华北地区主要灌丛群落物种组成及系统发育结构特征

doi:10.17521/cjpe.2018.0173

植物生态学报 ›› 2019, Vol. 43 ›› Issue (9): 793-805.DOI: 10.17521/cjpe.2018.0173

华北地区主要灌丛群落物种组成及系统发育结构特征

柴永福¹,许金石¹,刘鸿雁²,刘全儒³,郑成洋²,康慕谊^4,⁵,梁存柱⁶,王仁卿⁷,高贤明⁸,张峰⁹,福臣¹⁰,刘晓¹,岳明^1,^*()

¹西北大学西部资源生物与现代生物技术教育部重点实验室, 西安 710069
²北京大学城市与环境学院, 北京大学生态研究中心, 地表过程分析与模拟教育部重点实验室, 北京 100871
³北京师范大学生命科学学院, 北京 100875
⁴北京师范大学地表过程与资源生态国家重点实验室, 北京 100875
⁵北京师范大学地理科学学部自然资源学院, 北京 100875
⁶内蒙古大学生态与环境学院, 蒙古高原生态学与资源利用教育部重点实验室, 呼和浩特 010021
⁷山东大学生命科学学院, 济南 250100
⁸中国科学院植物研究所植被与环境变化国家重点实验室, 北京 100093
⁹山西大学黄土高原研究所, 太原 030006
¹⁰南开大学生命科学学院, 天津 300071

收稿日期:2018-07-30 接受日期:2018-10-29 出版日期:2019-09-20 发布日期:2020-01-03
通讯作者: 岳明
基金资助:
国家科技基础性工作专项(2011FY110300);国家自然科学基金(31700348)

Species composition and phylogenetic structure of major shrublands in North China

CHAI Yong-Fu¹,XU Jin-Shi¹,LIU Hong-Yan²,LIU Quan-Ru³,ZHENG Cheng-Yang²,KANG Mu-Yi^4,⁵,LIANG Cun-Zhu⁶,WANG Ren-Qing⁷,GAO Xian-Ming⁸,ZHANG Feng⁹,SHI Fu-Chen¹⁰,LIU Xiao¹,YUE Ming^1,^*()

¹Key Laboratory of Resource Biology and Biotechnology in Western China, Ministry of Education, Northwest University, Xi’an 710069, China
²Institute of Ecology, College of Urban and Environmental Sciences, Laboratory for Earth Surface Processes of the Ministry of Education, Peking University, Beijing 100871, China
³College of Life Sciences, Beijing Normal University, Beijing 100875, China
⁴State Key Laboratory of Earth Surface Processes and Resource Ecology, Beijing Normal University, Beijing 100875, China
⁵College of Resources Science & Technology, Faculty of Geographical Science, Beijing Normal University, Beijing 100875, China
⁶College of Ecology and Environment, Inner Mongolia University, Key Laboratory of Ecology and Resource Utilization of the Mongolian Plateau, Ministry of Education, Hohhot 010021, China
⁷School of Life Sciences, Shandong University, Jinan 250100, China
⁸State Key Laboratory of Vegetation and Environmental Change, Institute of Botany, Chinese Academy of Sciences, Beijing 100093, China
⁹Institute of Loess Plateau, Shanxi University, Taiyuan 030006, China
¹⁰College of Life Sciences, Nankai University, Tianjin 300071, China

Received:2018-07-30 Accepted:2018-10-29 Online:2019-09-20 Published:2020-01-03
Contact: YUE Ming
Supported by:
Supported by the National Basic Work of Science and Technology of China(2011FY110300);and the National Natural Science Foundation of China(31700348)

摘要/Abstract

摘要：

灌丛在中国北方广泛分布, 研究其物种组成特征及构建机制对植被恢复有重要意义。群落系统发育结构能有效反映群落组成的生态过程, 对揭示群落构建机制具有重要作用。该研究利用华北地区自然植物群落资源综合考察数据库的灌丛数据, 分析了中国华北地区主要灌丛群落的分布情况、物种组成及其系统发育结构特征。同时, 结合WorldClim数据库中的气候因子利用逐步回归和一般线性模型分析了气温和降水对灌丛不同垂直结构层物种系统发育结构特征的影响。经统计, 共调查木本植物75科207属570种; 草本植物99科491属1 221种。按植被型和群系类型分类后, 共有5种植被型, 195个群系, 其中分布最多的群系类型为虎榛子(Ostryopsis davidiana)灌丛、荆条(Vitex negundo var. heterophylla)灌丛、荆条+酸枣(Ziziphus jujuba)灌丛和沙棘(Hippophae rhamnoides)灌丛。整体上, 草本层的物种丰富度高于灌木层。草本层的系统发育结构表现为发散模式, 而灌木层的系统发育结构表现为聚集模式, 且不同植被型灌丛的系统发育结构不同。气候因子的变化对灌木层和草本层系统发育结构都表现出了明显的影响, 但对灌木层的影响要强于对草本层的影响, 且气候因子和植被型、群系类型都有明显的交互作用。分析结果表明, 区域尺度上气温和降水的变化会影响灌丛群落不同结构层的系统发育结构特征, 且对不同的植被型和群系类型的影响不同。

关键词: 华北地区, 灌丛, 系统发育结构, 群落构建, 环境过滤

Abstract:

Aims Shrublands are widely distributed in the North China. Understanding species composition and community assembly of the shrublands has important implications for promoting vegetation restoration in this area. Community phylogenetic structure can reflect community assembly process. In the present study, we explored the distribution patterns, species composition and phylogenetic structure of major shrubland types from North China, and then analyzed the effects of temperature and precipitation on these patterns with the help of the WorldClim dataset.
Methods A total 2 331 plots were setup to survey species composition and phylogenetic structure of major shrubland types in North China. A multiple stepwise regression was used to determine the effects of climatic factors on community phylogenetic structure, and a generalize linear model was used to test the interaction of environmental factors and formation types or vegetation types.
Important findings A total of 570 woody species belonging to 207 genera from 75 families, together with 1 221 herb species belonging to 491 genera from 99 families were recorded during the investigation. Five vegetation types and 195 formation types were identified, and the major formation types were Ostryopsis davidiana formation, Vitex negundo var. heterophylla, Vitex negundo var. heterophylla + Hippophae rhamnoides and Hippophae rhamnoides formation. The species richness of herb layer was higher than that of shrub layer. The phylogenetic structure was over-dispersed for herb layer and was convergent for shrub layer. The phylogenetic structure differed remarkable among different vegetation types. The phylogenetic structures of both shrub and herb layers were significantly correlated with climatic factors, and the interaction of climate factors and vegetation types or formation types as well. Our results indicates that large scale variation of climatic factors regulate community phylogenetic structure of different layers in shrub community, and the effects depend on vegetation types and formation types.

Key words: North China, shrubland, phylogenetic structure, community assembly, environmental filtering

柴永福, 许金石, 刘鸿雁, 刘全儒, 郑成洋, 康慕谊, 梁存柱, 王仁卿, 高贤明, 张峰, 福臣, 刘晓, 岳明. 华北地区主要灌丛群落物种组成及系统发育结构特征. 植物生态学报, 2019, 43(9): 793-805. DOI: 10.17521/cjpe.2018.0173

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. Species composition and phylogenetic structure of major shrublands in North China. Chinese Journal of Plant Ecology, 2019, 43(9): 793-805. DOI: 10.17521/cjpe.2018.0173

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2018.0173

https://www.plant-ecology.com/CN/Y2019/V43/I9/793

图/表 11

图1 华北地区灌丛调查样方分布图。

Fig. 1 Distribution of survey plots for shrubland in North China.

表1 华北地区灌丛研究中采用的环境因子

Table 1 Environmental variables used in the study of major shrublands in North China

编码 Code	环境因子 Environmental variables	缩写 Abbreviation
Bio1	年平均气温 Annual mean air temperature	AMT
Bio2	每月最高气温与最低气温差值的平均值 Mean diurnal air temperature range	MDTR
Bio3	等温性 Isothermality	ISO
Bio4	季节性气温变异 Air temperature seasonality	TS
Bio5	最热月的最高气温 Max temperature of the warmest month	MTWM
Bio6	最冷月的最低气温 Min temperature of the coldest month	MTCM
Bio7	气温的年较差 Temperature annual range	TAR
Bio8	最湿季度的平均气温 Mean temperature of the wettest quarter	MTWEQ
Bio9	最干季度的平均气温 Mean temperature of the driest quarter	MTDQ
Bio10	最热季度的平均气温 Mean temperature of the warmest quarter	MTWAQ
Bio11	最冷季度的平均气温 Mean temperature of the coldest quarter	MTCQ
Bio12	年降水量 Annual precipitation	AP
Bio13	最湿月的降水量 Precipitation of the wettest month	PWM
Bio14	最干月的降水量 Precipitation of the driest month	PDM
Bio15	降水季节性变化 Precipitation seasonality	PS
Bio16	最湿季度的降水量 Precipitation of the wettest quarter	PWEQ
Bio17	最干季度的降水量 Precipitation of the driest quarter	PDQ
Bio18	最热季度的降水量 Precipitation of the warmest quarter	PWAQ
Bio19	最冷季度的降水量 Precipitation of the coldest quarter	PCQ
Alt	海拔 Altitude	Alt

图2 华北地区灌丛的主要植被类型(A)和群系类型(B)。植被类型: a, 竹灌丛; b, 落叶灌丛; c, 灌丛荒漠; d, 常绿针叶灌丛; e, 常绿革叶灌丛。群系类型: 1, 虎榛子灌丛; 2, 荆条灌丛; 3, 荆条+酸枣灌丛; 4, 沙棘灌丛; 5, 山杏灌丛; 6, 黄刺玫灌丛; 7, 绣线菊灌丛; 8, 锦鸡儿灌丛; 9, 柠条锦鸡儿灌丛; 10, 酸枣灌丛; 11, 红砂荒漠; 12, 柽柳灌丛; 13, 野皂荚灌丛; 14, 白刺花灌丛; 15, 黄蔷薇灌丛。

Fig. 2 Major vegetation types (A) and formation types (B) of shrubland in North China. Vegetation types: a, bamboo thickets; b, deciduous thickets; c, desert thickets; d, evergreen needle-leaved thickets; e, evergreen leather-leaved thickets. Formation types: 1, Ostryopsis davidiana shrub; 2, Vitex negundo var. heterophylla shrub; 3, Vitex negundo var. heterophylla + Ziziphus jujube shrub; 4, Hippophae rhamnoides shrub; 5, Armeniaca sibirica shrub; 6, Rosa xanthine shrub; 7, Spiraea salicifolia shrub; 8, Caragana sinica shrub; 9, Caragana korshinskii shrub; 10, Ziziphus jujube shrub; 11, Reaumuria soongarica shrub; 12, Tamarix chinensis shrub; 13, Gleditsia microphylla shrub; 14, Sophora davidii shrub; 15, Rosa hugonis shrub.

图3 华北地区灌丛主要植被型在各地区的分布。BEJ, 北京; TIJ, 天津; HEB, 河北; HEN, 河南; SHX, 陕西; SAX, 山西; SHD, 山东; NIX, 宁夏; NMG, 内蒙古; GSU, 甘肃。

Fig. 3 Distribution of major vegetation types of shrubland in North China. BEJ, Beijing; TIJ, Tianjin; HEB, Hebei; HEN, Henan; SHX, Shaanxi; SAX, Shanxi; SHD, Shandong; NIX, Ningxia; NMG, Nei Mongol; GSU, Gansu.

表2 华北地区主要灌丛群落中物种数前10的科

Table 2 Top ten families with the most species of major shrubland in North China

序号 Rank	木本植物 Woody species		草本植物 Herbaceous species
序号 Rank	科名 Family	物种数 No. of species	科名 Family	物种数 No. of species
1	蔷薇科 Rosaceae	101	菊科 Asteraceae	219
2	豆科 Fabaceae	62	禾本科 Poaceae	138
3	忍冬科 Caprifoliaceae	24	豆科 Fabaceae	101
4	鼠李科 Rhamnaceae	23	毛茛科 Ranunculaceae	60
5	木犀科 Oleaceae	20	唇形科 Lamiaceae	55
6	杨柳科 Salicaceae	17	蔷薇科 Rosaceae	48
7	壳斗科 Fagaceae	16	苋科 Amaranthaceae	40
8	卫矛科 Celastraceae	14	伞形科 Apiaceae	37
9	榆科 Ulmaceae	12	十字花科 Brassicaceae	27
10	小檗科 Berberidaceae	11	蓼科 Polygonaceae	26
	合计 Total	300	合计 Total	751

表3 华北地区主要灌丛群落中物种频度前10的物种

Table 3 Top ten species with the most frequency of major shrubland in North China

序号 Rank	木本植物 Woody species		草本植物 Herbaceous species
序号 Rank	种名 Species	相对频度 Relative frequency	种名 Species	相对频度 Relative frequency
1	荆条 Vitex negundo var. heterophylla	9.53	白莲蒿 Artemisia sacrorum	2.35
2	酸枣 Ziziphus jujuba var. spinosa	4.76	大披针薹草 Carex lanceolata	2.24
3	虎榛子 Ostryopsis davidiana	3.98	狗尾草 Setaria viridis	1.74
4	山杏 Armeniaca sibirica	3.12	北京隐子草 Cleistogenes hancei	1.52
5	土庄绣线菊 Spiraea pubescens	2.78	委陵菜 Potentilla chinensis	1.52
6	沙棘 Hippophae rhamnoides	2.76	小红菊 Dendranthema chanetii	1.45
7	三裂绣线菊 Spiraea trilobata	2.71	阿尔泰狗娃花 Heteropappus altaicus	1.37
8	黄刺玫 Rosa xanthina	2.50	茜草 Rubia cordifolia	1.31
9	二色胡枝子 Lespedeza bicolor	2.16	山蒿 Artemisia brachyloba	1.31
10	绣线菊 Spiraea salicifolia	1.83	野青茅 Deyeuxia arundinacea	1.29
	合计 Total	36.13	合计 Total	16.10

图4 华北地区灌丛灌木层和草本层的物种丰富度(平均值±标准误差)。A, 所有群落草本层和灌木层物种丰富度之间的差异。B, 同一结构层不同植被型之间的差异。用t检验分析整体的差异, 用单因素方差分析检验不同植被型之间的差异。柱形上方不同字母代表相应处理之间在5%水平有显著性差异。a, 竹灌丛; b, 落叶灌丛; c, 灌丛荒漠; d, 常绿针叶灌丛; e, 常绿革叶灌丛。

Fig. 4 Species richness of shrub species and herb species of shrubland in North China (mean ± SE). A, Differences of species richness between herb and shrub species for all communities. B, Differences among vegetation types. Differences between herb and shrub species for all communities was texted by t test and differences among vegetation types were tested using a one-way ANOVA with a Tukey post hoc test of significance. Significance different at p < 0.05 was indicated by different letters. a, bamboo thickets; b, deciduous thickets; c, desert thickets; d, evergreen needle-leaved thickets; e, evergreen leather-leaved thickets.

图5 华北地区灌丛灌木层和草本层物种的净谱系亲缘关系指数(NRI)(平均值±标准误差)。A, 所有群落草本层和灌木层物种丰富度之间的差异。B, 同一结构层不同植被型之间的差异。用t检验分析整体的差异, 用单因素方差分析检验不同植被型之间的差异。柱形上方不同字母代表相应处理之间在5%水平有显著性差异。*表示NRI的均值和零值有显著差异(p < 0.05)。a, 竹灌丛; b, 落叶灌丛; c, 灌丛荒漠; d, 常绿针叶灌丛; e, 常绿革叶灌丛。

Fig. 5 Net relatedness index (NRI) of shrub species and herb species of shrubland in North China (mean ± SE). A, Differences of species richness between herb and shrub species for all communities. B, Differences among vegetation types. Differences between herb and shrub species for all communities was texted by t test and differences among vegetation types were tested using a one-way ANOVA with a Tukey post hoc test of significance. Significance different at p < 0.05 was indicated by different letters. * indicates that the mean of NRI was different with zero. a, bamboo thickets; b, deciduous thickets; c, desert thickets; d, evergreen needle-leaved thickets; e, evergreen leather-leaved thickets.

表4 环境因子对华北地区灌丛群落灌木层和草本层物种净谱系亲缘关系指数(NRI)的影响

Table 4 Effects of environmental factors on net relatedness index (NRI) of shrub layer and herb layer species of shrubland in North China

结构层 Structural layer	R²	p	显著环境因子 Significant environmental factors	标准化回归系数 Beta regression coefficient	t	p
灌木层 Shrub layer	0.18	<0.05	MTWEQ	6.81	7.91	<0.01
			PWEQ	5.08	5.16	<0.01
			MDTR	2.39	2.19	0.03
			MTCQ	1.74	2.73	0.01
			Alt	0.22	2.44	0.01
			ISO	-0.89	-2.71	0.01
			PWAQ	-5.45	-4.64	<0.01
			MTWM	-9.54	-8.32	<0.01
草本层 Herb layer	0.05	<0.05	TS	0.12	2.46	0.01
			MTWEQ	0.12	2.56	0.01
			PDM	0.12	2.84	<0.01
			PS	0.08	2.39	0.02

表5 华北地区植被型(VT)和环境因子对净谱系亲缘关系指数的交互作用

Table 5 Interactions of environmental factors and vegetation types on NRI in North China

交互作用	灌木 Shrub
交互作用	d.f.	F	p
MDTR × VT	4	39.15	<0.01
ISO × VT	4	36.33	<0.01
MTWM × VT	4	35.02	<0.01
MTWEQ × VT	4	36.63	<0.01
MTCQ × VT	4	42.16	<0.01
PWEQ × VT	4	28.76	<0.01
PWAQ × VT	4	28.70	<0.01
Alt × VT	4	39.59	<0.01
TS × VT	4	5.69	<0.01
MTWEQ × VT	4	4.33	<0.01
PDM × VT	4	1.86	<0.01
PS × VT	4	12.6	<0.01

表6 华北地区群系类型(FT)和环境因子对净谱系亲缘关系指数的交互作用

Table 6 Interactions of environmental factors and formation types on NRI in North China

交互作用	灌木 Shrub
交互作用	d.f.	F	p
MDTR × FT	194	4.15	<0.01
ISO × FT	194	4.12	<0.01
MTWM × FT	194	4.12	<0.01
MTWEQ × FT	194	4.09	<0.01
MTCQ × FT	194	4.02	<0.01
PWEQ × FT	194	3.95	<0.01
PWAQ × FT	194	3.94	<0.01
Alt × FT	194	4.17	<0.01
TS × FT	194	2.60	<0.01
MTWEQ × FT	194	2.54	<0.01
PDM × FT	194	2.33	<0.01
PS × FT	194	2.63	<0.01

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华北地区主要灌丛群落物种组成及系统发育结构特征

Species composition and phylogenetic structure of major shrublands in North China

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