Abstract
Purpose
Nitrogen (N) deposition is well known to affect plant community assembly, but the underlying mechanisms are still poorly understood. How different processes, including environmental and biotic filtering, drive plant community assembly can be quantified by trait range restrictions and patterns of trait convergence and divergence. However, few studies have combined species abundances data to simultaneously explore how these two processes function and further affect community assembly along N addition gradients by progressively examining multiple trait range restrictions and trait patterns.
Methods
We conducted a four-year N addition experiment with eight N addition levels in a temperate desert steppe, Inner Mongolia. We investigated species composition and then collected aboveground biomass (AGB). In addition, six soil properties and functional traits were measured. Community-weighted trait means (CWM) and community-weighted trait variances (CWV) at eight N addition levels were calculated based on species abundances and their corresponding traits. We quantified environmental filtering as a restriction of trait range, and trait patterns depended on CWV. We calculated the effect size (ES) values of six trait ranges and CWVs by comparing the observed community with two simulated null communities generated by a two-step trait-based approach. Using the one-tailed Wilcoxon signed-ranks tests (W) and the Spearman’s rank correlations (S) methods, we further explored changes in six trait ranges and trait patterns along experimental N addition gradients by the ES values.
Results
We found that high N addition altered species composition, and the dominant S-strategy species (conservative species) in high-barren and low-disturbance environments were gradually replaced by R-strategy species (acquisitive species) in low-barren and high-disturbance environments. The polynomial regression showed that AGB was highest at 27 g.m−2 N addition level. The responses of six CWM values under N24 and N48 were significantly different from those under other N addition treatments, and they showed two opposite trends along N addition gradients. The CWM values of three traits (i.e., plant height (H), leaf dry matter content (LDMC) and leaf carbon content (LCC)) significantly decreased, while the CWM values of other three traits (i.e., leaf thickness (LT), specific leaf area (SLA) and leaf nitrogen content (LNC)) significantly increased. The trait range of H, LDMC and LCC was wider than expected in N-poor communities, while narrower than expected in N-rich communities, and the opposite was true for LT and LNC. The trait range of SLA was overall narrower than expected at eight N addition levels. These results indicate that environmental filtering restricts the trait range of H, LDMC and LCC in N-rich communities, while restricts the trait range of LT and LNC in N-poor communities, and restricts the trait range of SLA under almost all N addition treatments. All six traits converged first, and then diverged along N addition gradients, indicating that high N addition induces functional trait differentiation in a temperate desert steppe.
Conclusion
Our study clearly demonstrates that environmental and biotic filtering play important roles in assembling communities along N addition gradients in desert steppe. Changes in trait ranges induced by environmental filtering strongly depend on the types of functional traits along environmental gradients. In the processes of biotic filtering, trait hierarchy drives the outcomes of competitive interactions among coexisting species in N-poor communities and further determines community assembly, as evidenced by trait convergence and large differences in competitive ability among species. In contrast, trait divergence and small differences in competitive ability among coexisting species in N-rich communities reveal that trait dissimilarity drives community assembly, corresponding to large functional trait differentiation. In summary, shifts in patterns of trait convergence and divergence along N addition gradients largely support the stress-dominance hypothesis, but our results also reveal that community assembly mechanisms are more complex. When attempting to explore assembly mechanisms along environmental gradients, not only species multiple traits and occurrences but also their abundances and specific environments should be considered.
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Data Availability
The datasets generated during and/or analyzed during the current study are available from the corresponding author on reasonable request.
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Acknowledgements
This study was supported by the National Natural Science Foundation of China (41622103 and 41571106), Youth Innovation Promotion Association CAS (E229880105) and the Second Tibetan Plateau Scientific Expedition and Research program (2019QZKK0305).
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Jingjuan Qiao collected and analyzed the data and wrote the paper.
Xiaoan Zuo designed the research and contributed to writing.
Ping Yue and Shaokun Wang assisted with paper writing.
Ya Hu, Xinxin Guo, Xiangyun Li, Peng Lv, Aixia Guo and Shanshan Sun participated in the experiment together.
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Qiao, J., Zuo, X., Yue, P. et al. High nitrogen addition induces functional trait divergence of plant community in a temperate desert steppe. Plant Soil 487, 133–156 (2023). https://doi.org/10.1007/s11104-023-05910-1
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DOI: https://doi.org/10.1007/s11104-023-05910-1