高寒草甸α与β多样性对增温的响应受海拔和土壤养分水平调控

doi:10.17520/biods.2025062

生物多样性 ›› 2025, Vol. 33 ›› Issue (12): 25062. DOI: 10.17520/biods.2025062 cstr: 32101.14.biods.2025062

• 研究报告: 植物多样性 • 上一篇下一篇

高寒草甸α与β多样性对增温的响应受海拔和土壤养分水平调控

李燚¹^,², 赵艳超¹, 陈立同¹^,^*()

¹ 中国科学院西北高原生物研究所青海海北高寒草地生态系统国家野外科学观测研究站, 西宁 810008
² 中国科学院大学, 北京 100049

收稿日期:2025-02-18 接受日期:2025-05-21 出版日期:2025-12-20 发布日期:2026-01-09
通讯作者: *E-mail: litong_chen@nwipb.cas.cn
基金资助:
青海省重点研发与转化计划科技国际合作专项(2022-HZ-817)

Responses of α and β diversity of alpine meadows to warming were modulated strongly by elevations and soil nutrients

Yi Li¹^,², Yanchao Zhao¹, Litong Chen¹^,^*()

¹ Qinghai Haibei National Field Research Station of Alpine Grassland Ecosystem, Northwest Institute of Plateau Biology, Chinese Academy of Sciences, Xining 810008, China
² University of Chinese Academy of Sciences, Beijing 100049, China

Received:2025-02-18 Accepted:2025-05-21 Online:2025-12-20 Published:2026-01-09
Supported by:
Key Research and Development and Transformation Program of Qinghai Province-Special Fund for International Scientific and Technological Cooperation(2022-HZ-817)

1. 附录.pdf(2405KB)

摘要/Abstract

摘要：

全球气候变暖会通过改变土壤水分和养分来影响植物群落多样性及其组成。青藏高原是全球气候变暖的敏感区, 且表现出海拔依赖性的增温特点。为准确理解、把握与预测未来全球变暖情景下青藏高原高寒草甸物种多样性的变化趋势与规律, 本文以祁连山北麓高寒草甸为研究对象, 基于低(3,200 m)、中(3,700 m)、高(4,050 m) 3个海拔建立的“增温-养分添加”实验平台, 于2021-2024年在生长季进行了群落物种组成调查, 以探究不同海拔植物群落多样性与组成对增温和养分添加的响应。结果表明: (1) α多样性对增温的响应具有海拔差异性, 即低海拔不变、中海拔降低而高海拔升高。养分添加影响了α多样性对增温的响应, 其主要是通过改变群落不同功能群植物的物种丰富度或相对多度实现的, 且具有海拔差异。具体而言, 在养分添加条件下, 增温降低了低海拔豆科和禾草、中海拔禾草、莎草和杂类草植物的丰富度, 而增加了高海拔禾草和杂类草植物的丰富度及其相对多度。(2)群落β多样性对增温的响应同样依赖于海拔, 并受到养分添加的调节。单独增温显著降低了高海拔群落β多样性, 但是在养分添加的条件下, 增温使得中海拔群落β多样性降低, 而高海拔群落β多样性增加。因此, 在未来全球变暖情景下, 为了更准确地预测生物多样性的响应, 必须要考虑不同地点的基线环境条件, 例如土壤水分和养分状况对生物多样性响应增温的影响。

关键词: α多样性, β多样性, 相对多度, 增温, 氮添加, 磷添加, 海拔, 青藏高原

Abstract

Aims: In this study, we aimed to test whether the responses of the community diversity and composition of alpine meadows were influenced by elevations and soil nutrient availability.

Methods: Based on a warming (open-top chambers, OTCs) and nutrient addition (nitrogen and phosphorus) experiment across elevations (3,200 m, 3,700 m, 4,050 m), plant community surveys were conducted from 2021 to 2024, and α and β diversity metrics and relative abundance of four functional groups were calculated. The linear mixed effects model, permutation multivariance analysis, one- and two-way ANOVAs, and Tukey’s HSD test were used to analyze the effects of warming and nutrient addition on the community α and β diversity.

Results: (1) The effects of warming on α diversity were elevation-dependent, showing no significant effects on it at 3,200 m, and reducing it at 3,700 m, and increasing it at 4,050 m. Meanwhile, nutrients addition also markedly influenced the responses of α diversity to warming by changing species richness and/or relative abundance of different functional groups. Specifically, under nutrient addition conditions, warming reduced the richness of legumes and grasses at 3,200 m, grasses, sedges, and forbs at 3,700 m, while increased the richness and relative abundance of grasses and forbs at 4,050 m. (2) Similarly, the responses of β diversity to warming were affected by elevations, and nutrients addition regulated such responses. For instance, warming significantly reduced β diversity at 4,050 m; by contrast, under nutrient addition conditions, warming reduced it at 3,700 m and increased it at 4,050 m, respectively.

Conclusion: Our findings suggest that elevations and soil nutrients strongly regulate the responses of alpine meadow diversity to warming through species richness and/or the relative abundance of four functional groups. Therefore, to more accurately predict the biodiversity response to global warming in the future, it is essential to consider the baseline environmental conditions (e.g., soil moisture and nutrients).

Key words: α diversity, β diversity, relative abundance, warming, nitrogen addition, phosphorus addition, elevation, Qinghai-Tibet Plateau

李燚, 赵艳超, 陈立同 (2025) 高寒草甸α与β多样性对增温的响应受海拔和土壤养分水平调控. 生物多样性, 33, 25062. DOI: 10.17520/biods.2025062.

Yi Li, Yanchao Zhao, Litong Chen (2025) Responses of α and β diversity of alpine meadows to warming were modulated strongly by elevations and soil nutrients. Biodiversity Science, 33, 25062. DOI: 10.17520/biods.2025062.

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链接本文: https://www.biodiversity-science.net/CN/10.17520/biods.2025062

https://www.biodiversity-science.net/CN/Y2025/V33/I12/25062

图/表 8

表1 利用线性混合效应模型分析海拔、增温、养分、年际及其交互作用对植物群落物种丰富度、Shannon-Wiener多样性指数和反辛普森指数的影响(F值)

Table 1 Effects of elevation, warming, nutrient, year and their interactions on species richness, Shannon-Wiener diversity index, inverse-Simpson index based on linear mixed effects model (F value)

处理 Treatment	分子自由度 numDF	分母自由度 denDF	物种丰富度 Species richness	Shannon-Wiener多样性指数 Shannon-Wiener diversity index	反辛普森指数 Inverse-Simpson index
氮添加 Nitrogen addition (N)	1	257	2.78^*	7.76^**	4.51^*
磷添加 Phosphorus addition (P)	1	257	11.68^**	2.04	1.62
增温 Warming (W)	1	257	1.33	0.23	0.12
海拔 Elevation (E)	2	257	77.04^***	0.23	3.93^*
年际 Year (Y)	3	257	18.23^***	11.78^***	10.24^***
N × P	1	257	0.02	3.79	1.06
N × W	1	257	5.26^*	6.90^**	3.84
P × W	1	257	17.07^***	19.76^***	14.87^***
N × E	2	257	12.92^***	4.92^**	1.52
P × E	2	257	11.55^***	20.81^***	12.78^***
W × E	2	257	6.59^**	6.37^**	7.16^**
N × Y	3	257	6.38^***	1.18	0.95
P × Y	3	257	2.10	0.40	0.06
W × Y	3	257	8.17^***	3.15^*	3.78^*
E × Y	6	257	11.09^***	11.66^***	8.75^***
N × P × W	1	257	1.77	0	1.25
N × P × E	2	257	4.72^*	3.66^*	5.46^**
N × W × E	2	257	1.91	4.58^*	1.83
P × W × E	2	257	4.26^*	2.42	1.34
N × P × Y	3	257	3.73^*	2.35	1.82
N × W × Y	3	257	3.69^*	0.51	0.23
P × W × Y	3	257	3.62^*	2.10	0.86
N × E × Y	6	257	0.62	0.26	0.30
P × E × Y	6	257	0.86	2.32^*	1.58
W × E × Y	6	257	0.80	2.45^*	1.92
N × P × W × E	2	257	4.94^**	5.26^**	4.35^*
N × P × W × Y	3	257	0.55	0.66	0.88
N × P × E × Y	6	257	0.38	0.91	0.71
N × W × E × Y	6	257	0.99	0.19	0.15
P × W × E × Y	6	257	1.24	0.23	0.68
N × P × W × E × Y	6	257	0.93	0.75	0.98

图1 2021-2024年植物群落物种丰富度响应比(a)、Shannon-Wiener多样性指数响应比(b)、反辛普森指数响应比(c)与海拔的关系。CK: 对照; N: 氮添加; P: 磷添加; NP: 氮磷共同添加。母图表示8个处理下响应比与海拔的关系, 子图表示单独增温处理下响应比与海拔的关系。

Fig. 1 Relationship between species richness response ratio (a), Shannon-Wiener diversity index response ratio (b), inverse-Simpson index response ratio (c) and elevation from 2021 to 2024. CK, Control; N, Nitrogen addition; P, Phosphorus addition; NP, Nitrogen and phosphorus co-addition. The parent figures represent the relationship between the response ratio and elevation under eight treatments, the subplot represent the relationship between the response ratio and elevation under warming.

图2 增温和养分添加及其交互作用对不同海拔物种丰富度(a)、Shannon-Wiener多样性指数(b)、反辛普森指数(c)的影响。CK: 对照; N: 氮添加; P: 磷添加; NP: 氮磷共同添加。不同小写和大写字母分别表示不增温和增温条件下4种养分处理对α多样性指标的影响差异显著; 星号代表同一养分条件下增温与不增温处理之间的差异显著性: * P < 0.05; ** P < 0.01; *** P < 0.001。

Fig. 2 Effects of warming, nutrient addition, and their interaction on species richness (a), Shannon-Wiener diversity index (b), and inverse-Simpson index (c) at different elevations. CK, Control; N, Nitrogen addition; P, Phosphorus addition; NP, Nitrogen and phosphorus co-addition. Different lowercase and uppercase letters indicate significant differences in the effects of four nutrient treatments on α diversity indices under non-warming and warming conditions, respectively; asterisks represent the significance of differences between warming and non-warming: * P < 0.05; ** P < 0.01; *** P < 0.001.

图3 增温、养分添加及其交互作用对不同海拔4种植物功能群相对多度的影响。CK: 对照; N: 氮添加; P: 磷添加; NP: 氮磷共同添加。不同小写和大写字母分别表示不增温和增温条件下4种养分处理对不同功能群相对多度的影响差异显著; 星号代表同一养分条件下增温处理与不增温处理之间的差异显著性: * P < 0.05; ** P < 0.01; *** P < 0.001。

Fig. 3 The effects of warming, nutrient addition and their interaction on the relative abundance of plant functional groups at different elevations. CK, Control; N, Nitrogen addition; P, Phosphorus addition; NP, Nitrogen and phosphorus co-addition. Different lowercase and uppercase letters indicate significant differences in the effects of four nutrient treatments on the relative abundance of plant functional groups under non-warming and warming conditions, respectively; asterisks represent the significance of differences between warming and non-warming: * P < 0.05; ** P < 0.01; *** P < 0.001.

表2 海拔、增温、养分、年际及其交互作用对植物群落Bray-Curtis距离(Dbray)和Jaccard距离(Djac)的影响

Table 2 Effect of elevation, warming, nutrient, year and their interactions on Bray-Curtis distance (Dbray) and Jaccard distance (Dbray)

处理 Treatment	df	D_bray		D_bray
处理 Treatment	df	R²	F	R²	F
增温 Warming (W)	1	0.01	7.79^***	0	5.19^**
氮添加 Nitrogen addition (N)	1	0.01	7.72^***	0	5.55^***
磷添加 Phosphorus addition (P)	1	0.03	24.51^***	0.01	14.06^***
海拔 Elevation (E)	2	0.21	98.28^***	0.13	72.78^***
年际 Year (Y)	3	0.15	46.74^***	0.53	428.51^***
W × N	1	0	1.50	0	1.62
W × P	1	0	1.81^*	0	1.56
N × P	1	0	4.32^***	0	1.63
W × E	2	0.01	4.34^***	0	1.17
N × E	2	0.01	3.89^***	0.01	2.94^**
P × E	2	0.02	7.86^***	0	2.00^*
W × Y	3	0.01	3.56^***	0	2.60^*
N × Y	3	0.01	2.31^***	0.01	4.45^***
P × Y	3	0.01	4.24^***	0	3.21^**
E × Y	6	0.13	21.22^***	0.08	21.86^***
W × N × P	1	0	1.60	0	1.34
W × N × E	2	0	0.88	0	0.71
W × P × E	2	0	1.86^**	0	0.83
N × P × E	2	0	1.80^*	0	0.52
W × N × Y	3	0	0.83	0	0.75
W × P × Y	3	0	1.06	0	2.16^*
N × P × Y	3	0	1.11	0	2.01^*
W × E × Y	6	0.02	2.40^***	0	0.99
N × E × Y	6	0.01	1.62^**	0.01	1.83
P × E × Y	6	0.02	2.66^***	0	0.97
W × N × P × E	2	0	1.75^*	0	0.56
W × N × P × Y	3	0	0.72	0	2.10
W × N × E × Y	6	0	0.75	0	0.77
W × P × E × Y	6	0.01	1.16	0	0.38
N × P × E × Y	6	0.01	1.06	0	0.74
W × N × P × E × Y	6	0.01	0.87	0	0.50

图4 2021-2024年增温、养分添加对3个海拔群落组成的非度量多维排序(NMDS)分析结果。CK: 对照; N: 氮添加; P: 磷添加; NP: 氮磷共同添加。蓝色实线表示不增温, 红色实线表示在4种养分处理下同时增温, 红色虚线表示单独增温。

Fig. 4 Results of non-metric multidimensional scaling (NMDS) analysis of the communities composition at different elevations by warming and nutrient addition in 2021-2024. CK, Control; N, Nitrogen addition; P, Phosphorus addition; NP, Nitrogen and phosphorus co-addition. Blue solid line indicates no warming, red solid line indicates warming under nutrient additions, and the red dashed line indicates warming without nutrient additions.

图5 增温、养分添加及其交互作用对不同海拔β多样性的影响, 包括Bray-Curtis距离(a)、Jaccard距离(b)。CK: 对照; N: 氮添加; P: 磷添加; NP: 氮磷共同添加。不同小写和大写字母分别表示不增温和增温条件下4种养分处理对β多样性指标影响的差异显著; 星号代表同一养分条件下增温处理与不增温处理之间的差异显著性: * P < 0.05; ** P < 0.01; *** P < 0.001。

Fig. 5 The effects of warming, nutrient addition and their interaction on β diversity at different elevations, including Bray-Curtis distance (a) and Jaccard distance without species abundance (b). CK, Control; N, Nitrogen addition; P, Phosphorus addition; NP, Nitrogen and phosphorus co-addition. Different lowercase and uppercase letters indicate significant differences in the effects of four nutrient treatments on β diversity indices under non-warming and warming conditions, respectively; asterisks represent the significance of differences between warming and non-warming: * P < 0.05; ** P < 0.01; *** P < 0.001.

表3 增温与年际对3个海拔Bray-Curtis距离的影响

Table 3 Effects of warming and year on the Bray-Curtis distance of three elevations

海拔 Elevation	处理 Treatment	df	R²	F
3,200 m	增温 Warming (W)	1	0.02	2.96^*
	年份 Year (Y)	1	0.24	41.61^***
	W × Y	1	0.02	2.72^*
3,700 m	W	1	0.03	4.70^***
	Y	1	0.19	30.67^***
	W × Y	1	0.01	2.08^*
4,050 m	W	1	0.02	3.05^**
	Y	1	0.21	33.25^***
	W × Y	1	0.01	1.38

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高寒草甸α与β多样性对增温的响应受海拔和土壤养分水平调控

Responses of α and β diversity of alpine meadows to warming were modulated strongly by elevations and soil nutrients

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