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Article

Warming Increases Ecological Niche of Leymus chinensis but Is Detrimental to Species Diversity in Inner Mongolia Temperate Grasslands

by
Xingbo Zhang
1,
Zhiqiang Wan
1,2,*,
Rui Gu
3,
Lingman Dong
1,
Xuemeng Chen
1,
Xi Chun
1,2,
Haijun Zhou
1,2 and
Weiqing Zhang
1
1
College of Geographical Science, Inner Mongolia Normal University, Hohhot 010020, China
2
Key Laboratory of Mongolian Plateau’s Climate System, Universities of Inner Mongolia Autonomous Region, Hohhot 010020, China
3
College of Grassland, Resources and Environment, Inner Mongolia Agricultural University, Hohhot 010020, China
*
Author to whom correspondence should be addressed.
Agronomy 2024, 14(10), 2425; https://doi.org/10.3390/agronomy14102425
Submission received: 12 September 2024 / Revised: 15 October 2024 / Accepted: 17 October 2024 / Published: 19 October 2024
(This article belongs to the Section Grassland and Pasture Science)
Browse Figures
Versions Notes

Abstract

:
Dominant species are crucial in regulating the structure and productivity of plant communities. Adaptation strategies to climate change vary among the dominant species of different life types. However, the responses of the ecological niches of dominant species to warming and precipitation in semi-arid grasslands and their impacts on community structure and function are unknown. This study involved conducting a long-term experimental simulation of warming and increased precipitation on grasslands in Inner Mongolia and studying population dynamics, ecological niches, and their responses to the structure and function of the community species of two dominant plants, L. chinensis (perennial rhizome grass) and S. krylovii (perennial clumped grass). The results show that the niche width of L. chinensis increased and S. krylovii decreased under warming and increased precipitation conditions. The overlap of L. chinensis and S. krylovii decreased under the same conditions. The niche widths of L. chinensis and S. krylovii were 1.22 for the control (C), 1.19 and 1.04 under warming (W) conditions, 1.27 and 0.97 under warming plus precipitation (WP) conditions, and 1.27 and 1.24 under the conditions of precipitation addition (P). The niche overlap of L. chinensis and S. krylovii were 0.72 in C, 0.69 in W, 0.68 in WP, and 0.82 in P. The biomass share and importance value of L. chinensis increased, and those of S. krylovii decreased in response to warming and precipitation. The effects of warming on species diversity and community stability are primarily influenced by the effects on the niche breadth of S. krylovii. Combined with our previous study, L. chinensis will offer more resources in communities in warmer and wetter steppe climates in the future. However, this is not conducive to community diversity.

1. Introduction

Global warming, characterized by increasing temperatures, has become more pronounced over the past century. The global surface average temperature in 2011–2020 was 1.1 °C higher than that in 1850–1900, and the rapid rate of increase in the global surface temperature since 1970 has been excessive in the last 2000 years [1], while warmer temperatures will indirectly change the surface atmospheric circulation patterns, causing changes in precipitation patterns regionally and globally and an increase in extreme precipitation events [2,3].
Grassland biomes cover approximately one quarter of the earth’s land area [4]. The impacts of global climate change, characterized by increases in temperature and precipitation [5], are particularly essential for grassland ecosystems [6,7,8]. Climate warming, accompanied by changes in precipitation patterns, impacts the morphological structure and physiological responses of plants in ecosystems, causing changes in the structure and function of plant communities [9,10,11]. The competition between species decreases with increased precipitation, increasing plant diversity [12]. Dominant species are crucial in regulating plant species and community stability during environmental changes [13,14]. Warming causes droughts and decreases species richness and community stability in temperate grasslands [15]. The magnitude of the warming effect on community variability depends on the level of drought at a site environment [16]. In many restored grasslands, there is an inverse relationship between the dominant grass species and diversity [17]. Several studies have shown that productivity stability in temperate grasslands in Inner Mongolia depends on dominant species with few but high diversity [18] and that increasing the dominance of particularly stable grasses can also increase the temporal stability of the community [19].
As crucial biological factors in this system, the responses of different dominant plant species to global changes could cause large differences in their spatial patterns and interspecific relationships [20,21]. Grasslands in Inner Mongolia are dominated by perennial rhizomatous and tufted grasses, Leymus chinensis (Trin.) Tzvelev (1968) (cited as L. chinensis) and Stipa krylovii Roshev (cited as S. krylovii) (1929), respectively, which differ significantly in their morphology, photosynthetic physiology [22,23], adaptability, and adaptation strategies to the environment [24,25,26]. Insufficient moisture can cause drought-tolerant tufted grasses to become dominant in grasslands [27], whereas warming and precipitation can cause the gradual dominance of rhizomes [28,29]. However, warming has also been shown to reduce the number of dominant species, including L. chinensis and Stipa capillata, in temperate grasslands [14,15]. The differences in the responses of these two dominant plant species to climate change may owe to differences in the uptake of the root water and nutrients in the soil layer [30,31]. Differences in plant adaptability affect interspecific relationships, distribution patterns, and community structures [32,33].
The status and significance of species in a community can be expressed using the ecological niche. The effects of ecological niches on habitats and their interactions can be used to quantify competitive relationships between species and their ability to utilize resources in the environment [34,35]. Niche breadths and overlaps have been used in various studies as indicators of plant characteristics [36,37]. Studies have shown that the niche breadth of a species is positively correlated with its adaptation to the environment and decreases with excessive nutrient limitations [38,39]. The niche breadth of a single species and the degree of niche overlaps between species are negatively correlated with species richness [40]. As the intensity of species competition increases, the niche width becomes narrower, and the niche overlap becomes smaller [41]. The overlap of climatic ecological niches among alpine plant species has increased over time, and the effects of precipitation on the climatic ecological niches of alpine plants will become more prominent in future warming [42]. Population spatial patterns and ecological niche changes in dominant species are correlated with the composition and community stability of species [43].
Differences in adaptation strategies to climate change among different plant life types indicate that interspecific relationships of dominant plants in the same community are affected by climate change, which regulates community structure and function. In this study, based on climate change trends in temperate grasslands, we conducted a temperature and rain-enhancement experiment and continuously observed experimental sample plots of grassland plant communities in the field for 11 years in Inner Mongolia, China, starting in 2013. We hypothesized that (1) an increase in warming and water levels would increase the niche breadth of L. chinensis and the niche overlap value between L. chinensis and S. krylovii. (2) An increase in the niche breadth of dominant species would decrease the species diversity of the community. In this study, we aimed to address (1) how the ecological niches among dominant species in grasslands respond to changes in temperature and precipitation and (2) how the ecological niches of dominant species regulate the structure and function of grassland communities in response to environmental changes.

2. Materials and Methods

2.1. Experimental Site

The study area was in the Mauden pasture (44°09′49.0″ N, 116°29′02.3″ E; elevation of 1102 m), 40 km east of Xilinhot City, Xilingol League (Figure 1). This area is in a semi-arid subregion of a mesothermal zone with a temperate continental climate and large annual temperature differences. The annual mean temperature of the study site during the experimental period was 2.6 °C, and the average precipitation was approximately 271 mm, which was primarily concentrated in May–September, accounting for 87.3% of the total annual precipitation. Data were obtained from the China Meteorological Data Sharing Service System of the China Meteorological Administration. We determined the dominant species based on their relative abundance. A dominant species was defined as that with an average relative aboveground biomass at the control sample site, accounting for over 5% of the total aboveground community biomass [44]. Two dominant species were identified, namely, perennial rhizomatous grass (31.71%) and perennial tufted grass (56.53%).

2.2. Experimental Design

Increases in warming and precipitation levels indirectly regulate the growth of plant communities by affecting soil moisture, which has significant lag and cumulative effects. Therefore, determining a longer time scale as a parameter is essential to accurately predict the changes in a plant community under climate change. An experiment for simulating temperature increases and precipitation changes began in April 2013, and a randomized block design was adopted. Four treatments were established, namely, control (C), warming (W), precipitation addition (P), and warming plus precipitation addition (WP). Each treatment plot was 3 × 3 m with four replicates, and 16 experimental areas were established. The warming treatments were performed using open-top growth chambers. The warming was set at 2 °C, according to the IPCC Fourth Assessment Report. The addition of precipitation was set at 20% of the average precipitation in each month between 1961 and 2010, and this precipitation supplementation was conducted twice a month at the beginning and the end of each month during the growing period (between May and September). We conducted the precipitation addition after 18:00 to limit evaporation. We increased the monthly precipitation of the OTC by 0.75 times to eliminate the shading effect caused by the difference between the top area and the bottom area of the OTC (0.75 m2). Each sample plot was replenished with 5, 9, 16, 13, and 5 mm of water between May and September for 48 mm of added precipitation. In addition, the increase in precipitation at OTCs partially offset the decrease in soil moisture caused by warming.

2.3. Measurement Methods

Each plot was surveyed once a year in August during the growing season to determine the composition of the community species, population density, and plant height. Density, a quantitative measurement of the spatial distribution of a population or community under certain environmental conditions, was determined using the number of individuals of each plant species in a sample plot. The identification of single plants varied with differences in their morphology and reproduction, with the individual plants of S. krylovii being easily distinguished and counted, while for L. chinensis, aboveground stems were used as an indicator of plant number [45]. We used a straightedge to obtain plant height measurements from the emergent portion of the plant and selected five individuals of each species to calculate their average height. To represent the degree of species dominance for each species, an importance value (IV) was calculated using the ratio of the height (or density) of the species to the sum of the heights (or densities) of all species (i.e., relative height or relative density). A linear regression model between height density and biomass was established and used to calculate the biomass in the sample (Table S1).

2.4. Data Computation and Analysis

The Shannon–Wiener index was used to determine the ecological niche width of dominant species [46], and the Schoener index was used to calculate the ecological niche overlap between dominant species [47].
B i = − ∑ j = 1 r ( P i j ln ⁡ P i j )
O i k = 1 − 1 2 ∑ j = 1 r P i j − P k j
where Pij = nij/Nij; Pij is the ratio of the number of individuals of species i to the number of individuals of the species in the jth resource state; nij is the number of individuals of species i utilizing the jth resource state, which is expressed by the importance value of species i in the jth sample in the present study; Nij is the total number of species i; and r is the sample number. Oik is the ecological niche overlap coefficient of species i and k, and Pij and Pkj are the importance values of species i and k in the jth sample, respectively.
The process in which the species composition of a community reaches a state of equilibrium and stabilizes again after a disturbance is called community diversity. Plant community stability is evaluated by calculating the inverse coefficient of variation (ICV) [48]. Species diversity is an essential characteristic of communities and a function of species richness and evenness. The Margalef Index (SD) considers only the number of species and the total number of individuals in the community and defines the number of species in a given sample size as a diversity index [49]. The formula is as follows:
I C V = μ σ
S D = S − 1 ln ⁡ N
where μ denotes the mean of the relative importance values of all plant species in a given sample, σ denotes the standard deviation of the mean of the relative importance values of all plant species in a given sample, N is the total number of species in a sample or community, and S is the total number of species in a sample or community.
Segmented structural equation modeling (SEM) was used to assess the effects of temperature on the ecological niches of dominant species and precipitation changes on the structure and function of grassland ecosystem communities. Before building the SEM, we developed a conceptual model with variables that could affect ecosystem diversity and stability for the ecological niches of both dominant species, after which we used Pearson’s correlation analyses to measure the correlation between plant variables and ecosystem diversity and stability. Variables that did not significantly correlate with ecosystem diversity or stability were excluded from the model. The path coefficients were obtained using maximum likelihood estimation techniques. The best model with the lowest Akaike information criterion value (p > 0.05) and RMSEA (< 0.05) was selected.
Microsoft Excel 2019 was used for data organization and statistical analyses. The niche breadth and overlap values were calculated using niche Width () and Overlap () in the interspecific linkage analysis package Spatia in RStudio (Version 4.1.3 https://github.com/rstudio/rstudio, accessed on 26 January 2024). Correlation heatmaps were used to analyze the relationships between the structural indicators of the dominant species, ecological niche data, and ecosystem stability. Structural model equations were calculated and plotted using the AMOS 22.0 software (AMOS Development Co., Armonk, NY, USA) [50].

3. Results

3.1. Effects of Increased Temperature and Rainfall on the Ecological Niches of Dominant Species

Warming plus precipitation addition increased the niche breadth of L. chinensis, decreased that of S. krylovii, and decreased the dominant species niche overlap. The niche breadth and overlap of the dominant species were reduced under warming alone and increased under precipitation addition alone (Figure 2). The Multi-year mean niche breadth values of L. chinensis and S. krylovii were 1.22 under the C condition, 1.19 and 1.04 under W conditions, 1.27 and 0.97 under WP conditions, 1.27 and 1.24 under P conditions, respectively (Figure 2a,b). The Multi-year mean niche overlap values between L. chinensis and S. krylovii were 0.72 in C, 0.69 in W, 0.68 in WP, and 0.82 in P (Figure 2c). The Multi-year mean niche breadth ratios of L. chinensis and S. krylovii were 1.007 and 1.003 in the C and P treatments, respectively. The W and WP treatments increased the niche breadth ratio of both species, and their Multi-year mean values were 1.225 and 1.205, respectively (Figure 2d). Since 2021, the niche breadth ratio of L. chinensis and S. krylovii has increased rapidly, reaching more than 2.

3.2. Effects of Increased Temperature and Rainfall on the Importance of Dominant Species in Community Structure and Functioning

Warming and precipitation addition increased the biomass percentage of L. chinensis and decreased that of S. krylovii. The relative biomasses of L. chinensis and S. krylovii were 25% and 64.6%, respectively, under C conditions. The relative biomasses of L. chinensis and S. krylovii were 48.9% and 42.4% in W and 48.9% and 37.4% in WP, respectively. The relative biomass of L. chinensis increased significantly in the experiment involving warming conditions, while that of S. krylovii decreased significantly under the same conditions (p < 0.01 for L. chinensis and S. krylovii). The relative biomass levels were similar and did not increase or decrease in the P treatments (43.1% and 45.6% for L. chinensis and S. krylovii, respectively (Figure 3a)).
Warming and precipitation addition increased the importance of L. chinensis and decreased that of S. krylovii. Compared with the C group, the W, WP, and P treatments increased the IV of L. chinensis by 54.9%, 53.6%, and 39%, respectively, and decreased that of S. krylovii by 28.6%, 39%, and 22.6%, respectively. Warming had a greater effect on the magnitude of IV increase for L. chinensis; however, for S. krylovii, warming plus precipitation addition caused the greatest decrease. Warming and precipitation addition reduced species diversity; however, warming alone reduced diversity more significantly. Species diversity was reduced by 24.6%, 13.7%, and 9.8% in the W, WP, and P treatments, respectively (Figure 3b).

3.3. Relationships Between Ecological Niches and Ecosystems Under Warming and Water-Augmentation Conditions

Warming and precipitation addition exacerbated the mutual inhibition between the IV of the two dominant species (p ≤ 0.001). There was a positive correlation between the biomass and IV of L. chinensis under the different treatment conditions (p ≤ 0.01). The negative correlation between the L. chinensis biomass and the importance of S. krylovii was exacerbated by the effect of warming alone (p ≤ 0.001) and decreased (p ≤ 0.05) with the effect of precipitation addition alone (Figure 4). The T (p ≤ 0.01) and TP (p ≤ 0.05) treatments caused a significant reciprocal inhibition of the biomass of the two dominant species; however, they were positively correlated (p ≤ 0.01) under the P condition. The warming treatment caused a significant negative correlation between the biomass of S. krylovii and the IV of L. chinensis while having the opposite relationship with its IV (p ≤ 0.001) (Figure 4b,c). Warming and precipitation addition exacerbated the significant positive correlation between the niche breadth of L. chinensis and its IV and biomass and a negative correlation with the IV of S. krylovii. The niche breadth of L. chinensis was negatively correlated with the biomass of S. krylovii under W and WP treatments, while the interaction between them was reduced under P treatments. Precipitation addition alone exacerbated the positive correlation between the niche breadth of S. krylovii and the importance and biomass of L. chinensis; the opposite correlation between the niche breadth and importance of S. krylovii was also intensified in the same treatment. Warming and precipitation addition reduced the positive correlations between the niche breadth of S. krylovii and its biomass and the niche breadth of L. chinensis. The niche breadth of the two dominant species and their overlap had a negative effect with warming and precipitation addition.
Warming alone caused a positive correlation between species diversity and the niche breadth of L. chinensis and its biomass and a negative correlation with the niche breadth of S. krylovii and its biomass. However, the treatments involving warming and precipitation addition treatments showed opposing trends. The species diversity and niche breadth of L. chinensis were correlated across treatments (Figure 4). Species diversity was positively correlated with the niche breadth of L. chinensis in the C (p ≤ 0.01) and W treatments (p ≤ 0.05) and negatively correlated with the niche breadth of L. chinensis in the WP (p ≤ 0.05) and P (p ≤ 0.01) treatments. Species diversity was positively (p ≤ 0.05) and negatively (p ≤ 0.01) correlated with niche breadth in the C treatment and the niche overlap in the WP (p ≤ 0.05) and P (p ≤ 0.01) treatments. In the WP treatment, species diversity was positively correlated (p ≤ 0.05) with the niche overlap of dominant species, while species diversity was negatively correlated (p ≤ 0.01) with the niche breadth of S. krylovii under the P treatment. Warming had a negative effect on the niche breadth of S. krylovii (p ≤ 0.01), and the niche breadth of S. krylovii had a positive effect on species diversity (p ≤ 0.001).
Changes in precipitation had a direct effect on species diversity (p ≤ 0.05). We found that precipitation was the primary factor significantly correlated with community stability in the ecological niche of the dominant species, and the value of niche overlap between dominant species was positively correlated with community stability (p ≤ 0.05) in the P treatment (Figure 4d). Precipitation changes influenced community stability (p ≤ 0.001) by affecting species diversity (p ≤ 0.05) (Figure 5) but had a direct effect on community stability (p ≤ 0.01). Warming had a negative effect on the niche breadth of S. krylovii (p ≤ 0.01), and the niche breadth of S. krylovii also had a negative effect on community stability (p ≤ 0.01). Temperature changes affected community stability (p ≤ 0.001) through the two steps of the niche breadth of S. krylovii (p ≤ 0.01) and species diversity (p ≤ 0.001).

4. Discussion

4.1. Effects of Warming and Increased Precipitation on the Ecological Niche of Dominant Species

The ecological niches used by species provide comprehensive insights into their status in a given ecosystem community [51]. L. chinensis and S. krylovii, which accounted for approximately 90% of the biomass of the observed community, were the dominant species in the grasslands of the temperate zone. Climate change has specific effects on plant ecological niches, and different environmental conditions have varying effects on different dominant plant species [52]. Niche changes in dominant species are critical for determining the sensitivity of ecosystems to climate change [53]. Our study shows that climate warming decreases the niche breadth of dominant species, whereas year-round warming increased the niche breadth of dominant species in alpine meadows on the Tibetan Plateau [54]. This is primarily because the dominant species of temperate grasslands are not the same as those of alpine meadow grasslands, and the difference in temperature and soil moisture also affects the results of the experiments. Decreased soil moisture owing to warming is a significant mechanism driving the impact of climate change on grasses [55]. Increased rainfall is crucial in stabilizing the growth of dominant plants in northern grasslands, which is consistent with the findings of [56]. The simultaneous effects of climate warming and increased precipitation levels will cause the niche breadth of dominant species to increase or even reverse in later stages [56], suggesting that increased precipitation will offset the effects of warming on the ecological niches significantly; however, warming will have a more profound effect on the ecological niche trend of the dominant species in the long term.
Classical ecological niche theory suggests that each species can survive under limited conditions [57], and the only condition regulating species coexistence is the degree of niche overlap between them [58]. Our results show that the niche breadth and overlap of the dominant species decreased under warming alone and increased under the conditions of precipitation addition alone (Figure 2a,b). In this study, we found that there was a niche overlap of major plant species in temperate grasslands, and the dominant species under moisture-increasing conditions had a more intense competition with overlap values ≥0.92 (Figure 2c). This may be because the ecological balance of the grassland was disrupted under water-augmentation conditions, and the plant species reallocated biological resources and increased competition under the rain-augmented environment. This is consistent with the findings of Silvertown et al., who found that the expected changes in the niche overlap of meadow plants on hydrological gradients are different and that ecological niche segregation can occur at all phylogenetic levels [59].
In our study, in grassland communities under warming conditions, the ecological niche of L. chinensis was higher than that of other grass species, encroaching on the survival space of other species, reducing the niche overlap of dominant species, and increasing the risk of biodiversity decline and species simplification. This phenomenon may be attributed to the well-developed rhizomes, tolerance to poor soils, and the photosynthetic properties of L. chinensis [39]. L. chinensis can utilize light and resources efficiently and is abundant and distributed in the community [60]; therefore, its ecological niche for each value was relatively higher than that of other grass species. Consistent with this, the growth of S. capillata is inhibited under prolonged warming [61,62,63]. However, the number of shoots in the underground shoot bank of L. chinensis increased under warming conditions [64], and the biomass increased [65,66]. Therefore, L. chinensis is crucial in maintaining the internal environment of the community.

4.2. Regulation of Community Structure and Function by Dominant Species Under Increased Temperature and Water Conditions

The biomasses of L. chinensis and S. krylovii in temperate grassland communities are most sensitive to warming and precipitation changes [66]. Warming and precipitation addition increased the biomass share of L. chinensis and decreased that of S. krylovii over time, and the niche breadths of L. chinensis and S. krylovii were also positively correlated with their biomasses and negatively correlated with the biomass of the other dominant species, suggesting that an increase in the biomass of the dominant species helps them to expand their niche breadth to obtain resources during climate change. In C, the biomass of the dominant species and the niche overlap were mutually inhibitory, which owed to the competition for light among fescue needles and grasses that increased with an increasing biomass, which is consistent with the results of [67]. The effects of warming on species diversity and community stability are primarily through effects on the niche breadth of S. krylovii. Climate warming reduces the temporal stability of biodiversity and community primary productivity (i.e., community stability) [68]. Long-term warming will significantly increase plant community diversity and productivity in all types of alpine grasslands in alpine saline meadows [69]. The effects of climate warming on grassland productivity depend on plant diversity [70]. Global warming in semi-natural grasslands may increase the relative importance of competitive interactions, favoring highly competitive, dominant species [71].
Species richness influences biomass production in grassland communities [72], which is consistent with the results of the control group in this experiment. For our second hypothesis, the niche breadth of L. chinensis increased in warming and watering environments but was detrimental to the species diversity of the community. Warming and precipitation addition increased the biomass of L. chinensis and decreased that of S. krylovii [73]. Increased precipitation offsets the negative effects of warming on plant biomasses in the Tibetan alpine grasslands [74]. The aboveground biomass, belowground biomass, and total biomass of the dominant species increased with increasing precipitation [25]. The niche breadth of L. chinensis had a significant negative effect on species diversity and community stability but was not significantly associated with climate change. This may be because the total, root, and leaf biomasses of S. krylovii are more sensitive to climate change than those of L. chinensis [66]. The high sensitivity of S. krylovii coneflower biomasses is reflected by changes in niche breadth and impacts species diversity and community stability. As an asexual plant, L. chinensis obtains stronger nutrients through a well-developed root system [75] and is relatively insensitive to environmental fluctuations. Changes in the niche breadth of L. chinensis primarily depend on its functional traits rather than climate change. There is a strong negative correlation between species diversity and niche breadth [36]. However, there was a strong positive correlation between species richness and the niche breadth of dominant species, which had a negative effect on niche overlap in the CK group. In contrast, the niche breadth between L. chinensis and S. krylovii under the condition of watering alone had an opposite correlation with that of the C group, and there was a significant positive correlation between the ecological niche overlap and stability of dominant species. This suggests that high precipitation levels prompted the plants to rapidly access nutrients and turnovers, which predicted a wider, rather than a narrower, ecological niche. This finding is corroborated by Schellenberger Costa et al. (2018) [76].

5. Conclusions

Changes in temperature and precipitation affect the niches of dominant species. Through multi-year temperature and control experiments, increased precipitation was found to be crucial in stabilizing the high-level niche of dominant plants in northern grasslands with the occurrence of a differentiation of ecological niches, while the temperature primarily affected the fluctuation of data on ecological niche changes. In the future, a warmer and wetter grassland climate would favor the expansion of the niche breadth of L. chinensis but not the maintenance of that of S. krylovii. Compared with the control, warming reduced the niche breadth of L. chinensis and S. krylovii by 0.03 and 0.17, respectively, and watering increased the niche breadth of L. chinensis and S. krylovii by 0.03 and 0.02, respectively. Warming and watering reduced species diversity, with warming alone reducing diversity by a greater magnitude. Long-term warming affected community stability through the niche breadth of S. krylovii. Increased precipitation had a direct effect on species diversity and community stability in temperate grasslands. In future warming, the response of grasslands to climate change is primarily reflected in an increase in dominance and a decrease in the species diversity index, which is not conducive to the maintenance of the species diversity of a community.
However, in this study, only 20% of the ecological niche of the dominant species was affected by the water-augmentation setup. Therefore, in the future, we need to explore the mechanisms by which different precipitation magnitudes (particularly the increase and decrease in precipitation) and warming affect the dominant species and this grassland community. It is unclear whether this rapid increase in the ratio of the ecological niche breadth of the dominant species over time will continue in the future. Therefore, more diverse environmental elements with more in-depth observations of the adaptation mechanisms of species over longer periods are required to more accurately explain a community’s adaptation to climate change.

Supplementary Materials

The following supporting information can be downloaded at: https://www.mdpi.com/article/10.3390/agronomy14102425/s1, Figure S1: A schematic diagram of the experimental arrangement; Figure S2: Changes of niche values of dominant species over time; Table S1: Productivity estimation models for different species. B indicates productivity (g m−2), H indicates height (cm), and D indicates plant density (number/m2).

Author Contributions

Z.W., conceptualization; Z.W. and R.G., methodology; Z.W., R.G. and X.Z., investigation. Z.W. and X.Z., writing—original draft; X.C. (Xuemeng Chen), L.D., X.Z., X.C. (Xi Chun), H.Z. and W.Z., writing—review and editing. All authors have read and agreed to the published version of the manuscript.

Funding

This research was funded by the National Natural Science Foundation of China (grant number 32260278 and 42161023) and First Class Discipline Research Special Project of Inner Mongolia Normal University (grant number YLXKZX-NSD-037).

Data Availability Statement

Data will be made available upon request.

Conflicts of Interest

The authors declare no conflicts of interest.

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Figure 1. This figure shows the geographical position of Inner Mongolia in China and the Xilingol League in Inner Mongolia. Our survey plots are also marked in the geographical map. On the right is a picture of the experimental plot.
Figure 1. This figure shows the geographical position of Inner Mongolia in China and the Xilingol League in Inner Mongolia. Our survey plots are also marked in the geographical map. On the right is a picture of the experimental plot.
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Figure 2. Niche breadth of L. chinensis (a) and S. krylovii (b), the niche overlap between L. chinensis and S. krylovii (c), and the ratio of niche breadth between L. chinensis and S. krylovii (d) under different treatment conditions. C: control; W: warming; WP: warming plus precipitation addition; P: precipitation addition; dots: The annual average of each index; lines: Normal curves for each type of data.
Figure 2. Niche breadth of L. chinensis (a) and S. krylovii (b), the niche overlap between L. chinensis and S. krylovii (c), and the ratio of niche breadth between L. chinensis and S. krylovii (d) under different treatment conditions. C: control; W: warming; WP: warming plus precipitation addition; P: precipitation addition; dots: The annual average of each index; lines: Normal curves for each type of data.
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Figure 3. Changes in the relative biomass of L. chinensis and S. krylovii under different treatment conditions (L. chinensis and S. krylovii biomass as a proportion of community biomass) (a). Effects of warming and precipitation addition on the importance value (IV) of L. chinensis and S. krylovii and species diversity (b). C: control; W: warming; WP: warming plus precipitation addition; P: precipitation addition.
Figure 3. Changes in the relative biomass of L. chinensis and S. krylovii under different treatment conditions (L. chinensis and S. krylovii biomass as a proportion of community biomass) (a). Effects of warming and precipitation addition on the importance value (IV) of L. chinensis and S. krylovii and species diversity (b). C: control; W: warming; WP: warming plus precipitation addition; P: precipitation addition.
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Figure 4. Correlation heatmap between the ecological niches of dominant species and community factors under C (a), W (b), WP (c), and P (d) conditions (statistical significance: * p ≤ 0.05, ** p ≤ 0.01, and *** p ≤ 0.001). LCIV: importance value of L. chinensis; SKIV: importance value of S. krylovii; LCB: biomass of L. chinensis; SKB: biomass of S. krylovii; LCND: niche breadth of L. chinensis; SKND: niche breadth of S. krylovii; NO: niche overlap between L. chinensis and S. krylovii; CS: community stability; SD: species diversity. C: control; W: warming; WP: warming plus precipitation; P: precipitation addition.
Figure 4. Correlation heatmap between the ecological niches of dominant species and community factors under C (a), W (b), WP (c), and P (d) conditions (statistical significance: * p ≤ 0.05, ** p ≤ 0.01, and *** p ≤ 0.001). LCIV: importance value of L. chinensis; SKIV: importance value of S. krylovii; LCB: biomass of L. chinensis; SKB: biomass of S. krylovii; LCND: niche breadth of L. chinensis; SKND: niche breadth of S. krylovii; NO: niche overlap between L. chinensis and S. krylovii; CS: community stability; SD: species diversity. C: control; W: warming; WP: warming plus precipitation; P: precipitation addition.
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Figure 5. Structural equation modeling of ecosystems using warming and precipitation addition through ecological niche width of dominant species (* p ≤ 0.05, ** p ≤ 0.01, and *** p ≤ 0.001). Negative and positive effects are indicated by red and green line.
Figure 5. Structural equation modeling of ecosystems using warming and precipitation addition through ecological niche width of dominant species (* p ≤ 0.05, ** p ≤ 0.01, and *** p ≤ 0.001). Negative and positive effects are indicated by red and green line.
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Zhang, X.; Wan, Z.; Gu, R.; Dong, L.; Chen, X.; Chun, X.; Zhou, H.; Zhang, W. Warming Increases Ecological Niche of Leymus chinensis but Is Detrimental to Species Diversity in Inner Mongolia Temperate Grasslands. Agronomy 2024, 14, 2425. https://doi.org/10.3390/agronomy14102425

AMA Style

Zhang X, Wan Z, Gu R, Dong L, Chen X, Chun X, Zhou H, Zhang W. Warming Increases Ecological Niche of Leymus chinensis but Is Detrimental to Species Diversity in Inner Mongolia Temperate Grasslands. Agronomy. 2024; 14(10):2425. https://doi.org/10.3390/agronomy14102425

Chicago/Turabian Style

Zhang, Xingbo, Zhiqiang Wan, Rui Gu, Lingman Dong, Xuemeng Chen, Xi Chun, Haijun Zhou, and Weiqing Zhang. 2024. "Warming Increases Ecological Niche of Leymus chinensis but Is Detrimental to Species Diversity in Inner Mongolia Temperate Grasslands" Agronomy 14, no. 10: 2425. https://doi.org/10.3390/agronomy14102425

APA Style

Zhang, X., Wan, Z., Gu, R., Dong, L., Chen, X., Chun, X., Zhou, H., & Zhang, W. (2024). Warming Increases Ecological Niche of Leymus chinensis but Is Detrimental to Species Diversity in Inner Mongolia Temperate Grasslands. Agronomy, 14(10), 2425. https://doi.org/10.3390/agronomy14102425

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