Compensatory dynamics among dominant species stabilize plant communities in Tibetan alpine steppes
communications earth & environment
Article
A Nature Portfolio journal
https://doi.org/10.1038/s43247-026-03596-8
Compensatory dynamics among dominant
species stabilize plant communities in
Tibetan alpine steppes
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Junfu Dong
1,2
, Lei Zhao
3
, Kai Xue
4,5
4,6
, Zhe Pang , Xiaoyong Cui
7
& Yanfen Wang
4,5
Plant community stability is often hindered by nitrogen and phosphorus deficiencies in high-altitude
alpine steppes. While nutrient addition can mitigate these constraints, the individual and interactive
effects of these elements to community stability remain unclear. Here we elucidated the underlying
mechanisms by decomposing asynchrony into compensatory and statistical-averaging effects across
undegraded and degraded grasslands on the Tibetan Plateau, under conditions with and without
nitrogen and phosphorus additions. In undegraded alpine steppes, phosphorus limits plant coverage,
while nitrogen joins as a limiting factor under degradation. Phosphorus addition threatens stability in
pristine areas, whereas nitrogen enhances degraded steppes’ biodiversity and statistical-averaging
effects. Despite varying nutrient scenarios, the compensatory effects among dominant species persist,
primarily regulating community stability, rather than population stability or statistical-averaging effects.
These results underscore the pivotal role of dominant species in community stability and their diverse
compensatory dynamics in undegraded and degraded steppes under nutrient addition conditions.
Grassland ecosystems, covering about 40% of the Earth’s land surface, serve
as an important reservoir of biodiversity and primary productivity to human
societies1. In these ecosystems, nitrogen (N) and phosphorus (P) are mostly
key nutrients that restrict plant growth and influence community stability2,3,
particularly in the most barren regions, such as the Tibetan alpine steppes4.
Compared to other ecosystems, this area is one of the most vulnerable and
elevated ecosystems in the world, responding to external disturbances
quickly5,6. Several lines of evidence indicate that Tibetan alpine steppes are
naturally limited in N and P6–8. With the increase of N and P additions from
atmospheric deposition and farming practices, the local ecosystems are
undergoing remarkable nutrient inputs in this region9–11. The nutrient
limitation can then be altered or partially mitigated, but it is imperative to
investigate how the plant community stability will change in comparison to
its current state.
The way that grassland plant communities respond to the addition
of N and P varies depending on the species-specific reactions. According
to a global study conducted over 45 grasslands, N addition reduced
legume cover in nitrogen-deficient soils while boosting non-nitrogenfixing plants. Conversely, P and other nutrients enhanced legume
abundance but failed to mitigate the adverse effects of N12. These varied
responses to nutrient additions are attributed to plant traits, such as
nutrient uptake strategies and competitive abilities, which ultimately
impact the plant community stability13,14. Depending on species composition, nutrient additions can either benefit certain plants or lead to
their competitive exclusion, thereby modulating its effect on the temporal stability of plant communities15. Such modifications result in
changes in community-level attributes, including species richness and
evenness, which are pivotal to the stability and function of ecosystems16.
Studies showing that N and P additions reduce the stability of grasses-a
dominating functional group in many grasslands-underscore the role of
plant functional types in mediating these responses13. In particular, N
addition increases the inter-annual variation of primary productivity
based on 1070 multi-year paired observations, indicating a shift towards
species with higher inter-annual variability in abundance. The concomitant reduction in species richness further intensifies this negative
impact of nutrient addition on community stability13. Despite these
insights, the relationship between species diversity and community
stability remains contentious across ecosystems.
1
School of Life Sciences, Shandong University, Qingdao, China. 2Qinghai Provincial Key Laboratory of Cold Regions Restoration Ecology, Northwest Institute of
Plateau Biology, Chinese Academy of Sciences, Xining, China. 3Beijing Key Laboratory of Biodiversity and Organic Farming, College of Resources and Environmental Sciences, China Agricultural University, Beijing, China. 4College of Resources and Environment, University of Chinese Academy of Sciences,
Beijing, China. 5Beijing Yanshan Earth Critical Zone National Research Station, University of Chinese Academy of Sciences, Beijing, China. 6Key Laboratory of
Adaptation and Evolution of Plateau Biota, Northwest Institute of Plateau Biology & Institute of Sanjiangyuan National Park, Chinese Academy of Sciences,
e-mail:
Xining, China. 7College of Life Sciences, University of Chinese Academy of Sciences, Beijing, China.
Communications Earth & Environment | (2026)7:433
1
Article
https://doi.org/10.1038/s43247-026-03596-8
Nevertheless, how plant species mediate community stability under
nutrient enrichment remains poorly understood. Hautier, Zhang15 bridge
this gap, showing that fertilization consistently weakens the stabilizing effect
of plant diversity across spatial scales. This occurs because fertilization
dampens asynchronous dynamics among species, a key mechanism for
stability in diverse plant communities15. Such responses to nutrient additions can lead to shifts in plant community composition, thereby affecting
grassland stability. Seabloom, Batzer17 further found that nutrient addition,
especially N, reduces local diversity and increases the minimal area of
coexistence, indicating changes in the scales at which species interact. Longterm N addition can also alter the β-diversity of plants in a semiarid
grassland3, and the above-ground net primary production (ANPP) can be
boosted by N and P additions in grasslands but decreased for community
stability14. Some studies suggest that this increase in productivity due to
nutrient additions can enhance community stability through the portfolio
effect18–20. Additionally, long-term warming and N addition can have
additive negative effects on community stability21. This suggests that the
resistance of grassland communities to N and P additions may be reduced
under certain conditions21, underscoring the potential impacts of N and P
additions on grassland ecosystems and the complexity of their effects on
plant community stability across different environmental contexts.
Dominant species also play an important stabilizing role in ecosystems
worldwide. Consistent with the mass ratio hypothesis, dominant species
exert a larger impact on ecosystem functions and community stability than
the non-dominant species22. N and P addit (...truncated)