Figure 6 Piecewise SEM describing how biotic and abiotic variables affect ecosystem multifunctionality. Numbers on the arrows represent all the path coefficients and squared multiple correlations (R 2) in the model. Solid lines indicate significant paths and dashed lines represent paths that were not significant. Positive path coefficients are pink and negative path coefficients are blue. A single asterisk (*) and double asterisk (***) indicate a significant difference between the variables at P < 0.05 and P < 0.001, respectively.
4 DISCUSSION
4.1 The diversity and composition of different trophic levels link to ecosystem functions
Exploring the spatial distribution patterns and mechanisms of biological species diversity on Earth has always been an important part of ecological and biodiversity research. Our study found a correlation between species diversity indices of different trophic levels, especially in terms of species richness indices (Figure S4), which is consistent with previous findings (Wang and Xing, 2021) indicating that species diversity can be an important indicator reflecting community dynamics. Meanwhile, plant communities undergo changes due to natural disturbances in the form of succession, resulting in changes to the community environment and effects on herbivores, pollinators and microbes (Beck et al., 2015; Tao et al., 2023). The relationship between plants and microbes can be understood from the mechanism of plant–soil feedback, where different plant communities produce different types of litter and root exudates, affecting soil organic carbon content and thereby changing the composition and structure of soil microbial communities (Smith et al., 2015). Researchers found that the influence of plant diversity on multifunctionality was indirectly influenced by shifts in soil biodiversity and plant cover (Delgado-Baquerizo et al., 2020). This underscores the importance of incorporating soil biodiversity into policy and management initiatives aimed at safeguarding the functionality of terrestrial ecosystems on a global scale. The activities of different types and quantities of rodents, through digging and seed dispersal behaviors, have been shown to influence the nutrient characteristics of the soil and plant species (Pech et al., 2007; Wesche et al., 2007), leading to changes in the composition of the surrounding plant community (Figure S6).
Ecosystem functions are closely related to the composition of keystone species in biological communities. The realization of ecosystem functions often depends on certain keystone species in the biological community, such as important predators, pollinators, and major tree species in forests (Raffard et al., 2021; Joshi et al., 2022). Our study found that keystone species in plant, animal, and microbial communities were interconnected with indicators of ecosystem functions (Figure S9), suggesting that biodiversity influences ecosystem functions and stability through interactions of species traits at multiple trophic levels (Turnbull et al., 2016). At the same time, our results provided evidence for the theory that ecological niche differentiation is necessary to maintain species coexistence (MacArthur, 1970). For example, the similarity of soil functional requirements of some plants (e.g., Thalictrum aquilegiifolium var. sibiricum ,Sibbaldianthe bifurca , Saussurea pulchra , andEuphorbia altotibetica , Figure S9a) and their different ecological niche widths in the community (Figure 1b) suggested that there was obvious competition for resources in the plant community, which affected the differences in the distribution of individual plants in the community. It has been shown that species ecological niche differentiation occurs during the construction of functional groups in grasslands due to environmental factor constraints, resulting in the coexistence of some species and the loss of others (Dray and Legendre, 2008), where environmental factors determine which traits are included in functional groups that could be clustered in localized communities (Ter Braak et al., 2012). Species ecological niche differentiation and interspecific interactions based on functional traits thus play a decisive role in plant community construction and diversity maintenance (Kraft et al., 2008). In different vegetation areas, microbial key species were also subject to limitations on ecosystem functioning (Figures S5, S9b and S9c). The planctomycete_A-2 belongs to Planctomycetes, which have the ability to utilize diverse carbon compounds and can survive solely on high-molecular-weight dextran as their carbon source, thereby participating in the carbon cycle (Boedeker et al., 2017). Consequently, it was unsurprising to find that planctomycete_A-2 was susceptible to nitrogen and carbon limitation in our study. Meanwhile, Pseudomonas_frederiksbergensis has shown potential to solubilize insoluble mineral phosphate (Zeng et al., 2016), which promotes the release of effective phosphorus, and plant uptake of phosphorus plays a significant role in promoting plant growth. We also found that Pseudomonas_frederiksbergensis and the total phosphorus content had a highly significant positive correlation.Xenodidymella_camporesii and Verrucaria_viridigrana belong to the phylum Ascomycota, which has been reported to tolerate stressful conditions such as low nutrient availability, leading to more efficient use of resources in challenging environments (Chen et al., 2017). This could explain the lack of significant correlation with most soil functions.
Rodents are the primary prey of many carnivores, while also consuming plant seeds and fruits, thereby converting plant energy into animal energy and dispersing plant seeds to different locations, promoting biodiversity maintenance (Yu et al., 2023). Our research found that the abundance of root voles was positively correlated with soil total phosphorus content, nitrate nitrogen content, and aboveground biomass (Figure S9d). A previous study suggested that the metabolic activities of root voles produce urea and provide abundant nitrogen resources for the areas surrounding their burrows, thus making the environment suitable for plant survival and growth (Li et al., 2021). Plateau pikas, as rodents mainly active underground, contribute to soil ventilation and water infiltration through gnawing, digging, and excretion behaviors, bringing organic matter and nutrients to the soil surface, enhancing ecosystem functions, and providing favorable conditions for plant growth (Zhang et al., 2003).
4.2 The pathways of multitrophic diversity drive ecosystem multifunctionality
Most studies have begun to focus on changes in ecosystem multifunctionality at different trophic levels. Our study showed that multitrophic-level diversity had a greater impact on ecosystem multifunctionality than the diversity of a single biome, suggesting that maintaining ecosystem multifunctionality requires the joint participation of biomes (Figures 4, 5 and 6), which is consistent with the results of others (Schuldt et al., 2018; Luo et al., 2022; Mori et al., 2023). Therefore, the relationship between biodiversity and ecosystem multifunctionality is more dependent on diversity at the level of multiple communities, and researchers should carry out their analyses at large spatial scales rather than in single communities when considering species diversity.
In addition to this, we found that ecosystem functions such as soil and water conservation, soil fertility, nutrient cycling and transformation, and community productivity were mediated by different single communities in addition to multitrophic level diversity (Figure 4). Due to the similarity in functional traits among different trophic groups and the low redundancy between trophic levels, different trophic groups support different functions (Soliveres et al., 2016; Luo et al., 2022). Plant diversity had a positive effect on water conservation. The increase in water conservation may result from increasing vegetation coverage, which could lead to a reduction in soil evaporation and enhancement of rainfall interception, as well as an increase in soil carbon and nitrogen levels (Figure S10). This is because the presence of soil organic matter can improve soil porosity, thereby boosting both soil water infiltration and retention (Zhu et al., 2019; Zhao et al., 2023). Plant community diversity, on the other hand, had a negative effect on community productivity, which may have been due to the fact that increased plant species richness increased competition among plants, and plant competition has been shown to stimulate an increase in plant litter and root secretions (Laganière et al., 2015). Our study in turn confirmed that there were differences in resource utilization strategies among different keystone species in the plant community, and therefore community productivity can be highly unstable in this situation, producing a downward trend. Soil microbes play an important role in driving EMF (Jing et al., 2015; Delgado-Baquerizo et al., 2020). The higher the soil bacterial diversity, the more important a role it plays in nutrient cycling and transformation functions, and different microbial species are strongly associated with different ecosystem functions. Maintaining a higher richness of microbial taxa is crucial for supporting increased functional redundancy and diversity, which helps explain why greater biodiversity is necessary to sustain more ecosystem functions (Wagg et al., 2019).
Here, we discovered that considering the characteristics of bacterial communities often proved to be a more effective predictor of ecosystem multifunctionality compared to considering fungal communities (Figure 4). This was due to the distribution of metabolic tasks among microorganisms, creating synergies between microorganisms with distinct physiological characteristics, such as those between fungi and bacteria. This highlights the significance of interconnections between microbial communities in influencing ecosystem functions, suggesting that these hidden synergies may have a broader and more significant ecological impact on soil microbial functions than previously recognized (Kohlmeier et al., 2005; Deveau et al., 2018). Our study also found that soil fertility, grassland productivity, and ecosystem multifunctionality increased significantly with rodent community diversity, and that, in general, mammals at low trophic levels, such as hares and rats, are dependent on food resources and shelter provided by good ecological niches. However, we also suspect that anthropogenic activities may be replacing top-down ecological effects that are partially independent of top predators. These ecological effects are not mediated by top predators but directly affect small mammals (Smiley et al., 2020; She et al., 2023).
Changes in the diversity of multiple trophic levels can be explained by variations in the aboveground plant niche breadths, and as the niche expands, the diversity of multiple trophic levels and ecosystem multifunctionality also increase (Figure 6). The response of plant species to interspecific competition involves altering their niches to reduce overlap with other species, making them more complementary in resource utilization (Eisenhauer et al., 2019). Plant communities with high diversity select for enhanced niche differentiation among species, reducing interspecific competition without increasing intraspecific competition. This mechanism may strengthen the relationship between biodiversity and ecosystem functioning (Amyntas et al., 2023). The relationship between plant diversity and ecosystem functioning depends not only on interactions between plants but also on interactions within and between different trophic levels of the food chain (Barnes et al., 2020; Albert et al., 2022). This emphasizes the importance of plants as primary producers in connecting aboveground consumers and belowground decomposers, thus influencing changes in multi-trophic level biological communities.
4.3 Altitude and pH influence ecosystem multifunctionality
Biodiversity is not the sole or primary driving factor of ecosystem multifunctionality (Giling et al., 2019); climate and abiotic factors also drive ecosystem functions (Grytnes and McCain, 2013). We found that biotic factor of altitude and pH directly explained changes in ecosystem multifunctionality (Figures 5 and 6), which aligned with previous evidence (Hu et al., 2020; Luo et al., 2022). Altitude serves as a predictive factor for patterns of species diversity and community composition under environmental changes. With increasing altitude, temperature decreases, solar radiation increases, and wind strength intensifies, putting significant stress on the physiological and survival strategies of species (Galván-Cisneros et al., 2023). Studies had shown that ecosystem multifunctionality significantly decreases with increasing altitude (Chen et al., 2022),which is contrary to our results. The reason for this discrepancy may be that the relationship between altitude and ecosystem multifunctionality is not simply linear but follows a single-peak curve. Research on the dependence of ecosystem multifunctionality on altitude in the Qinghai-Tibet Plateau indicated that there was a critical point around 3900 m where the relationship between ecosystem multifunctionality and altitude changes (Wang et al., 2023b). Our study sites were located near this critical point, which might explain why there was a positive correlation between altitude and ecosystem multifunctionality in our findings. Soil pH directly affects processes such as mineral weathering, organic matter mineralization, and humification in soils, which have a significant impact on the status of nutrient ions in soils. Most studies have confirmed that soil acidification reduces ecosystem functionality (Delgado-Baquerizo et al., 2016; Wei et al., 2022),which is inconsistent with our research findings. Through our study of sample points, we found that high-altitude grasslands with lower pH and higher ecosystem multifunctionality show obvious shrub encroachment. The pH content of shrub-encroached high-altitude grasslands tends to be acidic (Ma et al., 2022),and shrub encroachment can enhance ecosystem functionality, specifically in terms of carbon sequestration, soil fertility, and hydrological functions (Ding and Eldridge, 2023). These results support our conclusion.
5 CONCLUSIONS
In summary, our results indicate that key species at different trophic levels are interconnected with ecosystem functions, leading to species niche differentiation within communities and influencing the formation of biodiversity. The interconnections of diversity at a single trophic level, especially richness indices, highlight the importance of biodiversity in maintaining food web structures and functions within biological communities. The greater the number of species in an ecosystem, the more diverse the food base supply, ensuring ecosystem stability. Furthermore, we confirmed the role of multi-trophic diversity, namely biodiversity (i.e., plant diversity, bacterial diversity, and rodent diversity), in maintaining ecosystem functions, further emphasizing the critical importance of conserving biodiversity in sustaining ecosystem functions. Lastly, abiotic factors such as altitude alter ecosystem multifunctionality by affecting soil environments, and multitrophic diversity directly impacts ecosystem multifunctionality. Therefore, in the face of increasing human activities and climate change, the maintenance of multifunctionality through multitrophic diversity is crucial and requires our utmost attention.
AUTHOR CONTRIBUTIONS
All authors contributed to the article. H.S., H.Z., Z.W., and H.L., made efforts to define the topic of the study and guided the writing of the article. The help of R.Q. and T.C. solved the problem of rodent capture. L.M., Z.Z., X.H., and F.Y. solved the problems encountered during data processing and graphing. J.Y., X.L., S.L., J.W., H.A. and Z.S. participated in the field survey and sampling work.
DATA AVAILABILITY
Data will be made available on request.
ACKNOWLEDGMENTS
This study was funded by the National Natural Science Foundation of China (32371684), the National Natural Science Foundation of China Joint Fund Project (U21A20186, U20A2006), the Second Tibetan Plateau Scientific Expedition and Research (STEP) Program (2019QZKK0302–02), and the International Cooperation Project of Key Research and Development and Transformation in Qinghai Province (2024–HZ–810).
CONFLICTS OF INTEREST
The authors declare no competing interests.
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