, 空间分布格局, 大熊猫国家公园

Abstract

Aims Landslide mass is a widely existing habitat type in the Minshan Mountains, but insufficient attention has been drawn to its importance in biodiversity research and protection. Orchid species are abundant in the landslide masses of the Wanglang National Nature Reserve, especially in the Baishagou landslide mass. By studying the spatial distribution pattern of orchids in the Baishagou landslide mass, we aimed to infer the approximate distribution pattern of orchids in the landslide mass of the Minshan Mountains and provide guidance for the protection of orchids in this area.

Methods We collected data from 186 quadrats belonging to 4 transects in the Baishagou landslide mass, and conducted interspecific association, correlation analysis, detrended correspondence analysis (DCA), and two-way indicator species analysis (TWINSPAN).

Results The results indicated that the herb community of the Baishagou landslide mass is primarily affected by environmental factors, and the competition among species is relatively weak. Canopy closure and water erosion both play important roles on the distribution of orchids in landslide masses. Landslide masses can be divided into three types: landslide-forest edge habitat (Type 1), landslide habitat (Type 2) and high canopy closure landslide habitat (Type 3). Adaptive differentiation has occurred between orchid species from Type 1 and Type 3.

Conclusion We suggest implementing protection points to monitor the richness of orchid distribution. In addition, detailed investigations should be carried out on other landslide masses which formed at different points in time and with different soil substrates to find out the distribution pattern of orchid species in the Wanglang National Nature Reserve, or even the entire Minshan area.

Key words: landslide mass, orchid, spatial distribution pattern, Giant Panda National Park

王重阳, 赵联军, 孟世勇 (2022) 王朗国家级自然保护区滑坡体兰科植物分布格局及其保护策略. 生物多样性, 30, 21313. DOI: 10.17520/biods.2021313.

Chongyang Wang, Lianjun Zhao, Shiyong Meng (2022) Spatial distribution pattern and protection strategy for orchids in landslide mass of the Wanglang National Nature Reserve. Biodiversity Science, 30, 21313. DOI: 10.17520/biods.2021313.

导出引用管理器 EndNote|Ris|BibTeX

链接本文: https://www.biodiversity-science.net/CN/10.17520/biods.2021313

https://www.biodiversity-science.net/CN/Y2022/V30/I2/21313

图/表 8

图1 野外调查的地点以及样带划分示意图

Fig. 1 The location of the field investigation and schematic diagram of transects

表1 每条样带的样方、物种和多样性指数总结表: 以Shannon-Wiener为例

Table 1 A summary of quadrats, species and diversity indices (Shannon-Wiener index, specifically) of each transect

	样带A Transect A	样带B Transect B	样带C Transect C	样带D Transect D
样方数 Quadrat quantity	20	38	68	60
总物种数 Total species quantity	30	40	62	40
兰科植物物种数 Orchid species quantity	4	5	8	9
Shannon-Wiener指数 Shannon-Wiener index	3.040	3.309	3.582	3.125

表2 每条样带中兰科植物平均盖度(× 100)分布列表

Table 2 Distribution table of average coverage (multiplied 100 times) for orchid species in each transect

物种 Species	样带A Transect A	样带B Transect B	样带C Transect C	样带D Transect D
抱茎叶无柱兰 Amitostigma amplexifolium	0.48	0	0.51	0
一花无柱兰 Amitostigma monanthum	0	0	0	4.92
无苞杓兰 Cypripedium bardolphianum	0	5.13	0	0
华西杓兰 Cypripedium farreri	0	0	0	4.1
毛杓兰 Cypripedium franchetii	107.14	8.21	5.94	0
紫点杓兰 Cypripedium guttatum	0	1.28	0	0
西藏杓兰 Cypripedium tibeticum	0	0	0	27.21
凹舌掌裂兰 Dactylorhiza viridis	0	0	0	0.16
西南手参 Gymnadenia orchidis	0	0	6.09	0
北方盔花兰 Galearis roborowskyi	0	0	0.2	0.16
二叶盔花兰 Galearis spathulata	0	0	4.83	3.28
短梗山兰 Oreorchis erythrochrysea	0	0	2.93	1.48
少花鹤顶兰 Phaius delavayi	61.9	78.46	27.54	16.39
广布小红门兰 Ponerorchis chusua	2.86	4.1	4.09	49.34

表3 优势非兰科植物和优势兰科植物物种重要值排序表(样方数N = 186)

Table 3 Characteristics and importance values of dominant non-orchid and orchid species (number of quadrats N = 186)

物种 Species		平均高度 Average height (cm)	总盖度 Total coverage	频度 Frequency	相对高度 Relative height (%)	相对盖度 Relative coverage (%)	相对频度 Relative frequency (%)	重要值 Importance value (%)	排名 Order	物种编号 Species no.
主要非兰科植物 Major non-orchid plants
	独花黄精 Polygonatum hookeri	5.24	1210.85	177	0.69	26.39	12.47	13.18	1	1
	圆穗蓼 Polygonum macrophyllum	4.80	596.97	119	0.63	13.01	8.39	7.34	2	2
	绢毛绣线菊 Spiraea sericea	13.02	632.10	46	1.72	13.77	3.24	6.25	3	3
	膨囊薹草 Carex lehmannii	5.86	362.80	83	0.77	7.91	5.85	4.84	4	4
	北极果 Arctous alpinus	5.59	296.70	56	0.74	6.47	3.95	3.72	5	5
	樱草杜鹃 Rhododendron primuliflorum	13.43	183.20	30	1.78	3.99	2.11	2.63	6	6
	百蕊草 Thesium chinense	5.93	109.99	66	0.78	2.40	4.65	2.61	7	7
	当归 Angelica sinensis	4.14	65.70	63	0.55	1.43	4.44	2.14	8	8
	柳属某种 Salix sp.	15.54	113.90	26	2.05	2.48	1.83	2.12	9	9
	尼泊尔蓼 Polygonum nepalense	5.24	114.70	41	0.69	2.50	2.89	2.03	10	10
	窄叶鲜卑花 Sibiraea angustata	17.84	47.90	19	2.36	1.04	1.34	1.58	11	11
	薄叶铁线莲 Clematis gracilifolia	9.00	60.50	31	1.19	1.32	2.18	1.56	12	12
	钟花龙胆 Gentiana nanobella	1.09	16.92	58	0.14	0.37	4.09	1.53	13	13
	方枝柏 Juniperus saltuaria	18.82	27.28	17	2.49	0.59	1.20	1.43	16	14
	圆叶小堇菜 Viola biflora var. rockiana	1.71	29.09	48	0.23	0.63	3.38	1.41	17	15
主要兰科植物 Major orchid plants
	少花鹤顶兰 Phaius delavayi	10.36	73.60	22	1.37	1.60	1.55	1.51	14	16
	广布小红门兰 Ponerorchis chusua	5.73	35.12	41	0.76	0.77	2.89	1.47	15	17
	毛杓兰 Cypripedium franchetii	6.73	30.30	30	0.89	0.66	2.11	1.22	20	18
	西藏杓兰 Cypripedium tibeticum	7.50	16.60	12	0.99	0.36	0.85	0.73	40	19
	西南手参 Gymnadenia orchidis	11.57	4.20	7	1.53	0.09	0.49	0.70	41	20
	一花无柱兰 Amitostigma monanthum	8.50	3.00	2	1.12	0.07	0.14	0.44	56	21
	短梗山兰 Oreorchis erythrochrysea	6.33	2.92	6	0.84	0.06	0.42	0.44	57	22
	华西杓兰 Cypripedium farreri	6.67	2.50	3	0.88	0.05	0.21	0.38	62	23
	二叶盔花兰 Galearis spathulata	4.67	3.33	6	0.62	0.07	0.42	0.37	63	24
	抱茎叶无柱兰 Amitostigma amplexifolium	6.50	0.20	2	0.86	<0.01	0.14	0.33	66	25

表3 优势非兰科植物和优势兰科植物物种重要值排序表(样方数N = 186)

Table 3 Characteristics and importance values of dominant non-orchid and orchid species (number of quadrats N = 186)

物种 Species		平均高度 Average height (cm)	总盖度 Total coverage	频度 Frequency	相对高度 Relative height (%)	相对盖度 Relative coverage (%)	相对频度 Relative frequency (%)	重要值 Importance value (%)	排名 Order	物种编号 Species no.
主要非兰科植物 Major non-orchid plants
	独花黄精 Polygonatum hookeri	5.24	1210.85	177	0.69	26.39	12.47	13.18	1	1
	圆穗蓼 Polygonum macrophyllum	4.80	596.97	119	0.63	13.01	8.39	7.34	2	2
	绢毛绣线菊 Spiraea sericea	13.02	632.10	46	1.72	13.77	3.24	6.25	3	3
	膨囊薹草 Carex lehmannii	5.86	362.80	83	0.77	7.91	5.85	4.84	4	4
	北极果 Arctous alpinus	5.59	296.70	56	0.74	6.47	3.95	3.72	5	5
	樱草杜鹃 Rhododendron primuliflorum	13.43	183.20	30	1.78	3.99	2.11	2.63	6	6
	百蕊草 Thesium chinense	5.93	109.99	66	0.78	2.40	4.65	2.61	7	7
	当归 Angelica sinensis	4.14	65.70	63	0.55	1.43	4.44	2.14	8	8
	柳属某种 Salix sp.	15.54	113.90	26	2.05	2.48	1.83	2.12	9	9
	尼泊尔蓼 Polygonum nepalense	5.24	114.70	41	0.69	2.50	2.89	2.03	10	10
	窄叶鲜卑花 Sibiraea angustata	17.84	47.90	19	2.36	1.04	1.34	1.58	11	11
	薄叶铁线莲 Clematis gracilifolia	9.00	60.50	31	1.19	1.32	2.18	1.56	12	12
	钟花龙胆 Gentiana nanobella	1.09	16.92	58	0.14	0.37	4.09	1.53	13	13
	方枝柏 Juniperus saltuaria	18.82	27.28	17	2.49	0.59	1.20	1.43	16	14
	圆叶小堇菜 Viola biflora var. rockiana	1.71	29.09	48	0.23	0.63	3.38	1.41	17	15
主要兰科植物 Major orchid plants
	少花鹤顶兰 Phaius delavayi	10.36	73.60	22	1.37	1.60	1.55	1.51	14	16
	广布小红门兰 Ponerorchis chusua	5.73	35.12	41	0.76	0.77	2.89	1.47	15	17
	毛杓兰 Cypripedium franchetii	6.73	30.30	30	0.89	0.66	2.11	1.22	20	18
	西藏杓兰 Cypripedium tibeticum	7.50	16.60	12	0.99	0.36	0.85	0.73	40	19
	西南手参 Gymnadenia orchidis	11.57	4.20	7	1.53	0.09	0.49	0.70	41	20
	一花无柱兰 Amitostigma monanthum	8.50	3.00	2	1.12	0.07	0.14	0.44	56	21
	短梗山兰 Oreorchis erythrochrysea	6.33	2.92	6	0.84	0.06	0.42	0.44	57	22
	华西杓兰 Cypripedium farreri	6.67	2.50	3	0.88	0.05	0.21	0.38	62	23
	二叶盔花兰 Galearis spathulata	4.67	3.33	6	0.62	0.07	0.42	0.37	63	24
	抱茎叶无柱兰 Amitostigma amplexifolium	6.50	0.20	2	0.86	<0.01	0.14	0.33	66	25

图2 有效样方分布的除趋势对应分析(DCA)二维排序图。图中A、B、C、D代表4条样带, 数字代表样方在样带中的编号。

Fig. 2 Detrended correspondence analysis (DCA) two-dimension sorting result of valid quadrat distribution. A, B, C and D represent four transects, respectively, and numbers represent the position of the quadrat in each transect.

图3 双向指示种分析聚类结果。(a)双向指示种分析聚类结果在除趋势对应分析(DCA)排序图中的叠加, 灰色点代表类型1, 粉红色点代表类型2, 绿色点代表类型3, 蓝色点代表类型4, 橙色点代表类型5。聚类结果中, 含样方数量较多并且在排序平面中具有明显分布范围的类型, 全部用多边形沿边界线框起、涂色并标上编号, 作为主要类型。(b)样方聚类结果在样带中的空间位置示意图, 横轴表示样方的数字编号, 纵轴表示4条样带, 颜色与类型的对应关系与(a)相同。

Fig. 3 The results of two-way indicator species analysis (TWINSPAN) clustering. (a) The superposition of TWINSPAN clustering and detrended correspondence analysis (DCA) sorting results, in which gray colored spots represent Group 1, pink represents Group 2, green represents Group 3, blue represents Group 4, and orange represents Group 5, respectively. (b) The spatial positions of TWINSPAN clustering groups in transects in a general view. The horizontal axis represents the serial number of each belt, and the vertical axis indicates four transects. The correlation between colour and groups are identical to that in (a).

图4 种间关联卡方检验结果的半矩阵图。*表示正相关(* P ≤ 0.05; ** P ≤ 0.01), #表示负相关(# P ≤ 0.05; ## P ≤ 0.01); 1-25表示物种编号, 详见表3。

Fig. 4 Semi-matrix diagram of the chi-square test result showing interspecies association. * indicates that correlation between species is positive (*P ≤ 0.05; ** P ≤ 0.01); # indicates negative correlation between species (# P ≤ 0.05; ## P ≤ 0.01). Numbers 1–25 represent the species numbers, as shown in Table 3. Numbers 1-25 represent the species numbers, as shown in Table 3.

图5 种间Spearman相关性的半矩阵图。蓝色代表正相关, 红色代表负相关, 颜色越深相关性越显著。×表示相应的物种对相关性不显著, 1-25表示物种编号, 详见表3。

Fig. 5 Semi-matrix diagram of Spearman correlation between species. Blue represents positive correlation, and red represents negative correlation. The darker the color, the more significant the relevance between species. × represent insignificant correlation between relative species. Numbers 1-25 represent the species numbers, as shown in Table 3.

参考文献 34

[1]	Cui YH, Han YZ, Zhang MT, Yang XQ, Zhao ZH (2021) Spatial pattern and interspecific association of tree species in mixed coniferous and deciduous broad-leaved mixed forest under different disturbance intensities. Chinese Journal of Applied Ecology, 32, 2053-2060. (in Chinese with English abstract)
	[ 崔玉华, 韩有志, 张梦弢, 杨秀清, 赵占合 (2021) 不同干扰强度下针阔混交林树种空间格局及种间关联性. 应用生态学报, 32, 2053-2060.]
[2]	Deng S, Wu YN, Wu KL, Fang L, Li L, Zeng SJ (2020) Breeding characteristics and artificial propagation of 14 species of Wild Plant with Extremely Small Populations (WPESP) in China. Biodiversity Science, 28, 385-400. (in Chinese with English abstract) DOI
	[ 邓莎, 吴艳妮, 吴坤林, 房林, 李琳, 曾宋君 (2020) 14种中国典型极小种群野生植物繁育特性和人工繁殖研究进展. 生物多样性, 28, 385-400.] DOI
[3]	Højsgaard S, Halekoh U (2021) doBy: Groupwise Statistics, LSmeans, Linear Contrasts, Utilities. R package version 4.6.9.. https://CRAN.R-project.org/package=doBy/. (accessed on 2021-02-25)
[4]	Huang BQ, Luo YB, An DJ, Kou Y (2010) Characteristics of plant species in herb community in Huanglong Valley, Sichuan. Bulletin of Botanical Research, 30, 543-548. (in Chinese with English abstract)
	[ 黄宝强, 罗毅波, 安德军, 寇勇 (2010) 四川黄龙沟草本层植物种类组成与数量特征. 植物研究, 30, 543-548.]
[5]	Huang BQ, Luo YB, Yu FH, Tang SY, Dong L, An DJ (2007) Interspecific relationships of dominant species in orchid communities of forest vegetation in Huanglong Valley, Sichuan, China. Journal of Plant Ecology (Chinese Version), 31, 865-872. (in Chinese with English abstract)
	[ 黄宝强, 罗毅波, 于飞海, 唐思远, 董立, 安德军 (2007) 四川黄龙沟森林植被中兰科植物群落优势种种间联结和相关分析. 植物生态学报, 31, 865-872.] DOI
[6]	Huang BQ, Yang XQ, Yu FH, Luo YB, Tai YD (2008) Surprisingly high orchid diversity in travertine and forest areas in the Huanglong Valley, China, and implications for conservation. Biodiversity and Conservation, 17, 2773- 2786.
[7]	Janečková P, Wotavová K, Schödelbauerová I, Jersáková J, Kindlmann P (2006) Relative effects of management and environmental conditions on performance and survival of populations of a terrestrial orchid, Dactylorhiza majalis. Biological Conservation, 129, 40-49.
[8]	Li P, Tang SY, Dong L, Luo YB, Kou Y, Yang XQ, Perner H (2005) Species diversity and flowering phenology of Orchidaceae in Huanglong Valley, Sichuan. Biodiversity Science, 13, 255-261. (in Chinese with English abstract) DOI URL
	[ 李鹏, 唐思远, 董立, 罗毅波, 寇勇, 杨小琴,Perner H (2005) 四川黄龙沟兰科植物的多样性及其保护. 生物多样性, 13, 255-261.] DOI
[9]	Li S, Wang DJ, Gu XD, McShea WJ (2010) Beyond pandas, the need for a standardized monitoring protocol for large mammals in Chinese nature reserves. Biodiversity and Conservation, 19, 3195-3206. DOI URL
[10]	Li SG, Qu H (2007) Practical Biostatistics, pp. 112-121. Peking University Press, Beijing. (in Chinese)
	[ 李松岗, 曲红 (2007) 实用生物统计, 112-121页. 北京大学出版社, 北京.]
[11]	Li WL, Wu J, Lü BX (2011) Research on landslide triggered by earthquake: Review and prospect. Journal of Catastrophology, 26, 103-108. (in Chinese with English abstract)
	[ 李为乐, 伍霁, 吕宝雄 (2011) 地震滑坡研究回顾与展望. 灾害学, 26, 103-108.]
[12]	Lin HQ, Yang PY, Liu GY, Cheng YH, Tan YC, He TM (2020) Diversity and conservation of Orchidaceae plants in Wolong National Nature Reserve. Journal of Sichuan Forestry Science and Technology, 41(3), 14-22. (in Chinese with English abstract)
	[ 林红强, 杨攀艳, 刘桂英, 程跃红, 谭迎春, 何廷美 (2020) 卧龙国家级自然保护区兰科植物多样性及保护研究. 四川林业科技, 41(3), 14-22.]
[13]	Liu LX, Li XQ, Zhou ZC, Hou XW, Wang WZ, Wang ZX (2011) An experimental study of the initiation mechanism of landslide debris flow under a strong earthquake. Hydrogeology & Engineering Geology, 38, 104-109. (in Chinese with English abstract)
	[ 刘玲霞, 李向全, 周志超, 侯新伟, 王文中, 王振兴 (2011) 强震条件下谢家店滑坡碎屑流发生机制试验研究. 水文地质工程地质, 38, 104-109.]
[14]	Liu SJ, Zhang B, Yang QW, Hu CH, Shu CQ (2010) Research on the natural recovery of vegetation on the non-normative landslide mass in Wenchuan earthquake-Take landslide mass in Xiejiadian of Yingchanggou in Pengzhou as an example. Journal of Mountain Science, 28, 373-378. (in Chinese with English abstract)
	[ 刘守江, 张斌, 杨清伟, 胡翠华, 舒成强 (2010) 汶川地震非规范滑坡体上植被的自然恢复能力研究--以彭州银厂沟谢家店子滑坡体为例. 山地学报, 28, 373-378.]
[15]	Liu ZH, Tian YP, An DJ, Wang HJ, Tang S, Zhang JL, Sun HL, Liu Y, Zhang QM (2009) Formation and evolution of the travertine landscape at Huanglong, Sichuan, one of the world natural heritages. Acta Geoscientica Sinica, 30, 841-847. (in Chinese with English abstract)
	[ 刘再华, 田友萍, 安德军, 王海静, 唐淑, 张金流, 孙海龙, 刘彦, 张清明 (2009) 世界自然遗产-四川黄龙钙华景观的形成与演化. 地球学报, 30, 841-847.]
[16]	Luo YB, Jia JS, Wang CL (2003) A general review of the conservation status of Chinese orchids. Biodiversity Science, 11, 70-77. (in Chinese with English abstract) DOI URL
	[ 罗毅波, 贾建生, 王春玲 (2003) 中国兰科植物保育的现状和展望. 生物多样性, 11, 70-77.] DOI
[17]	Medici EP, Desbiez ALJ (2012) Population viability analysis: Using a modeling tool to assess the viability of tapir populations in fragmented landscapes. Integrative Zoology, 7, 356-372.
[18]	Oksanen J, Blanchet FG, Friendly M, Kindt R, Legendre P, McGlinn D, Minchin PR, O’Hara RB, Simpson GL, Solymos P, Stevens HMH, Szoecs E, Wagner H (2020) vegan: Community Ecology Package. R package version 2.5-7. https://CRAN.R-project.org/package=vegan/. (accessed on 2021-02-03)
[19]	Pielou EC (1966) The measurement of diversity in different types of biological collections. Journal of Theoretical Biology, 13, 131-144. DOI URL
[20]	Qin WH, Jiang MK, Xu WG, He ZH (2012) Assessment of in situ conservation of 1,334 native orchids in China. Biodiversity Science, 20, 177-183. (in Chinese with English abstract) DOI URL
	[ 秦卫华, 蒋明康, 徐网谷, 贺昭和 (2012) 中国1,334种兰科植物就地保护状况评价. 生物多样性, 20, 177-183.] DOI
[21]	Revelle W (2020) psych: Procedures for Personality and Psychological Research, Northwestern University, Evanston, Illinois, USA, https://CRAN.R-project.org/package=psychVersion=2.1.3/ (accessed on 2021-02-03)
[22]	Roleček J, Tichý L, Zelený D, Chytrý M (2009) Modified TWINSPAN classification in which the hierarchy respects cluster heterogeneity. Journal of Vegetation Science, 20, 596-602. DOI URL
[23]	Shannon CE, Wiener W (1963) The Mathematical Theory of Communication. University of Illinois Press. Urbana.
[24]	Simpson EH (1949) Measurement of diversity. Nature, 163, 688. DOI URL
[25]	Wan SL, Zhang JL, Liu MJ, He WM, Li HL, Guo CB, Li ZM (2020) Seismogenic structure and seismic activity analysis of Minshan block. Earthquake, 40(2), 49-70. (in Chinese with English abstract)
	[ 万森林, 张军龙, 刘明军, 贺为民, 李海龙, 郭长宝, 李智敏 (2020) 岷山断块的发震构造与地震活动性分析. 地震, 40(2), 49-70.]
[26]	Wei TY, Simko V (2017) R package “corrplot”: Visualization of a Correlation Matrix (Version 0. 84). https://github.com/taiyun/corrplot/. (accessed on 2021-02-03)
[27]	Yang CS, Dong JH, Zhu SN, Xiong GH (2021) Detection, identification and deformation characteristics of landslide groups by InSAR in Batang section of Jinsha River convergence zone, China. Journal of Earth Sciences and Environment, 43, 398-408. (in Chinese with English abstract)
	[ 杨成生, 董继红, 朱赛楠, 熊国华 (2021) 金沙江结合带巴塘段滑坡群InSAR探测识别与形变特征. 地球科学与环境学报, 43, 398-408.]
[28]	Zeleny D (2021) twinspanR: TWo-way INdicator SPecies ANalysis (and its modified version) in R. R package version 0. 22. . (accessed on 2021-02-02)
[29]	Zeng SJ, Huang WC, Wu KL, Zhang JX, Teixeira da Silva JA, Duan J (2016) In vitro propagation of Paphiopedilum orchids. Critical Reviews in Biotechnology, 36, 521-534.
[30]	Zhang F, Wang JZ, Li CC, Yang QY (2021) Analysis of deformation and failure characteristics and formation mechanism of Zheduotang landslide on the Sichuan-Tibet Railway. Journal of Geological Hazards and Environment Preservation, 32(2), 3-7. (in Chinese with English abstract)
	[ 张凡, 王家柱, 李长春, 杨清云 (2021) 川藏线折多塘滑坡变形破坏特征及其形成机制分析. 地质灾害与环境保护, 32(2), 3-7.]
[31]	Zhang JL (2016) spaa: SPecies Association Analysis. R package version 0.2.2. https://CRAN.R-project.org/package=spa/. (accessed on 2021-02-03)
[32]	Zhang JT (2018) Quantitative Ecology, pp 33-34. Science Press, Beijing. (in Chinese)
	[ 张金屯 (2018) 数量生态学, 33-34页. 科学出版社, 北京.]
[33]	Zhang YB, Du HD, Jin XH, Ma KP (2015) Species diversity and geographic distribution of wild Orchidaceae in China. Chinese Science Bulletin, 60, 179-188. (in Chinese with English abstract)
	[ 张殷波, 杜昊东, 金效华, 马克平 (2015) 中国野生兰科植物物种多样性与地理分布. 科学通报, 60, 179-188.]
[34]	Zhu XW, Zhang R, Zhou XL (1989) Basic characteristics of karst in Minshan Mt., west Sichuan. Carsologica Sinica, 8, 253-260. (in Chinese with English abstract)
	[ 朱学稳, 张任, 周绪伦 (1989) 川西岷山岩溶的基本特征. 中国岩溶, 8, 253-260.]