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

doi:10.17520/biods.2025062

Abstract

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

Yi Li, Yanchao Zhao, Litong Chen. Responses of α and β diversity of alpine meadows to warming were modulated strongly by elevations and soil nutrients[J]. Biodiv Sci, 2025, 33(12): 25062.

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URL: https://www.biodiversity-science.net/EN/10.17520/biods.2025062

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

Figures/Tables 8

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

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.

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.

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.

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

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.

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.

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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