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Journal of Arid Land
Journal of Arid Land  2026, Vol. 18 Issue (5): 851-867    DOI: 10.1016/j.jaridl.2026.05.007     CSTR: 32276.14.JAL.20250492
Research article     
Branch architecture of Tetraena mongolica Maxim. controls particle size distribution of nebkha sediments
ZHAI Bo1, DANG Xiaohong2,*(), LIU Jing3, LIU Xiangjie4, CHEN Xiaona4, LIU Yajing4
1 School of Geographical Sciences and Planning, Jining Normal University, Ulanqab 012000, China
2 College of Desert Science and Engineering, Inner Mongolia Agricultural University, Hohhot 010018, China
3 Institute of Water Resources for Pastoral Area of the Ministry of Water Resources of China, Hohhot 010020, China
4 Experimental Center for Desert Forestry, Chinese Academy of Forestry, Bayannur 015200, China
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Abstract  

The formation of desert shrub sand piles (nebkhas) is attributed to the obstruction and subsequent deposition of migrating sand by the shrub itself. However, the relationship between sediment particle size distribution and shrub branch architecture remains inadequately understood. In August 2020, field investigations were conducted on Tetraena mongolica Maxim. shrubs in the Bayan Engger Desert Nature Reserve, located on the Ordos Plateau in Inner Mongolia Autonomous Region, China. Crown morphological parameters of T. mongolica shrubs and associated nebkhas were systematically measured alongside branch architectures. A one-way analysis of variance (ANOVA) was used to identify differences in branch architectures among various levels, while correlation analysis and model fitting were applied to establish the relationship between crown and nebkha morphological parameters. Path analysis was utilized to identify the key branch architectures that influence crown development. Furthermore, sediment redistribution characteristics of nebkhas were quantified, and principal component analysis combined with regression models was utilized to elucidate the contributions of key branch architectures and sensitive particle size fractions to nebkha deposition. Results indicated that the step-by-step branch ratio (SBR) initially increased from the lower branches to the outermost branches before subsequently decreasing. Additionally, branch angle significantly increased (P<0.0500), whereas both the branch length and the ratio of branch diameters (RBD) significantly decreased toward the exterior of the shrub (P<0.0500). Expansion of crown area significantly enhanced nebkha volume, demonstrating a strong linear relationship (P<0.0010). As the primary contact surface for trapping wind-blown sand, the silhouette area of the shrub initially increased and then decreased from bottom to top. Notably, the silhouette area of the 10-30 cm height layer played a crucial role in promoting nebkha volume expansion (P<0.0100). Path analysis further revealed that the key branch architectures promoting crown area expansion were the step-by-step branch ratio between the third-level and fourth-level branches (SBR3:4), followed by the fourth-level branch length (BLL4), the third-level branch angle (BAL3), and the ratio of branch diameters between the fourth-level and third-level branches (RBD4:3). Under the continuous interception of sediments by branches and leaves, the proportion of surface sediment with a particle size of 100.00-250.00 μm reached 51.07%, indicating a significant increase in fine-sized particles. Further analysis confirmed that SBR3:4, BLL4, BAL3, and sediments within the 50.00-100.00 μm particle size range were the primary contributors to nebkha deposition. These results demonstrate that the branch characteristics of T. mongolica shrubs near the ground surface promote fine sediment accumulation and nebkha development by regulating crown expansion. The findings reveal the unique adaptation mechanisms of rare and endangered plants in nebkha microhabitats and provide a scientific basis for ecological windbreak and sand-fixation projects in the desert transition zones of arid and semi-arid regions.

Key words： Tetraena mongolica Maxim.      nebkhas      branch architecture      particle size distribution      nebkha formation      Ordos Plateau     
Received: 10 October 2025      Published: 31 May 2026
Corresponding Authors: *DANG Xiaohong (E-mail: dangxiaohong1986@126.com)
About author: Author contributions

Conceptualization: ZHAI Bo, DANG Xiaohong; Data curation: ZHAI Bo, DANG Xiaohong, LIU Xiangjie; Formal analysis: ZHAI Bo, LIU Jing, CHEN Xiaona; Funding acquisition: ZHAI Bo, DANG Xiaohong; Investigation: ZHAI Bo, DANG Xiaohong, LIU Jing, LIU Xiangjie, CHEN Xiaona, LIU Yajing; Methodology: ZHAI Bo, LIU Jing, LIU Xiangjie, CHEN Xiaona, LIU Yajing; Project administration: DANG Xiaohong; Resources: ZHAI Bo, DANG Xiaohong; Software: ZHAI Bo, LIU Jing, CHEN Xiaona, LIU Yajing; Supervision: DANG Xiaohong; Validation: ZHAI Bo, LIU Jing, LIU Xiangjie, CHEN Xiaona; Visualization: ZHAI Bo, LIU Xiangjie, LIU Yajing; Writing - original draft: ZHAI Bo; Writing - review and editing: ZHAI Bo, DANG Xiaohong, LIU Xiangjie, CHEN Xiaona. All authors approved the manuscript.

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ZHAI Bo
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ZHAI Bo, DANG Xiaohong, LIU Jing, LIU Xiangjie, CHEN Xiaona, LIU Yajing. Branch architecture of Tetraena mongolica Maxim. controls particle size distribution of nebkha sediments. Journal of Arid Land, 2026, 18(5): 851-867.

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http://jal.xjegi.com/10.1016/j.jaridl.2026.05.007     OR     http://jal.xjegi.com/Y2026/V18/I5/851

Fig. 1 Photographs showing the Tetraena mongolica Maxim. (a) and its nebkha (b), the annual wind rose diagram of the study area (c), and a schematic diagram indicating sampling points for collecting sediments from individual nebkhas (d). N, north; S, south; W, west; E, east.
OBR SBR1:2 SBR2:3 SBR3:4
Branch ratio 4.90±0.68 4.79±0.81b 5.79±0.95a 4.51±1.02b
Table 1 Single-factor variance analysis of the overall and stepwise branch ratios of Tetraena mongolica Maxim.
Fig. 2 Branch length and angle (n=60; a) and the ratio of branch diameters (RBD) across the four branch levels (n=60; b) of T. mongolica shrubs. On the x-axis of the left panel, the 1st, 2nd, 3rd, and 4th refer to the first-level to fourth-level branches, respectively. RBD2:1, ratio of branch diameters between the second-level and first-level branches; RBD3:2, ratio of branch diameters between the third-level and second-level branches; RBD4:3, ratio of branch diameters between the fourth-level and third-level branches. Error bars mean standard deviations. Different uppercase letters indicate significant differences in branch length between different branch levels (P<0.0500); different lowercase letters indicate significant differences in branch angle or RBD between different branch levels (P<0.0500).
Fig. 3 Fitting relationship of the crown area of T. mongolica shrubs with nebkha bottom area and volume (a), and correlation analysis between shrub morphological parameters and nebkha morphological indices. ***, P<0.0010 level. The flatter the red ellipse, the stronger the correlation.
0-10 cm height layer 10-30 cm height layer 30-50 cm height layer Total
Silhouette area (m2) 1.30±0.43c 3.10±0.89a 2.38±1.08b 7.48±3.84
Table 2 Silhouette area of each height layer of T. mongolica shrubs and the total shrub silhouette area
Fig. 4 Correlation analysis of shrub silhouette area with nebkha bottom area and volume. SA0-10, SA10-30, and SA30-50 represent the silhouette areas of the 0-10, 10-30, and 30-50 cm height layers of T. mongolica shrubs, respectively. *, P<0.0100 level. The flatter the red ellipse, the stronger the correlation.
Fig. 5 Path analysis showing the relationship between branch architecture and crown area of T. mongolica shrubs. BLL4, fourth-level branch length; SBR3:4, step-by-step branch ratio between the third-level and fourth-level branches; BAL3, third-level branch angle; RBD4:3, ratio of branch diameters between the fourth-level and third-level branches; r, correlation coefficient.
Fig. 6 Comparison of sediment structures among the interior, base, and exterior of the nebkha
Indicator Principal component loading
PC1 PC2 PC3
BLL4 0.87 0.30 0.20
SBR3:4 0.95 0.11 0.10
BAL3 0.91 0.19 0.12
RBD4:3 -0.37 0.11 -0.16
<2.00 μm -0.46 -0.72 0.35
2.00-50.00 μm 0.02 -0.82 0.51
50.00-100.00 μm 0.93 0.07 0.14
100.00-250.00 μm 0.52 -0.10 -0.58
250.00-500.00 μm -0.41 -0.14 -0.71
500.00-1000.00 μm -0.67 0.66 0.33
1000.00-2000.00 μm -0.64 0.69 0.31
Contribution rate (%) 45.44 20.70 14.06
Accumulated contribution rate (%) 45.44 66.14 80.20
Table 3 Principal component analysis (PCA) of nebkha formation based on the branch architectures of T. mongolica shrubs and intercepted sediment fractions
No. Equation R2 P F
1 y1=3.26x1+2.52 0.87 0.0001 143.47
2 y1=1.22x2+4.99 0.68 0.0001 46.22
3 y1=0.48x3-8.93 0.72 0.0001 55.81
4 y2=268066x1-850373 0.64 0.0001 39.57
5 y2=106430x2-712955 0.57 0.0001 29.02
6 y2=106430x3-1619457 0.45 0.0001 18.03
Table 4 Fitting equations relating the main branch architectures of T. mongolica shrubs with sediment particle size and nebkha volume
Fig. S1 Wind tunnel simulation test to examine the effects of T. mongolica shrubs on wind speed, sand transport disturbance, and nebkha accumulation. (a), average acceleration rate of wind speed at different heights behind the shrub cluster when the incident wind speeds are 6.0, 8.0, 10.0, and 12.0 m/s; (b), sediment transport rate at heights ranging from 0 to 50 cm behind the shrub; (c), a small wind shadow sand pile formed beneath and behind the shrub after the wind tunnel simulation test. H, shrub height (H=27.00 cm in this study).
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