养分添加对不同海拔高寒草甸物种多样性及组成的影响

doi:10.17521/cjpe.2025.0137

植物生态学报 ›› 2026, Vol. 50 ›› Issue (4): 870-882.DOI: 10.17521/cjpe.2025.0137 cstr: 32100.14.cjpe.2025.0137

养分添加对不同海拔高寒草甸物种多样性及组成的影响

李燚¹^,³^,^*, 黄欢²^,^*, 赵艳超¹, 陈立同¹^,^**()()

¹ 中国科学院西北高原生物研究所, 西宁 810008
² 青海大学生态环境工程学院, 西宁 810016
³ 中国科学院大学, 北京 100049

收稿日期:2025-04-14 接受日期:2025-10-17 出版日期:2026-04-20 发布日期:2026-06-25
通讯作者: ^**陈立同(litong_chen@nwipb.cas.cn)
作者简介:第一联系人：
^*同等贡献
基金资助:
青海省应用基础研究项目(2020-ZJ-751);第二次青藏高原综合科学考察研究(2019QZKK0302)

Effects of nutrient addition on species diversity and composition of alpine meadows at different altitudes

LI Yi¹^,³^,^*, HUANG Huan²^,^*, ZHAO Yan-Chao¹, CHEN Li-Tong¹^,^**()()

¹ Northwest Institute of Plateau Biology, Chinese Academy of Sciences, Xining 810008, China
² College of Eco-Environmental Engineering, Qinghai University, Xining 810016, China
³ University of Chinese Academy of Sciences, Beijing 100049, China

Received:2025-04-14 Accepted:2025-10-17 Online:2026-04-20 Published:2026-06-25
Contact: ^**CHEN Li-Tong(litong_chen@nwipb.cas.cn)
About author:First author contact:
^*Contributed equally to this work
Supported by:
Applied Basic Research Project in Qinghai Province(2020-ZJ-751);Second Tibetan Plateau Scientific Expedition and Research (STEP) Program(2019QZKK0302)

摘要/Abstract

摘要： 青藏高原土壤速效养分匮乏, 养分添加能通过影响不同的物种或功能群改变群落的物种多样性及组成。由于海拔梯度上具有明显的环境异质性, 群落多样性及组成对养分添加的响应可能存在着垂直分异。因此, 为更好地理解养分添加对不同海拔高寒草甸群落多样性及组成的影响, 2021-2024年, 对低(3 200 m)、中(3 700 m)、高(4 050 m) 3个海拔进行氮添加、磷添加、氮磷共同添加实验, 并在每年生物量顶峰时期进行群落调查。分别计算了物种丰富度、香农-威纳多样性指数、反辛普森指数和毗卢均匀度指数以表征群落的α多样性; 计算了群落的Bray-Curtis相似性和Jaccard相似性以表征群落的β多样性; 同时量化群落不同功能群的物种丰富度及相对多度。结果表明: 低海拔群落α多样性对养分添加不敏感。中海拔群落α多样性对养分添加的响应分异, 表现为在氮添加时降低、磷添加时不变、氮磷共同添加时增加。高海拔的α多样性在养分添加时显著增加。α多样性对养分添加的响应差异主要是由于不同功能群物种的丰富度及多度受养分添加的影响, 特别是部分稀有种的定植和局域灭绝。养分添加降低了低、中海拔群落的相似性, 但导致高海拔群落趋同, 且3个海拔的Jaccard相似性对养分添加的响应更敏感。该研究揭示了青藏高原不同海拔高寒草甸对养分添加的差异化响应, 阐明养分添加对群落多样性指标及物种组成的影响, 为区域生物多样性海拔差异化保护策略制定提供了科学依据。

关键词: 高寒草甸, α多样性, β多样性, 氮添加, 磷添加

Abstract:

Aims Due to the deficiency of soil available nutrients on the Qingzang Plateau, nutrient addition can change the species diversity and composition of alpine meadows by differently affecting species or functional groups. Meanwhile, given an apparent heterogeneity in environmental conditions across elevations, the responses of species diversity and composition to nutrient addition may be vertically differentiated. The aim of this study was to investigate the effects of nutrient addition on the species diversity and composition of alpine meadow communities at different altitudes.

Methods From 2021 to 2024, we added nitrogen (N), phosphorus (P) and both nitrogen and phosphorus at three altitudes (3 200 m, 3 700 m and 4 050 m), and conducted community surveys at the peak of growing season each year. We then calculated α diversity index (species richness, Shannon-Weiner index, Invsimpson index and Pielou evenness index) and β diversity index (Bray-Curtis similarity and Jaccard similarity), and quantified the relative abundance and species richness of different functional groups.

Important findings (1) The α diversity of low elevation communities was insensitive to nutrient addition, but of middle elevation communities decreased with N addition, remained unchanged with P addition, and increased with both N and P additions. By contrast, at high elevations α diversity increased significantly with nutrient addition. The differences in the response of α diversity to nutrient addition are mainly due to the abundance and diversity of plant functional groups being affected differently by nutrient addition, in particular, the colonization and local extinction of some rare species at different elevations. (2) Nutrient addition reduced the community similarity of low and mid-elevations, whereas it resulted in community homogenization at high elevations.

Key words: alpine meadows, α diversity, β diversity, nitrogen addition, phosphorus addition

李燚, 黄欢, 赵艳超, 陈立同. 养分添加对不同海拔高寒草甸物种多样性及组成的影响. 植物生态学报, 2026, 50(4): 870-882. DOI: 10.17521/cjpe.2025.0137

LI Yi, HUANG Huan, ZHAO Yan-Chao, CHEN Li-Tong. Effects of nutrient addition on species diversity and composition of alpine meadows at different altitudes. Chinese Journal of Plant Ecology, 2026, 50(4): 870-882. DOI: 10.17521/cjpe.2025.0137

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https://www.plant-ecology.com/CN/Y2026/V50/I4/870

图/表 8

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处理 Treatment	自由度 df	物种丰富度 Species richness		香农威纳多样性指数 Shannon-Weiner index		毗卢均匀度指数 Pielou evenness index		反辛普森指数 Invsimpson index
处理 Treatment	自由度 df	F	p	F	p	F	p	F	p
氮添加 N	1	2.089	0.150	2.402	0.123	1.360	0.245	1.482	0.225
磷添加 P	1	24.506	<0.001	9.291	0.003	3.163	0.077	6.211	0.014
海拔 E	2	45.611	<0.001	0.734	0.393	1.465	0.228	2.534	0.113
年际 Y	3	3.578	0.060	7.338	0.008	13.822	<0.001	6.557	0.011
N × P	1	0.010	0.921	1.645	0.202	2.429	0.121	0.001	0.980
N × E	2	1.049	0.307	10.891	0.001	10.678	0.001	6.906	0.009
P × E	2	23.048	<0.001	21.883	<0.001	13.700	<0.001	13.238	<0.001
N × Y	3	0.683	0.410	0.441	0.508	0.291	0.591	0.029	0.864
P × Y	3	1.000	0.319	1.129	0.290	0.602	0.439	1.108	0.294
E × Y	6	21.433	<0.001	11.054	0.001	4.915	0.028	6.492	0.012
N × P × E	2	2.190	0.141	3.590	0.060	2.666	0.105	1.072	0.302
N × P × Y	3	4.822	0.030	3.483	0.064	1.798	0.182	0.686	0.409
N × E × Y	6	0.753	0.387	0.644	0.423	0.457	0.500	0	0.995
P × E × Y	6	6.619	0.011	4.422	0.037	2.265	0.134	3.202	0.076
N × P × E × Y	6	0.548	0.460	0.003	0.959	0.088	0.768	0.012	0.914

处理 Treatment	自由度 df	物种丰富度 Species richness		香农威纳多样性指数 Shannon-Weiner index		毗卢均匀度指数 Pielou evenness index		反辛普森指数 Invsimpson index
处理 Treatment	自由度 df	F	p	F	p	F	p	F	p
氮添加 N	1	2.089	0.150	2.402	0.123	1.360	0.245	1.482	0.225
磷添加 P	1	24.506	<0.001	9.291	0.003	3.163	0.077	6.211	0.014
海拔 E	2	45.611	<0.001	0.734	0.393	1.465	0.228	2.534	0.113
年际 Y	3	3.578	0.060	7.338	0.008	13.822	<0.001	6.557	0.011
N × P	1	0.010	0.921	1.645	0.202	2.429	0.121	0.001	0.980
N × E	2	1.049	0.307	10.891	0.001	10.678	0.001	6.906	0.009
P × E	2	23.048	<0.001	21.883	<0.001	13.700	<0.001	13.238	<0.001
N × Y	3	0.683	0.410	0.441	0.508	0.291	0.591	0.029	0.864
P × Y	3	1.000	0.319	1.129	0.290	0.602	0.439	1.108	0.294
E × Y	6	21.433	<0.001	11.054	0.001	4.915	0.028	6.492	0.012
N × P × E	2	2.190	0.141	3.590	0.060	2.666	0.105	1.072	0.302
N × P × Y	3	4.822	0.030	3.483	0.064	1.798	0.182	0.686	0.409
N × E × Y	6	0.753	0.387	0.644	0.423	0.457	0.500	0	0.995
P × E × Y	6	6.619	0.011	4.422	0.037	2.265	0.134	3.202	0.076
N × P × E × Y	6	0.548	0.460	0.003	0.959	0.088	0.768	0.012	0.914

海拔 Elevation (m)	处理 Treatment	自由度 df	物种丰富度 Species richness		香农威纳多样性指数 Shannon-Weiner index		毗卢均匀度指数 Pielou evenness index		反辛普森指数 Invsimpson index
海拔 Elevation (m)	处理 Treatment	自由度 df	F	p	F	p	F	p	F	p
3 200	氮添加 N	1	1.335	0.278	6.157	0.035	6.239	0.034	3.100	0.112
	磷添加 P	1	0.649	0.441	0.882	0.372	0.747	0.410	0.503	0.496
	年际 Y	3	14.429	<0.001	20.549	<0.001	14.866	<0.001	13.169	<0.001
	N × P	1	0.148	0.709	0.157	0.701	0.419	0.534	0.841	0.383
	N × Y	3	0.814	0.372	1.075	0.305	0.968	0.330	0.012	0.915
	P × Y	3	2.890	0.096	1.737	0.194	0.942	0.337	1.076	0.305
	N × P × Y	3	1.748	0.193	2.723	0.106	2.024	0.162	0.819	0.370
3 700	N	1	0.454	0.517	0.554	0.476	0.334	0.578	0.120	0.737
	P	1	2.345	0.160	1.775	0.215	0.853	0.380	2.261	0.167
	Y	3	26.243	<0.001	0.024	0.877	2.663	0.110	1.024	0.317
	N × P	1	3.605	0.090	1.749	0.219	0.436	0.526	4.686	0.059
	N × Y	3	2.203	0.145	0.044	0.834	0.056	0.814	0.006	0.938
	P × Y	3	2.496	0.121	3.276	0.077	2.050	0.159	2.010	0.163
	N × P × Y	3	0.057	0.812	0.051	0.822	0.167	0.684	0.434	0.514
4 050	N	1	3.154	0.109	1.661	0.230	1.144	0.313	0.654	0.439
	P	1	19.193	0.002	9.929	0.012	6.027	0.036	8.982	0.015
	Y	3	6.608	0.014	0.002	0.968	1.049	0.311	0.004	0.951
	N × P	1	5.061	0.051	5.523	0.043	4.868	0.055	4.311	0.068
	N × Y	3	0.280	0.600	0.007	0.936	0.027	0.871	0.016	0.900
	P × Y	3	3.805	0.057	2.569	0.116	1.190	0.281	2.340	0.133
	N × P × Y	3	5.531	0.023	3.721	0.060	1.933	0.171	1.769	0.190

海拔 Elevation (m)	处理 Treatment	自由度 df	物种丰富度 Species richness		香农威纳多样性指数 Shannon-Weiner index		毗卢均匀度指数 Pielou evenness index		反辛普森指数 Invsimpson index
海拔 Elevation (m)	处理 Treatment	自由度 df	F	p	F	p	F	p	F	p
3 200	氮添加 N	1	1.335	0.278	6.157	0.035	6.239	0.034	3.100	0.112
	磷添加 P	1	0.649	0.441	0.882	0.372	0.747	0.410	0.503	0.496
	年际 Y	3	14.429	<0.001	20.549	<0.001	14.866	<0.001	13.169	<0.001
	N × P	1	0.148	0.709	0.157	0.701	0.419	0.534	0.841	0.383
	N × Y	3	0.814	0.372	1.075	0.305	0.968	0.330	0.012	0.915
	P × Y	3	2.890	0.096	1.737	0.194	0.942	0.337	1.076	0.305
	N × P × Y	3	1.748	0.193	2.723	0.106	2.024	0.162	0.819	0.370
3 700	N	1	0.454	0.517	0.554	0.476	0.334	0.578	0.120	0.737
	P	1	2.345	0.160	1.775	0.215	0.853	0.380	2.261	0.167
	Y	3	26.243	<0.001	0.024	0.877	2.663	0.110	1.024	0.317
	N × P	1	3.605	0.090	1.749	0.219	0.436	0.526	4.686	0.059
	N × Y	3	2.203	0.145	0.044	0.834	0.056	0.814	0.006	0.938
	P × Y	3	2.496	0.121	3.276	0.077	2.050	0.159	2.010	0.163
	N × P × Y	3	0.057	0.812	0.051	0.822	0.167	0.684	0.434	0.514
4 050	N	1	3.154	0.109	1.661	0.230	1.144	0.313	0.654	0.439
	P	1	19.193	0.002	9.929	0.012	6.027	0.036	8.982	0.015
	Y	3	6.608	0.014	0.002	0.968	1.049	0.311	0.004	0.951
	N × P	1	5.061	0.051	5.523	0.043	4.868	0.055	4.311	0.068
	N × Y	3	0.280	0.600	0.007	0.936	0.027	0.871	0.016	0.900
	P × Y	3	3.805	0.057	2.569	0.116	1.190	0.281	2.340	0.133
	N × P × Y	3	5.531	0.023	3.721	0.060	1.933	0.171	1.769	0.190

处理 Treatment	自由度 df	Bray-Curtis相似性 Bray-Curtis similarity			Jaccard相似性 Jaccard similarity
处理 Treatment	自由度 df	R²	F	p	R²	F	p
氮添加 N	1	0.01	3.61	0.001	0	1.48	0.166
磷添加 P	1	0.03	14.02	0.001	0.01	7.84	0.001
年际 Y	3	0.14	22.45	0.001	0.13	35.05	0.001
海拔 E	2	0.24	58.13	0.001	0.54	213.51	0.001
N × P	1	0.01	2.81	0.004	0	1.50	0.174
N × Y	3	0.01	1.23	0.171	0	0.81	0.648
P × Y	3	0.02	2.66	0.001	0.01	1.34	0.160
N × E	2	0.01	2.09	0.002	0	1.42	0.154
P × E	2	0.03	6.16	0.001	0.01	2.73	0.007
Y × E	6	0.15	12.20	0.001	0.09	11.75	0.001
N × P × Y	3	0.01	0.80	0.796	0	0.87	0.559
N × P × E	2	0.01	1.72	0.017	0.01	2.14	0.023
N × Y × E	6	0.01	1.06	0.336	0.01	0.76	0.828
P × Y × E	6	0.03	2.10	0.001	0.01	1.04	0.378
N × P × Y × E	6	0.01	0.73	0.966	0	0.44	1.000

养分添加对不同海拔高寒草甸物种多样性及组成的影响

Effects of nutrient addition on species diversity and composition of alpine meadows at different altitudes

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海拔 Elevation (m)	处理 Treatment	自由度 df	Bray-Curtis相似性 Bray-Curtis similarity			Jaccard相似性 Jaccard similarity
海拔 Elevation (m)	处理 Treatment	自由度 df	R²	F	p	R²	F	p
3 200	氮添加 N	1	0.02	2.71	0.031	0.01	1.73	0.125
	磷添加 P	1	0.10	14.73	0.001	0.01	1.81	0.096
	年际 Y	3	0.44	21.74	0.001	0.53	21.95	0.001
	N × P	1	0.01	1.76	0.103	0.01	0.88	0.452
	N × Y	3	0.02	1.01	0.397	0.02	0.76	0.685
	P × Y	3	0.06	3.09	0.003	0.02	0.70	0.723
	N × P × Y	3	0.02	1.11	0.331	0.01	0.62	0.817
3 700	N	1	0.01	1.65	0.081	0.01	0.85	0.565
	P	1	0.06	7.23	0.001	0.03	3.46	0.002
	Y	3	0.38	14.58	0.001	0.44	16.21	0.001
	N × P	1	0.02	2.00	0.039	0.02	2.20	0.026
	N × Y	3	0.02	0.82	0.723	0.02	0.77	0.776
	P × Y	3	0.06	2.20	0.003	0.03	0.98	0.491
	N × P × Y	3	0.02	0.66	0.938	0.01	0.43	0.997
4 050	N	1	0.03	3.40	0.002	0.01	1.92	0.070
	P	1	0.06	6.18	0.001	0.05	7.10	0.001
	Y	3	0.34	12.42	0.001	0.49	21.75	0.001
	N × P	1	0.02	2.39	0.021	0.02	2.24	0.036
	N × Y	3	0.04	1.47	0.079	0.02	0.79	0.733
	P × Y	3	0.05	1.83	0.022	0.04	1.60	0.045
	N × P × Y	3	0.02	0.60	0.959	0.02	0.73	0.822

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