Chin J Plant Ecol ›› 2026, Vol. 50 ›› Issue (4): 870-882.DOI: 10.17521/cjpe.2025.0137 cstr: 32100.14.cjpe.2025.0137
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LI Yi1,3,*, HUANG Huan2,*, ZHAO Yan-Chao1, CHEN Li-Tong1,**(
)(
)
Received:2025-04-14
Accepted:2025-10-17
Online:2026-04-20
Published:2026-06-25
Contact:
CHEN Li-Tong
About author:First author contact:*Contributed equally to this work
Supported by: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[J]. Chin J Plant Ecol, 2026, 50(4): 870-882.
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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 | ||||
|---|---|---|---|---|---|---|---|---|---|
| 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 |
Table 1 Results of linear mixed model on the effects of nitrogen addition, phosphorus addition, elevation, year and their interactions on taxonomic α diversity
| 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 | ||||
|---|---|---|---|---|---|---|---|---|---|
| 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 | ||||
|---|---|---|---|---|---|---|---|---|---|---|
| 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 | |
Table 2 Results of linear mixed model on the effects of nitrogen addition, phosphorus addition, year and their interactions on taxonomic alpha diversity at different elevations
| 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 | ||||
|---|---|---|---|---|---|---|---|---|---|---|
| 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 | |
Fig. 1 Effects of no nutrient addition (CK), nitrogen addition (N), phosphorus addition (P), and combined nitrogen and phosphorus addition (NP) on the α diversity of plant communities at three altitudes from 2021 to 2024 (mean ± SE). Different lowercase letters indicate the significance of differences in the effects of the four nutrient treatments on α diversity indices (p < 0.05).
Fig. 2 Effects of no nutrient addition (CK), nitrogen addition (N), phosphorus addition (P), and combined nitrogen and phosphorus addition (NP) on the relative abundance of plant functional groups at three altitudes from 2021 to 2024 (mean ± SE). Different lowercase letters indicate the significance of differences in the effects of the four nutrient treatments on the relative abundance of plant functional groups (p < 0.05).
Fig. 3 Effects of no nutrient addition (CK), nitrogen addition (N), phosphorus addition (P), and combined nitrogen and phosphorus addition (NP) on species richness of plant functional groups at three altitudes from 2021 to 2024 (mean ± SE). Different lowercase letters indicate the significance of differences in the effects of the four nutrient treatments on species richness of plant functional groups (p < 0.05).
| 处理 Treatment | 自由度 df | Bray-Curtis相似性 Bray-Curtis similarity | Jaccard相似性 Jaccard similarity | ||||
|---|---|---|---|---|---|---|---|
| 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 |
Table 3 Results of PERMANOVA on the effect of elevation, fertilizer, year and their interactions on community composition
| 处理 Treatment | 自由度 df | Bray-Curtis相似性 Bray-Curtis similarity | Jaccard相似性 Jaccard similarity | ||||
|---|---|---|---|---|---|---|---|
| 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 |
| 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 | ||||
|---|---|---|---|---|---|---|---|---|
| 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 | |
Table 4 Results of PERMANOVA on the effect of fertilizer, year and their interactions on community composition of three elevation
| 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 | ||||
|---|---|---|---|---|---|---|---|---|
| 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 | |
Fig. 4 Effects of no nutrient addition (CK), nitrogen addition (N), phosphorus addition (P), and combined nitrogen and phosphorus addition (NP) on β-diversity at three altitudes from 2021 to 2024 (mean ± SE). Different lowercase letters indicate the significance of differences in the effects of the four nutrient treatments on β-diversity of plant functional groups (p < 0.05).
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