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Journal of Arid Land
Journal of Arid Land  2025, Vol. 17 Issue (10): 1443-1457    DOI: 10.1007/s40333-025-0057-z     CSTR: 32276.14.JAL.0250057z
Research article     
Shrub leaf stoichiometry and its driving factors in the grasslands of the Altay Mountains, Northwest China
MA Xuexi1,2,3,4, JING Wei1,5, LIANG Yuanye1,3, ZHANG Wenbo1,3, LIU Haoyu1,5, YAN Meng2, MAO Jiefei1,3,4, FAN Lianlian1,3,4, LI Yaoming1,3, CHEN Xi1,3, Georgy A LAZKOV6, GAO Yingzhi2,7,*()
1Key Laboratory of Ecological Safety and Sustainable Development in Arid Lands, Xinjiang Institute of Ecology and Geography, Chinese Academy of Sciences, Urumqi 830011, China
2Key Laboratory of Grassland Resources and Ecology of Western Arid Desert Area of the Ministry of Education, College of Grassland Science, Xinjiang Agricultural University, Urumqi 830052, China
3Research Center for Ecology and Environment of Central Asia, Chinese Academy of Sciences, Urumqi 830011, China
4Bayanbulak Alpine Grassland Observation and Research Station of Xinjiang, Bayanbulak 841314, China
5Xinjiang Vocational and Technical College of Construction, Urumqi 830054, China
6Laboratory of Flora, Institute of Biology and Soil Science, Kyrgyz Academy of Sciences, Bishkek 720071, Kyrgyzstan
7Key Laboratory of Vegetation Ecology of the Ministry of Education, State Environmental Protection Key Laboratory of Wetland Ecology and Vegetation Restoration, Institute of Grassland Science, Northeast Normal University, Changchun 130024, China
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Abstract  

Grassland shrub encroachment is a phenomenon that is prevalent in arid and semi-arid regions worldwide, impacting grassland ecosystems in several ways. In the context of escalating climate change and human activities, examining the nutrient and stoichiometric characteristics of Spiraea shrubs in grassland ecosystems, along with their relationships with environmental factors, can yield valuable insights into the nutrient utilization and survival strategies of these shrubs. This, in turn, offers a scientific foundation for developing future conservation measures. This study was conducted in July 2023 in the Altay Mountains, Northwest China, where Spiraea shrubs thrive across five grassland types: temperate steppe desert, temperate desert steppe, temperate steppe, temperate meadow steppe, and mountain meadow. Leaf and soil samples were collected from each grassland type to analyze the concentrations of carbon (C), nitrogen (N), and phosphorus (P), as well as the stoichiometric characteristics of both the leaves and soil. Subsequently, correlation analysis and redundancy analysis (RDA) were conducted to investigate the variations in leaf C, N, and P concentrations and leaf stoichiometry of Spiraea shrubs as well as their influencing factors. The results indicated the presence of significant or highly significant differences (P<0.050) in the leaf C, N, and P concentrations and leaf stoichiometry (C:N, C:P, and N:P ratios) of Spiraea shrubs across the five grassland types. The N:P ratios of Spiraea shrub leaves in the five grassland types ranged from 7.37 to 11.77, suggesting that N availability generally limits the growth of Spiraea shrubs. Results of RDA revealed that the most significant contributors to the C, N, and P concentrations and stoichiometric characteristics of Spiraea shrub leaves were in the following order: soil total N>mean annual precipitation>elevation>soil pH>soil organic C>mean annual temperature. These factors had contribution rates of 35.32%, 13.19%, 10.20%, 8.82%, 8.34%, and 6.48%, respectively. It was determined that soil nutrients have a greater impact on the growth and nutrient accumulation of Spiraea shrubs compared to climatic factors. This study makes an important contribution to the theoretical basis and data support, enabling a deeper understanding of the response mechanisms of shrub species in the grassland ecosystems of the Altay Mountains to climate change.

Key words： Spiraea shrubs      shrub encroachment      leaf stoichiometry      soil stoichiometry      redundancy analysis (RDA)      Altay mountains     
Received: 19 February 2025      Published: 31 October 2025
Corresponding Authors: *GAO Yingzhi (E-mail: gaoyz108@nenu.edu.cn)
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MA Xuexi
JING Wei
LIANG Yuanye
ZHANG Wenbo
LIU Haoyu
YAN Meng
MAO Jiefei
FAN Lianlian
LI Yaoming
CHEN Xi
Georgy A LAZKOV
GAO Yingzhi
Cite this article:

MA Xuexi, JING Wei, LIANG Yuanye, ZHANG Wenbo, LIU Haoyu, YAN Meng, MAO Jiefei, FAN Lianlian, LI Yaoming, CHEN Xi, Georgy A LAZKOV, GAO Yingzhi. Shrub leaf stoichiometry and its driving factors in the grasslands of the Altay Mountains, Northwest China. Journal of Arid Land, 2025, 17(10): 1443-1457.

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http://jal.xjegi.com/10.1007/s40333-025-0057-z     OR     http://jal.xjegi.com/Y2025/V17/I10/1443

Fig. 1 Overview of the study area based on the elevation and locations of field sampling plots in the five grassland types of the Altay Mountains (a), as well as the photos showing the landscape of the five grassland types (b-f). (b), temperate steppe desert (TSD); (c), temperate desert steppe (TDS); (d), temperate steppe (TS); (e), temperature meadow steppe (TMS); (f), mountain meadows (MM).
Grassland type Sampling
plot		 Longitude Latitude Elevation (m) Coverage (%) Patch size (m2) Plant height (cm)
TSD I 87°43′25″E 47°51′16″N 832 4.30±1.32 0.08±0.03 14.62±4.53
II 86°21′52″E 48°12′56″N 736 20.30±1.60 0.21±0.05 19.54±2.50
III 90°03′17″E 46°31′40″N 828 46.40±4.58 0.47±0.13 50.52±1.49
TDS I 87°00′28″E 48°03′44″N 853 7.90±1.99 0.16±0.04 30.61±1.27
II 90°04′47″E 46°30′09″N 1243 21.50±5.77 0.16±0.05 23.97±2.26
III 90°23′41″E 46°35′06″N 1259 42.20±3.70 0.14±0.04 24.84±3.42
TS I 87°34′26″E 48°02′12″N 1262 10.20±3.32 0.15±0.02 19.27±3.02
II 86°43′32″E 48°27′46″N 1110 19.10±2.50 0.19±0.04 18.92±1.98
III 86°44′01″E 48°18′20″N 1266 47.00±8.95 0.61±0.20 49.04±6.97
TMS I 90°15′36″E 46°56′30″N 1533 7.55±3.75 0.07±0.05 17.49±6.42
II 87°00′53″E 48°11′18″N 1191 16.65±2.75 0.09±0.02 23.57±3.33
III 86°23′32″E 48°21′29″N 1251 45.58±5.57 0.52±0.20 38.66±3.25
MM I 90°13′55″E 47°05′16″N 1855 8.30±2.99 0.06±0.01 30.16±6.86
II 88°17′14″E 48°00′14″N 1732 20.62±1.51 0.26±0.10 49.57±8.23
III 88°10′38″E 47°57′39″N 1589 39.24±2.70 0.48±0.04 60.62±6.02
Table 1 Details of the sampling plots for the five grassland types
Grassland type BD (g/cm3) pH SWC (%) EC (μS/cm) SAN (mg/kg)
TSD 1.43±0.06a 6.82±0.35a 2.10±0.72c 30.42±8.95b 61.99±18.56b
TDS 1.38±0.13a 6.88±0.29a 3.31±1.79c 45.08±23.27ab 102.87±61.09b
TS 1.14±0.12b 6.71±0.34a 6.74±2.64b 39.06±20.61ab 150.13±44.36b
TMS 1.16±0.09b 6.76±0.26a 7.63±2.21b 40.88±18.79ab 135.84±28.07b
MM 1.05±0.17b 6.51±0.66a 11.95±6.53a 56.41±29.52a 302.89±250.47a
Grassland type SAP (mg/kg) Clay (%) Silt (%) Sand (%)
TSD 12.57±4.42a 3.96±2.05ab 41.70±20.18b 54.34±22.16a
TDS 9.54±3.63b 3.42±3.35b 37.84±22.17b 58.74±25.49a
TS 4.77±2.23c 5.54±1.23ab 70.52±15.48a 23.94±16.61b
TMS 5.09±2.47c 5.22±2.76ab 60.91±24.89a 33.86±27.39b
MM 6.16±2.46c 6.29±1.46a 78.12±3.47a 15.59±4.65b
Table 2 Soil physical and chemical properties of the five grassland types
Fig. 2 Soil stoichiometric characteristics of five grassland types. (a), SOC (soil organic carbon); (b), STN (soil total nitrogen); (c), STP (soil total phosphorus); (d), soil C:N ratio (ratio of SOC to STN); (e), soil C:P ratio (ratio of SOC to STP); (f), soil N:P ratio (ratio of STN to STP). Different lowercase letters indicate significant differences of soil stoichiometry among the grassland types (P<0.050) based on the Duncan's multiple range test. Bar indicates the standard error of the mean.
Fig. 3 Leaf stoichiometric characteristics of Spiraea shrubs across five grassland types. (a), leaf C (carbon) concentration; (b), leaf N (nitrogen) concentration; (c), leaf P (phosphorus) concentration; (d), leaf C:N ratio; (e), leaf C:P ratio; (f), leaf N:P ratio. Different lowercase letters indicate significant differences of leaf stoichiometry among the grassland types (P<0.050) based on the Duncan's multiple range test. Bar indicates the standard error of the mean.
Fig. 4 Heatmap of correlations of leaf C, N, and P concentrations and stoichiometric characteristics of Spiraea shrubs with environmental factors (including soil properties, climatic factors, and elevation). BD, bulk density; SWC, soil water content; EC, electrical conductivity; SAN, soil available nitrogen; SAP, soil available phosphorus; MAT, mean annual temperature; MAP, mean annual precipitation. Red circle indicates a positive correlation, and blue circle indicates a negative correlation; the larger the circle area, the stronger the correlation. *, P<0.050 level; **, P<0.010 level; ***, P<0.001 level.
Fig. 5 Redundancy analysis (RDA) of leaf C, N, and P concentrations and stoichiometric characteristics of Spiraea shrubs with different environmental factors
Environmental factor Explanatory rate (%) Contribution
rate (%)		 P Environmental factor Explanatory rate (%) Contribution rate (%) P
STN 23.22 35.32 0.002** Soil N:P ratio 1.58 2.37 0.240
MAP 8.74 13.19 0.002** SWC 1.33 2.14 0.322
Elevation 6.68 10.20 0.006** EC 1.02 1.63 0.500
pH 5.84 8.82 0.024* STP 0.83 1.18 0.552
SOC 5.42 8.34 0.010** SAN 0.74 1.01 0.600
MAT 4.23 6.48 0.048* BD 0.58 1.02 0.640
Soil C:N ratio 2.93 4.41 0.110 Soil C:P ratio 0.39 0.61 0.800
SAP 2.26 3.43 0.196
Table 3 Explanation and contribution rates of environmental factors in the variations leaf C, N, and P concentrations and stoichiometric characteristics of Spiraea shrubs
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