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

doi:10.17520/biods.2025251

Abstract

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

Weiyu Lu, Xu Chen, Riqian Zhang, Yunhai Zhang, Zhao Li. Effects of nitrogen and cow dung input on plant species-area relationship in a typical grassland[J]. Biodiv Sci, 2025, 33(12): 25251.

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URL: https://www.biodiversity-science.net/EN/10.17520/biods.2025251

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

Figures/Tables 6

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).

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.

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

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.

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.

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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