地形异质性对云南普洱季风常绿阔叶林物种多样性的影响

doi:10.17521/cjpe.2020.0148

植物生态学报 ›› 2020, Vol. 44 ›› Issue (10): 1015-1027.DOI: 10.17521/cjpe.2020.0148

地形异质性对云南普洱季风常绿阔叶林物种多样性的影响

王艳红¹, 李帅锋¹, 郎学东¹, 黄小波¹, 刘万德¹, 徐崇华², 苏建荣¹^,^*()

¹中国林业科学研究院资源昆虫研究所, 国家林业和草原局云南普洱森林生态系统国家定位观测研究站, 昆明 650224
²云南省太阳河自然保护区管理局, 云南普洱 665000

收稿日期:2020-05-12 接受日期:2020-08-21 出版日期:2020-10-20 发布日期:2020-11-24
通讯作者: 苏建荣
作者简介:*E-mail:jianrongsu@vip.sina.com
基金资助:
云南省科技创新人才计划(2018HC013);云南省基础研究计划(2019FB058);云南省基础研究计划(202001AS070005)

Effects of topographic heterogeneity on species diversity in a monsoon evergreen broad- leaved forest in Puʼer, Yunnan, China

WANG Yan-Hong¹, LI Shuai-Feng¹, LANG Xue-Dong¹, HUANG Xiao-Bo¹, LIU Wan-De¹, XU Chong-Hua², SU Jian-Rong¹^,^*()

¹Research Institute of Resource Insects, Chinese Academy of Forestry, Pu’er Forest Ecosystem Research Station, National Forestry and Grassland Administration of China, Kunming 650224
²Taiyanghe Provincinal Nature Reserve of Yunnan, Pu’er, Yunnan 665000, China

Received:2020-05-12 Accepted:2020-08-21 Online:2020-10-20 Published:2020-11-24
Contact: SU Jian-Rong
Supported by:
Yunnan Science and Technology Innovation Talent Program(2018HC013);Applied Basic Research Foundation of Yunnan Province(2019FB058);Applied Basic Research Foundation of Yunnan Province(202001AS070005)

摘要/Abstract

摘要：

地形异质性通过调控树木生长所需的养分、水分和光照等而成为亚热带森林结构与物种组成的重要驱动因子。但是, 地形异质性对季风常绿阔叶林物种多样性及其分布影响的研究还相对较少。该文基于云南普洱30 hm²森林动态监测样地(大样地) 750个20 m × 20 m的样方调查数据, 以海拔、坡度、凹凸度和坡向4个地形因子为变量, 采用C均值模糊聚类分析大样地的地形类型, 进而分析不同地形条件下的群落物种组成及群落物种多样性; 采用Torus转换检验法, 探讨物种与地形关联性, 为季风常绿阔叶林生物多样性保护提供科学依据。研究结果表明, 大样地可分为山脊、陡坡、缓坡、高谷和沟谷等5类地形, 地形面积分别是8.00、6.04、7.68、2.76和5.52 hm²。大样地中胸径(DBH) ≥ 1 cm的木本植物个体153 418株, 分属79科179属271种。5类地形中, 物种丰富度、不同径级的植株密度和比例明显不同, 多样性及优势物种多度分布具有较大差异。种-面积曲线表明, 同等面积条件下, 随着取样面积增加, 山脊的物种丰富度始终最小, 高谷次之, 沟谷的物种丰富度始终最大。种-个体数累积曲线表明, 随着个体数增加, 山脊物种丰富度的累积速率最小, 种丰富度增加缓慢, 高谷次之。在被检验的123个物种中, 与地形相关的物种有83个, 高达67.5%的物种与至少一类地形存在显著相关关系。山脊和缓坡中与地形具有显著负相关关系的物种数超过显著正相关的物种数; 而与陡坡、高谷和沟谷显著正相关的物种数高于显著负相关的物种数。普洱大样地地形异质性对物种多样性维持的贡献率为7.8%。

关键词: 地形异质性, 季风常绿阔叶林, 物种组成, 群落结构, 重要值, 物种多样性, 物种地形关联

Abstract:

Aims In subtropical forests, topographic heterogeneity drives ecosystem structure and species composition by regulating the available nutrients, water and light needed for tree growth. A gap in current research is the role of topographic heterogeneity in plant species diversity and distribution in monsoon evergreen broad-leaved forests. Our study aims to fill this gap, providing scientific grounds for the conservation of biodiversity through an exploration of floral structure and ecosystem development in monsoon evergreen broad-leaved forests.
Methods Within a 30 hm² dynamics plot, we established 750 20 m × 20 m sample plots, using fuzzy C-mean clustering to classify different topographic types based on elevations, convexities, slopes, and aspects of each sample plots. We compared community composition and community species diversity between different topographic types, using Torus-translation tests to examine the relationship between floral species and topography.
Important findings We identified five distinct topography types: ridge (8.00 hm²), steep slope (6.04 hm²), less-steep slope (7.68 hm²), high plateau (2.76 hm²), and valley (5.52 hm²). We counted a total of 153 418 woody plant individuals with a diameter at breast height (DBH) ≥ 1 cm, belonging to 271 species, 179 genera and 79 families. Among the topographic types, we noted differences in species richness, plant density, proportion of different diameters, species diversity and the abundance of dominant species. When comparing species-area relationships between the five topographic types, we noticed that the ridge had the lowest number of species, while the valley had the highest. The cumulative species-individual relationships within the topographic types revealed that the ridge possesses the lowest cumulative rate. Of the 123 examined species, 67.5% were significantly related to at least one type of topography. Among species significantly related to topography, those species with significant negative correlation were more abundant in the ridge and steep slope topographic types. The opposite was true in steep slope, high plateau and valley. Based on our data, we determined that topographic heterogeneity contributed 7.8% to the maintenance of species diversity in our study area.

Key words: topographic heterogeneity, monsoon broad-leaved evergreen forest, species composition, community structure, importance value, species diversity, topographic association of woody plants

王艳红, 李帅锋, 郎学东, 黄小波, 刘万德, 徐崇华, 苏建荣. 地形异质性对云南普洱季风常绿阔叶林物种多样性的影响. 植物生态学报, 2020, 44(10): 1015-1027. DOI: 10.17521/cjpe.2020.0148

WANG Yan-Hong, LI Shuai-Feng, LANG Xue-Dong, HUANG Xiao-Bo, LIU Wan-De, XU Chong-Hua, SU Jian-Rong. Effects of topographic heterogeneity on species diversity in a monsoon evergreen broad- leaved forest in Puʼer, Yunnan, China. Chinese Journal of Plant Ecology, 2020, 44(10): 1015-1027. DOI: 10.17521/cjpe.2020.0148

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2020.0148

https://www.plant-ecology.com/CN/Y2020/V44/I10/1015

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地形 Topography	海拔 Elevation (m)	坡度 Slope (°)	凹凸度 Convexity (m)	坡向 Slope aspect (°)	样地数 Sample plot number
山脊 Ridge	1 547.0 ± 0.96^a	19.6 ± 0.33^a	0.67 ± 0.14^a	164.41 ± 1.56^a	200
陡坡 Steep slope	1 525.7 ± 1.62^b	19.4 ± 0.50^a	-0.17 ± 0.18^b	324.74 ± 2.19^b	151
缓坡 Less-steep slope	1 504.5 ± 1.18^c	18.4 ± 0.34^b	-0.42 ± 0.12^b	185.70 ± 1.27^c	192
高谷 High plateau	1 548.7 ± 3.10^a	19.5 ± 0.56^a	0.41 ± 0.21^a	52.53 ± 4.52^d	69
沟谷 Valley	1 498.3 ± 1.89^d	18.8 ± 0.52^a	-0.91 ± 0.20^bc	294.68 ± 2.23^e	138

地形 Topography	海拔 Elevation (m)	坡度 Slope (°)	凹凸度 Convexity (m)	坡向 Slope aspect (°)	样地数 Sample plot number
山脊 Ridge	1 547.0 ± 0.96^a	19.6 ± 0.33^a	0.67 ± 0.14^a	164.41 ± 1.56^a	200
陡坡 Steep slope	1 525.7 ± 1.62^b	19.4 ± 0.50^a	-0.17 ± 0.18^b	324.74 ± 2.19^b	151
缓坡 Less-steep slope	1 504.5 ± 1.18^c	18.4 ± 0.34^b	-0.42 ± 0.12^b	185.70 ± 1.27^c	192
高谷 High plateau	1 548.7 ± 3.10^a	19.5 ± 0.56^a	0.41 ± 0.21^a	52.53 ± 4.52^d	69
沟谷 Valley	1 498.3 ± 1.89^d	18.8 ± 0.52^a	-0.91 ± 0.20^bc	294.68 ± 2.23^e	138

参数 Parameter	面积 Area (hm²)	个体数 Number of individuals	植株密度 Plant density (individual·hm^-2)	根萌生 Root sprouting	干基萌生 Stem basal sprouting	干萌生 Stem epicormic	枝萌生 Branch epicormic
大样地 Big sample plot	30.00	153 418	5 114	14 691	675	721	-
山脊 Ridge	8.00	44 861	5 608	5 222	98	142	-
陡坡 Steep slope	6.04	29 703	4 918	2 687	118	113	1
缓坡 Less-steep slope	7.68	39 320	5 120	3 295	203	207	2
高谷 High plateau	2.76	13 843	5 016	1 682	62	80	-
沟谷 Valley	5.52	25 691	4 654	1 805	195	179	38

参数 Parameter	面积 Area (hm²)	个体数 Number of individuals	植株密度 Plant density (individual·hm^-2)	根萌生 Root sprouting	干基萌生 Stem basal sprouting	干萌生 Stem epicormic	枝萌生 Branch epicormic
大样地 Big sample plot	30.00	153 418	5 114	14 691	675	721	-
山脊 Ridge	8.00	44 861	5 608	5 222	98	142	-
陡坡 Steep slope	6.04	29 703	4 918	2 687	118	113	1
缓坡 Less-steep slope	7.68	39 320	5 120	3 295	203	207	2
高谷 High plateau	2.76	13 843	5 016	1 682	62	80	-
沟谷 Valley	5.52	25 691	4 654	1 805	195	179	38

地形 Topography	物种 Species	相对频度 Relative frequency	相对多度 Relative abundance	相对胸高断面积 Relative basal area at the breast height	重要值 Importance value
山脊 Ridge	短刺锥 Castanopsis echidnocarpa	3.7	28.5	34.0	22.0
	泥柯 Lithocarpus fenestratus	3.7	14.5	6.5	8.2
	截果柯 Lithocarpus truncatus	3.7	5.3	7.1	5.3
	西南木荷 Schima wallichii	3.6	2.6	9.7	5.3
	茶梨 Anneslea fragrans	3.5	4.4	5.4	4.4
	隐距越桔 Vaccinium exaristatum	3.5	5.4	3.6	4.2
	红梗润楠 Machilus rufipes	3.6	4.6	2.5	3.6
	红叶木姜子 Litsea rubescens	3.6	4.3	2.5	3.5
	红皮水锦树 Wendlandia tinctoria subsp. intermedia	3.3	3.0	2.8	3.0
	深绿山龙眼 Helicia nilagirica	3.5	3.3	2.1	3.0
陡坡 Steep slope	短刺锥 Castanopsis echidnocarpa	2.8	16.4	22.8	14.0
	泥柯 Lithocarpus fenestratus	2.6	9.8	4.2	5.5
	西南木荷 Schima wallichii	2.8	3.4	9.8	5.3
	红梗润楠 Machilus rufipes	2.8	5.5	4.4	4.2
	枹丝锥 Castanopsis calathiformis	1.9	5.6	5.0	4.2
	截果柯 Lithocarpus truncatus	2.6	4.1	5.4	4.0
	红叶木姜子 Litsea rubescens	2.7	4.5	2.6	3.2
	西桦 Betula alnoides	0.9	0.5	7.4	3.0
	茶梨 Anneslea fragrans	2.5	3.4	2.9	2.9
	隐距越桔 Vaccinium exaristatum	2.2	2.8	1.8	2.2
缓坡 Less-steep slope	短刺锥 Castanopsis echidnocarpa	2.8	21.8	24.7	16.5
	西南木荷 Schima wallichii	2.8	3.2	14.3	6.8
	泥柯 Lithocarpus fenestratus	2.8	10.1	6.3	6.4
	枹丝锥 Castanopsis calathiformis	2.5	7.2	4.4	4.7
	红梗润楠 Machilus rufipes	2.9	6.0	4.8	4.6
	截果柯 Lithocarpus truncatus	2.7	4.2	6.3	4.4
	茶梨 Anneslea fragrans	2.6	4.0	3.7	3.5
	红叶木姜子 Litsea rubescens	2.7	3.7	3.8	3.4
	隐距越桔 Vaccinium exaristatum	2.0	2.4	1.7	2.0
	艾胶算盘子 Glochidion lanceolarium	2.7	2.4	0.6	1.9
高谷 High plateau	短刺锥 Castanopsis echidnocarpa	3.1	16.7	24.0	14.6
	泥柯 Lithocarpus fenestratus	3.0	17.2	5.7	8.6
	西南木荷 Schima wallichii	3.2	3.2	13.5	6.7
	红梗润楠 Machilus rufipes	3.2	5.7	5.0	4.7
	截果柯 Lithocarpus truncatus	2.8	3.9	5.7	4.1
	红叶木姜子 Litsea rubescens	3.1	5.4	2.9	3.8
	西桦 Betula alnoides	1.4	0.8	7.7	3.3
	隐距越桔 Vaccinium exaristatum	2.7	4.0	2.4	3.0
	茶梨 Anneslea fragrans	3.1	2.8	2.7	2.9
	枹丝锥 Castanopsis calathiformis	2.0	2.6	2.7	2.5
沟谷 Valley	短刺锥 Castanopsis echidnocarpa	2.5	12.9	15.6	10.4
	西南木荷 Schima wallichii	2.4	3.4	10.9	5.6
	泥柯 Lithocarpus fenestratus	2.3	9.5	4.2	5.4
	红梗润楠 Machilus rufipes	2.6	5.6	6.2	4.8
	截果柯 Lithocarpus truncatus	2.4	3.7	6.3	4.1
	枹丝锥 Castanopsis calathiformis	1.9	5.6	3.1	3.5
	云南瘿椒树 Tapiscia yunnanensis	1.8	1.4	4.7	2.6
	茶梨 Anneslea fragrans	2.2	3.2	2.2	2.6
	红叶木姜子 Litsea rubescens	2.2	3.2	2.2	2.5
	齿叶黄杞 Engelhardtia serrata	1.7	2.4	3.2	2.4

地形异质性对云南普洱季风常绿阔叶林物种多样性的影响

Effects of topographic heterogeneity on species diversity in a monsoon evergreen broad- leaved forest in Puʼer, Yunnan, China

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地形 Topography	植株密度(比例) Plant density (Percentage)
地形 Topography	平均胸径 Average DBH (cm)	1-5 cm	5-10 cm	10-20 cm	20-30 cm	30-40 cm	40-50 cm	≥50 cm
大样地 Big sample plot	6.34	3 213 (62.83)	1 041 (20.35)	589 (11.51)	184 (3.60)	57 (1.12)	20 (0.40)	10 (0.19)
山脊 Ridge	6.33 ± 0.05^a	3 376 (60.20)	1 226 (21.86)	723 (12.89)	210 (3.74)	53 (0.95)	15 (0.27)	5 (0.09)
陡坡 Steep slope	6.28 ± 0.07^a	3 162 (64.31)	950 (19.32)	532 (10.83)	178 (3.61)	63 (1.27)	24 (0.48)	9 (0.18)
缓坡 Less-steep slope	6.38 ± 0.07^a	3 254 (63.56)	1 012 (19.76)	584 (11.41)	180 (3.51)	54 (1.06)	22 (0.42)	14 (0.28)
高谷 High plateau	6.42 ± 0.15^a	3 171 (63.22)	983 (19.61)	576 (11.49)	190 (3.78)	65 (1.30)	21 (0.41)	10 (0.19)
沟谷 Valley	6.35 ± 0.09^a	2 998 (64.40)	941 (20.21)	466 (10.02)	157 (3.37)	57 (1.21)	22 (0.49)	13 (0.30)

参数 Parameter	科数 No. of family	属数 No. of genera	丰富度 Richness	香农威纳指数 Shannon-Wiener index	稀有种 Rare species	稀有物种比例 Rare species ratio (%)	偶见种 Occasional species	偶见种比例 Occasional species ratio (%)
大样地 Big sample plot	79	178	271	-	148	54.6	66	24.4
山脊 Ridge	48	104	145	2.47 ± 0.02^d	63	43.4	44	30.3
陡坡 Steep slope	67	148	198	2.87 ± 0.03^b	78	39.4	56	28.3
缓坡 Less-steep slope	69	147	217	2.81 ± 0.03^b	101	46.5	67	30.9
高谷High plateau	51	103	143	2.69 ± 0.04^c	29	20.3	65	45.5
沟谷 Valley	64	143	200	3.05 ± 0.03^a	74	37.0	62	31.0

物种 Species	山脊 Ridge	缓坡 Less-steep slope	高谷 High plateau	沟谷 Valley
艾胶算盘子 Glochidion lanceolarium	0	0	0	-
枹丝锥 Castanopsis calathiformis	-	0	+	0
茶梨 Anneslea fragrans	0	0	0	0
齿叶黄杞 Engelhardtia serrata	-	0	0	+
短刺锥 Castanopsis echidnocarpa	+	0	0	0
红梗润楠 Machilus rufipes	-	0	0	0
红皮水锦树 Wendlandia tinctoria subsp. intermedia	+	0	0	0
红叶木姜子 Litsea rubescens	0	0	0	0
泥柯 Lithocarpus fenestratus	0	+	0	0
截果柯 Lithocarpus truncatus	0	0	0	0
深绿山龙眼 Helicia nilagirica	+	0	0	0
西桦 Betula alnoides	0	0	0	0
西南木荷 Schima wallichii	0	0	0	0
隐距越桔 Vaccinium exaristatum	+	0	0	0
云南瘿椒树 Tapiscia yunnanensis	0	-	0	0

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