氮和牛粪输入对典型草原植物种-面积关系的影响

doi:10.17520/biods.2025251

生物多样性 ›› 2025, Vol. 33 ›› Issue (12): 25251. DOI: 10.17520/biods.2025251 cstr: 32101.14.biods.2025251

• 研究报告: 植物多样性 • 上一篇下一篇

氮和牛粪输入对典型草原植物种-面积关系的影响

卢炜煜¹^,²^,³, 陈旭¹^,³, 张日谦¹^,³, 张云海¹^,³, 李昭⁴^,^*()

¹ 中国科学院植物研究所植被与环境变化重点实验室, 北京 100093
² 中南安全环境技术研究院股份有限公司, 武汉 430061
³ 中国科学院大学, 北京 100049
⁴ 国家林业和草原局国家公园(自然保护地)发展中心, 北京 100714

收稿日期:2025-07-02 接受日期:2025-11-20 出版日期:2025-12-20 发布日期:2026-01-09
通讯作者: *E-mail: 48612673@qq.com
基金资助:
国家重点研发计划(2022YFF1302800);国家自然科学基金(32122055)

Effects of nitrogen and cow dung input on plant species-area relationship in a typical grassland

Weiyu Lu¹^,²^,³, Xu Chen¹^,³, Riqian Zhang¹^,³, Yunhai Zhang¹^,³, Zhao Li⁴^,^*()

¹ Key Laboratory of Vegetation and Environmental Change, Institute of Botany, Chinese Academy of Sciences, Beijing 100093, China
² Central Southern Safety and Environment Technology Institute Co., Ltd., Wuhan 430061, China
³ University of Chinese Academy of Sciences, Beijing 100049, China
⁴ Development Centre of National Park (Natural Protected Areas), National Forestry and Grassland Administration, Beijing 100714, China

Received:2025-07-02 Accepted:2025-11-20 Online:2025-12-20 Published:2026-01-09
Supported by:
National Key Research and Development Program of China(2022YFF1302800);National Natural Science Foundation of China(32122055)

摘要/Abstract

摘要：

氮富集通常会降低样方尺度的植物多样性, 而牛粪输入被认为能维持天然草原植物多样性。但氮沉降增加和牛粪周转时间延长等复合情景如何影响草原植物多样性及种-面积关系尚不明确。本研究依托中国科学院内蒙古草原生态系统定位研究站, 于2022年在附近典型草原建立了氮和牛粪添加实验, 并于实验第3年即2024年8月测定了每个实验小区1-8 m²植物物种数及表层土壤理化特性指标。结果表明: 围封显著降低了18.9%的空间尺度平均植物物种数; 显著降低了种-面积关系截距(单位面积植物多样性), 但并未改变物种共存最小面积和种-面积关系斜率(群落组成空间异质性)。氮和牛粪添加均未显著改变植物种-面积关系的截距、斜率和物种共存最小面积。结构方程模型结果显示, 围封通过降低土壤pH值使种-面积关系斜率增加, 但通过增加植物地上生产力降低了种-面积关系截距; 牛粪添加通过增加土壤铵态氮含量同时降低种-面积关系斜率和物种共存最小面积。本文结果表明, 典型草原生态系统牛粪输入会带来适量的养分输入, 有利于物种共存。因此, 应在保障草畜平衡的基础上评估牛粪输入对草原生态系统功能和畜牧生产的协同效益。

关键词: 草地, 植物多样性, 牛粪, 氮沉降, 种-面积关系

Abstract

Aims: Nitrogen (N) enrichment can reduce grassland plant diversity, while dung decomposition and nutrient release are important for maintaining plant diversity in native grassland ecosystems. Grasslands have been subjected to both atmospheric N deposition and dung inputs. However, it remains unclear how these factors affect plant richness and species-area relationship (SAR).

Methods: In 2022, we conducted a field experiment at the Inner Mongolia Grassland Ecosystem Research Station, Chinese Academy of Sciences. A split-plot completely randomized design was used, with enclosure and grazing as the two main effects. Within each main effect, four treatments were applied: control, cow dung addition, N addition, combined N and cow dung addition. In August 2024, species richness was recorded at four spatial scales (1 m², 2 m², 4 m², and 8 m²). At the same time, aboveground net primary productivity (ANPP) and relevant soil physicochemical indicators were measured.

Results: We found that enclosure significantly reduced the mean plant species richness at multiple spatial scales by 18.9%. Enclosure significantly reduced the intercept of the species-area relationship (i.e., plant diversity per unit area), but did not affect the minimum area for species coexistence or the slope (i.e., spatial heterogeneity of community composition). Neither N nor cow dung addition significantly changed the intercept, or slope of the species-area relationship. Results from the structural equation model showed that enclosure had a positive effect on the slope by lowering soil pH. It also exerted a negative effect on the intercept by increasing aboveground net primary productivity. Cow dung addition had a negative effect on both the slope and intercept of the species-area relationship by increasing soil ammonium nitrogen content.

Conclusions: Our findings suggest that dung inputs in typical steppe ecosystems can facilitate species coexistence through moderate nutrient supplementation, highlighting the importance of evaluating the ecological, productive, and livelihood benefits of dung coverage under the premise of grass-livestock balance.

Key words: grassland, plant diversity, cow dung, nitrogen deposition, species-area relationship

卢炜煜, 陈旭, 张日谦, 张云海, 李昭 (2025) 氮和牛粪输入对典型草原植物种-面积关系的影响. 生物多样性, 33, 25251. DOI: 10.17520/biods.2025251.

Weiyu Lu, Xu Chen, Riqian Zhang, Yunhai Zhang, Zhao Li (2025) Effects of nitrogen and cow dung input on plant species-area relationship in a typical grassland. Biodiversity Science, 33, 25251. DOI: 10.17520/biods.2025251.

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链接本文: https://www.biodiversity-science.net/CN/10.17520/biods.2025251

https://www.biodiversity-science.net/CN/Y2025/V33/I12/25251

图/表 6

图1 实验样地及嵌套取样概览图。本实验以放牧和围封为主效应, 氮和牛粪添加为副效应。图中编号1-8为进行植物物种调查的8个1 m × 1 m小样方。种-面积关系包括4个调查面积: 1 m2 (红色)、2 m2 (黄色)、4 m2 (绿色)和8 m2 (蓝色)。

Fig. 1 Overview of experimental plots and nested sampling. This experiment primarily focuses on grazing and enclosure as the main effects, with N and cow dung addition as the secondary effects. The numbers 1-8 in the figure refer to eight 1 m × 1 m small plots for plant species sampling. The species- area relationship includes four sampling areas: 1 m2 (red), 2 m2 (yellow), 4 m2 (green), and 8 m2 (blue).

图2 氮和牛粪添加对植物种-面积关系的影响。线性拟合为围封和放牧条件下物种数与调查面积的相关性。共有1 m2、2 m2、4 m2和8 m2 4个调查面积。4种颜色对应4种处理: 对照、氮添加、牛粪添加及氮和牛粪同时添加。拟合为实线, 代表两者具有显著相关关系(P < 0.05)。阴影部分表示95%置信区间。

Fig. 2 Effects of N and cow dung addition on the plant species-area relationship. Linear regressions illustrate the relationship between species number and sampling area under both enclosure and grazing conditions. Four spatial scales were examined: 1 m2, 2 m2, 4 m2, and 8 m2. Four colors represent four treatments: control, N addition, cow dung addition, and combined N and cow dung addition. Solid lines indicate significant relationships (P < 0.05), and shaded areas represent 95% confidence intervals of the fitted lines.

表1 围封和放牧条件下氮和牛粪添加对植物群落和土壤理化性质影响的方差分析结果(F值)

Table 1 Variance analysis results of the effects of N and cow dung addition on plant communities and soil physicochemical properties under enclosure and grazing conditions (F-value). Amin, Minimum area for species coexistence; SWC, Soil water content. SIN, Soil inorganic nitrogen.

	df	物种数 Species number	斜率 Slope	截距 Intercept	物种共存最小面积 A_min	地上净初级生产力 Aboveground net primary production	pH	土壤含水量 SWC	铵态氮含量 NH+ 4-N	硝态氮含量 NO- 3-N	土壤无机氮含量 SIN
面积 Area	1, 191	954.7^***	-	-	-	-	-	-	-	-	-
围封 Enclosure (E)	1, 56	30.9^***	1.7	28.3^***	2.0	102.7^***	33.4^***	31.5^***	3.1	0.0	0.0
氮添加 N addition (N)	1, 56	0.1	0.1	0.0	0.0	0.0	0.2	0.4	0.7	0.8	0.5
牛粪添加 Cow dung addition (D)	1, 56	0.2	0.9	0.7	2.8	2.3	0.6	0.2	5.5^*	20.7^***	24.0^***
围封 × 氮添加 E × N	1, 56	0.2	0.0	0.3	0.3	4.4^***	0.1	1.3	1.4	0.1	0.2
围封 × 牛粪添加 E × D	1, 56	0.0	0.4	0.1	0.2	0.1	1.7	0.2	2.5	0.1	0.4
氮添加 × 牛粪添加 N × D	1, 56	3.9	0.1	3.9	0.8	1.7	1.2	0.4	1.7	0.5	0.9
E × N × D	1, 56	1.7	1.6	3.6	2.9	8.0^**	0.2	1.3	1.6	1.5	0.9

图3 氮和牛粪添加对植物种-面积关系的斜率(a)、截距(b)和物种共存最小面积(c)的影响(平均值 ± 标准误差)。不同小写字母表示围封处理间差异显著(P < 0.05), 不同大写字母表示放牧处理间差异显著(P < 0.05)。星号表示围封和放牧条件下同一处理间差异显著: * P < 0.05; ** P < 0.01; NS, 不显著。

Fig. 3 Effects of N and cow dung addition on the slope (a), intercept (b), and minimum area for species coexistence (Amin) (c) of the plant species-area relationship (mean ± SE). Different lowercase letters indicate significant differences between treatments during enclosure (P < 0.05), while different uppercase letters indicate significant differences between treatments during grazing (P < 0.05). Asterisks denote significant differences between enclosure and grazing within the same treatment: * P < 0.05; ** P < 0.01; NS, not significant.

图4 氮和牛粪添加对植物地上净初级生产力(a)、土壤pH (b)、土壤含水量(c)、土壤铵态氮含量(d)、土壤硝态氮含量(e)和土壤无机氮含量(f)的影响(平均值 ± 标准误差)。不同小写字母表示围封处理间差异显著(P < 0.05), 不同大写字母表示放牧处理间差异显著(P < 0.05)。星号表示围封和放牧条件下同一处理间差异显著: * P < 0.05; ** P < 0.01; *** P < 0.001; NS, 不显著。

Fig. 4 Effects of cow dung and nitrogen addition on aboveground net primary productivity (a), soil pH (b), soil water content (c), soil ammonium nitrogen (NH4+-N) content (d), soil nitrate nitrogen (NO3--N) content (e), and soil inorganic nitrogen (f) (mean ± SE). Different lowercase letters indicate significant differences between treatments during enclosure (P < 0.05), while different uppercase letters indicate significant differences between treatments during grazing (P < 0.05). Asterisks denote significant differences between enclosure and grazing within the same treatment: * P < 0.05; ** P < 0.01; *** P < 0.001; NS, not significant.

图5 影响种-面积关系的斜率(a)、截距(b)和物种共存最小面积(Amin) (c)的结构方程模型。黑线表示正相关, 红线表示负相关。* P < 0.05; ** P < 0.01; *** P < 0.001。RMSEA (近似误差均方根)用于评价模型拟合度。

Fig. 5 Structural equation models illustrating the factors influencing the slope (a), intercept (b), and minimum area for species coexistence (Amin) (c) of the species-area relationship. SWC, Soil water content. Black arrows represent positive effects, while red arrows represent negative effects. * P < 0.05; ** P < 0.01; *** P < 0.001. RMSEA (root mean square error of approximation) is used to evaluate the model fit.

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氮和牛粪输入对典型草原植物种-面积关系的影响

Effects of nitrogen and cow dung input on plant species-area relationship in a typical grassland

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