
Biodiv Sci ›› 2025, Vol. 33 ›› Issue (11): 25158. DOI: 10.17520/biods.2025158 cstr: 32101.14.biods.2025158
• Original Papers: Plant Diversity • Previous Articles Next Articles
Xiaoqian Ju1,2,3, Yun Tian1,2,3,*(
), Mingze Xu4,5, Yuanmeng Dai6, Manle Li7, Yuhan Zhou1,2, Peng Liu1,2,3, Xin Jia1,2,3, Tianshan Zha1,2,3
Received:2025-04-29
Accepted:2025-09-16
Online:2025-11-20
Published:2025-12-26
Contact:
Yun Tian
Supported by:Xiaoqian Ju, Yun Tian, Mingze Xu, Yuanmeng Dai, Manle Li, Yuhan Zhou, Peng Liu, Xin Jia, Tianshan Zha. How soil environmental factors shape leaf traits in arid-land vegetation[J]. Biodiv Sci, 2025, 33(11): 25158.
| 固沙阶段 Sand-fixation stages | 半固定沙地 Semi-fixed dune (D1) | 固定沙地 Fixed dune (D2) | 土壤结皮固定沙地 Fixed dune with biological soil crusts (D3) | 草本植物固定沙地 Fixed dune with abundant herbaceous plants (D4) |
|---|---|---|---|---|
| 样地图 Plot photograph | ![]() | ![]() | ![]() | ![]() |
| 经纬度 Latitude and longitude | 107.23° N, 37.70° E | 107.23° N, 37.71° E | 107.23° N, 37.71° E | 107.22° N, 37.70° E |
| 海拔 Elevation (m) | 1,549 | 1,541 | 1,508 | 1,549 |
| 土壤容重 Soil bulk density (BD) (g/cm) | 1.54 ± 0.03a | 1.43 ± 0.12b | 1.62 ± 0.03a | 1.57 ± 0.02a |
| 土壤碳氮比 Soil carbon to nitrogen ratio (SC : SN) | 2.53 ± 0.42c | 7.44 ± 1.70a | 3.97 ± 0.65b | 4.84 ± 1.04b |
| 土壤碳磷比 Soil carbon to phosphorus ratio (SC : SP) | 12.63 ± 9.35a | 14.33 ± 4.80a | 13.85 ± 3.90a | 13.41 ± 4.47a |
| 土壤氮磷比 Soil nitrogen to phosphorus ratio (SN : SP) | 4.98 ± 3.63a | 2.02 ± 0.79b | 3.59 ± 1.29ab | 2.95 ± 1.26ab |
| 土壤有机碳含量 Soil organic carbon content (SOC) (g/kg) | 0.75 ± 0.07c | 2.69 ± 1.32a | 1.20 ± 0.27bc | 1.91 ± 0.64ab |
| 土壤温度 Soil temperature (ST) (℃) | 25.18 ± 1.37a | 23.22 ± 2.82a | 24.87 ± 3.06a | 24.47 ± 2.11a |
| 土壤含水量 Soil water content (SWC) (%) | 0.08 ± 0.02a | 0.10 ± 0.03a | 0.03 ± 0.01b | 0.04 ± 0.02b |
| 土壤全氮含量 Soil total nitrogen content (TN) (g/kg) | 0.22 ± 0.03b | 0.31 ± 0.12a | 0.24 ± 0.02ab | 0.30 ± 0.02ab |
| 土壤全磷含量 Soil total phosphorus content (TP) (g/kg) | 0.05 ± 0.02b | 0.19 ± 0.14a | 0.07 ± 0.02b | 0.12 ± 0.06ab |
Table 1 Location information and variation characteristics of soil physicochemical properties across different sand-fixation stages in Mu Us sandy land
| 固沙阶段 Sand-fixation stages | 半固定沙地 Semi-fixed dune (D1) | 固定沙地 Fixed dune (D2) | 土壤结皮固定沙地 Fixed dune with biological soil crusts (D3) | 草本植物固定沙地 Fixed dune with abundant herbaceous plants (D4) |
|---|---|---|---|---|
| 样地图 Plot photograph | ![]() | ![]() | ![]() | ![]() |
| 经纬度 Latitude and longitude | 107.23° N, 37.70° E | 107.23° N, 37.71° E | 107.23° N, 37.71° E | 107.22° N, 37.70° E |
| 海拔 Elevation (m) | 1,549 | 1,541 | 1,508 | 1,549 |
| 土壤容重 Soil bulk density (BD) (g/cm) | 1.54 ± 0.03a | 1.43 ± 0.12b | 1.62 ± 0.03a | 1.57 ± 0.02a |
| 土壤碳氮比 Soil carbon to nitrogen ratio (SC : SN) | 2.53 ± 0.42c | 7.44 ± 1.70a | 3.97 ± 0.65b | 4.84 ± 1.04b |
| 土壤碳磷比 Soil carbon to phosphorus ratio (SC : SP) | 12.63 ± 9.35a | 14.33 ± 4.80a | 13.85 ± 3.90a | 13.41 ± 4.47a |
| 土壤氮磷比 Soil nitrogen to phosphorus ratio (SN : SP) | 4.98 ± 3.63a | 2.02 ± 0.79b | 3.59 ± 1.29ab | 2.95 ± 1.26ab |
| 土壤有机碳含量 Soil organic carbon content (SOC) (g/kg) | 0.75 ± 0.07c | 2.69 ± 1.32a | 1.20 ± 0.27bc | 1.91 ± 0.64ab |
| 土壤温度 Soil temperature (ST) (℃) | 25.18 ± 1.37a | 23.22 ± 2.82a | 24.87 ± 3.06a | 24.47 ± 2.11a |
| 土壤含水量 Soil water content (SWC) (%) | 0.08 ± 0.02a | 0.10 ± 0.03a | 0.03 ± 0.01b | 0.04 ± 0.02b |
| 土壤全氮含量 Soil total nitrogen content (TN) (g/kg) | 0.22 ± 0.03b | 0.31 ± 0.12a | 0.24 ± 0.02ab | 0.30 ± 0.02ab |
| 土壤全磷含量 Soil total phosphorus content (TP) (g/kg) | 0.05 ± 0.02b | 0.19 ± 0.14a | 0.07 ± 0.02b | 0.12 ± 0.06ab |
| 固沙阶段 Sand-fixation stages | 禾本科 Poaceae | 非禾本科 Non-Poaceae | 灌木 Shrub |
|---|---|---|---|
| 半固定沙地 Semi-fixed dune (D1) | 赖草 Leymus secalinus | 虫实 Corispermum puberulum 草木樨状黄芪 Astragalus melilotoides 沙蓬 Agriophyllum squarrosum 乳白黄芪 Astragalus galactites | 黑沙蒿 Artemisia ordosica 蒙古羊柴 Corethrodendron fruticosum 沙芥 Pugionium cornutum |
| 固定沙地 Fixed dune (D2) | 赖草 Leymus secalinus 沙生针茅 Stipa glareosa 隐子草 Cleistogenes caespitosa | 虫实 Corispermum puberulum 狗娃花 Aster hispidus 乳白黄芪 Astragalus galactites 草木樨状黄芪 Astragalus melilotoides 砂珍棘豆 Oxytropis racemosa 大翅蓟 Onopordum acanthium | 黑沙蒿 Artemisia ordosica 华北白前 Cynanchum komarovii 柠条锦鸡儿 Caragana arborescens 达乌里胡枝子 Lespedeza daurica |
| 土壤结皮固定沙地 Fixed dune with biological soil crusts (D3) | 沙生针茅 Stipa glareosa 中华草沙蚕 Tripogon chinensis 隐子草 Cleistogenes caespitosa | 虫实 Corispermum puberulum 角蒿 Incarvillea sinensis 乳白黄芪 Astragalus galactites 狗娃花 Aster hispidus 银灰旋花 Convolvulus ammannii | 黑沙蒿 Artemisia ordosica 蒙古羊柴 Corethrodendron fruticosum 华北白前 Cynanchum komarovii 达乌里胡枝子 Lespedeza daurica |
| 草本植物固定沙地 Fixed dune with abundant herbaceous plants (D4) | 赖草 Leymus secalinus 沙生针茅 Stipa glareosa 隐子草 Cleistogenes caespitosa 沙芦苇 Agropyron mongolicum 中华草沙蚕 Tripogon chinensis | 虫实 Corispermum puberulum 角蒿 Incarvillea sinensis 狗娃花 Aster hispidus 甘草 Glycyrrhiza uralensis 大翅蓟 Onopordum acanthium 苦豆子 Sophora alopecuroides 鹅绒藤 Cynanchum chinense 雾冰藜 Grubovia dasyphylla | 黑沙蒿 Artemisia ordosica 蒙古羊柴 Corethrodendron fruticosum 华北白前 Cynanchum komarovii |
Table 2 Species composition of different plant functional groups across different sand-fixation stages in Mu Us sandy land
| 固沙阶段 Sand-fixation stages | 禾本科 Poaceae | 非禾本科 Non-Poaceae | 灌木 Shrub |
|---|---|---|---|
| 半固定沙地 Semi-fixed dune (D1) | 赖草 Leymus secalinus | 虫实 Corispermum puberulum 草木樨状黄芪 Astragalus melilotoides 沙蓬 Agriophyllum squarrosum 乳白黄芪 Astragalus galactites | 黑沙蒿 Artemisia ordosica 蒙古羊柴 Corethrodendron fruticosum 沙芥 Pugionium cornutum |
| 固定沙地 Fixed dune (D2) | 赖草 Leymus secalinus 沙生针茅 Stipa glareosa 隐子草 Cleistogenes caespitosa | 虫实 Corispermum puberulum 狗娃花 Aster hispidus 乳白黄芪 Astragalus galactites 草木樨状黄芪 Astragalus melilotoides 砂珍棘豆 Oxytropis racemosa 大翅蓟 Onopordum acanthium | 黑沙蒿 Artemisia ordosica 华北白前 Cynanchum komarovii 柠条锦鸡儿 Caragana arborescens 达乌里胡枝子 Lespedeza daurica |
| 土壤结皮固定沙地 Fixed dune with biological soil crusts (D3) | 沙生针茅 Stipa glareosa 中华草沙蚕 Tripogon chinensis 隐子草 Cleistogenes caespitosa | 虫实 Corispermum puberulum 角蒿 Incarvillea sinensis 乳白黄芪 Astragalus galactites 狗娃花 Aster hispidus 银灰旋花 Convolvulus ammannii | 黑沙蒿 Artemisia ordosica 蒙古羊柴 Corethrodendron fruticosum 华北白前 Cynanchum komarovii 达乌里胡枝子 Lespedeza daurica |
| 草本植物固定沙地 Fixed dune with abundant herbaceous plants (D4) | 赖草 Leymus secalinus 沙生针茅 Stipa glareosa 隐子草 Cleistogenes caespitosa 沙芦苇 Agropyron mongolicum 中华草沙蚕 Tripogon chinensis | 虫实 Corispermum puberulum 角蒿 Incarvillea sinensis 狗娃花 Aster hispidus 甘草 Glycyrrhiza uralensis 大翅蓟 Onopordum acanthium 苦豆子 Sophora alopecuroides 鹅绒藤 Cynanchum chinense 雾冰藜 Grubovia dasyphylla | 黑沙蒿 Artemisia ordosica 蒙古羊柴 Corethrodendron fruticosum 华北白前 Cynanchum komarovii |
| 固沙阶段 Sand-fixation stages | 功能群 Functional groups | 相对多度 Relative abundance (%) | 相对盖度 Relative coverage (%) | 相对频度 Relative frequency (%) | Shannon-Wiener 多样性指数 Shannon-Wiener diversity index (H′) | 物种丰富度指数 Species richness index (S) |
|---|---|---|---|---|---|---|
| 半固定沙地 Semi-fixed dune (D1) | 灌木 Shrub | 11 | 94 | 48 | 0.70 ± 0.28b | 5.00 ± 1.00b |
| 禾本科 Poaceae | 3 | 1 | 8 | |||
| 非禾本科 Non-Poaceae | 86 | 5 | 44 | |||
| 固定沙地 Fixed dune (D2) | 灌木 Shrub | 38 | 79 | 32 | 1.86 ± 0.22a | 11.67 ± 0.58a |
| 禾本科 Poaceae | 34 | 2 | 28 | |||
| 非禾本科 Non-Poaceae | 28 | 19 | 40 | |||
| 土壤结皮固定沙地 Fixed dune with biological soil crusts (D3) | 灌木 Shrub | 68 | 98 | 46 | 1.57 ± 0.29a | 8.67 ± 2.08ab |
| 禾本科 Poaceae | 9 | 1 | 20 | |||
| 非禾本科 Non-Poaceae | 23 | 1 | 34 | |||
| 草本植物固定沙地 Fixed dune with abundant herbaceous plants (D4) | 灌木 Shrub | 36 | 89 | 30 | 1.89 ± 0.19a | 12.33 ± 1.15a |
| 禾本科 Poaceae | 35 | 8 | 30 | |||
| 非禾本科 Non-Poaceae | 29 | 3 | 40 |
Table 3 Diversity indices of different plant functional groups across different sand-fixation stages in Mu Us sandy land
| 固沙阶段 Sand-fixation stages | 功能群 Functional groups | 相对多度 Relative abundance (%) | 相对盖度 Relative coverage (%) | 相对频度 Relative frequency (%) | Shannon-Wiener 多样性指数 Shannon-Wiener diversity index (H′) | 物种丰富度指数 Species richness index (S) |
|---|---|---|---|---|---|---|
| 半固定沙地 Semi-fixed dune (D1) | 灌木 Shrub | 11 | 94 | 48 | 0.70 ± 0.28b | 5.00 ± 1.00b |
| 禾本科 Poaceae | 3 | 1 | 8 | |||
| 非禾本科 Non-Poaceae | 86 | 5 | 44 | |||
| 固定沙地 Fixed dune (D2) | 灌木 Shrub | 38 | 79 | 32 | 1.86 ± 0.22a | 11.67 ± 0.58a |
| 禾本科 Poaceae | 34 | 2 | 28 | |||
| 非禾本科 Non-Poaceae | 28 | 19 | 40 | |||
| 土壤结皮固定沙地 Fixed dune with biological soil crusts (D3) | 灌木 Shrub | 68 | 98 | 46 | 1.57 ± 0.29a | 8.67 ± 2.08ab |
| 禾本科 Poaceae | 9 | 1 | 20 | |||
| 非禾本科 Non-Poaceae | 23 | 1 | 34 | |||
| 草本植物固定沙地 Fixed dune with abundant herbaceous plants (D4) | 灌木 Shrub | 36 | 89 | 30 | 1.89 ± 0.19a | 12.33 ± 1.15a |
| 禾本科 Poaceae | 35 | 8 | 30 | |||
| 非禾本科 Non-Poaceae | 29 | 3 | 40 |
Fig. 1 Differences in leaf traits of plant functional groups across different sand-fixation stages. LA, Leaf area; SLA, Specific leaf area; LTD, Leaf tissue density; LDMC, Leaf dry matter content; LCC, Leaf carbon content; LNC, Leaf nitrogen content; LPC, Leaf phosphorus content; LC : LN, Leaf carbon to nitrogen ratio; LN : LP, Leaf nitrogen to phosphorus ratio; LC : LP, Leaf carbon to phosphorus ratio; D1, Semi-fixed dune; D2, Fixed dune; D3, Fixed dune with biological soil crusts; D4, Fixed dune with abundant herbaceous plants. Upper case letters indicate significant differences among different sand-fixation stages and lowercase letters indicate significant differences among different functional groups (P < 0.05).
Fig. 2 Redundancy analysis biplots showing the distribution of leaf trait assemblages in relation to soil environmental factors (A-C), and the relationships between individual leaf traits and soil environmental factors (E-G), across different plant functional groups and sand-fixation stages. BD, Soil bulk density; SOC, Soil organic carbon content; TN, Soil total nitrogen content; TP, Soil total phosphorus content; SC : SN, Soil carbon to nitrogen ratio; SN : SP, Soil nitrogen to phosphorus ratio; SC : SP, Soil carbon to phosphorus ratio; SWC, Soil water content; ST, Soil temperature; LA, Leaf area; SLA, Specific leaf area; LTD, Leaf tissue density; LDMC, Leaf dry matter content; LCC, Leaf carbon content; LNC, Leaf nitrogen content; LPC, Leaf phosphorus content; LC : LN, Leaf carbon to nitrogen ratio; LN : LP, Leaf nitrogen to phosphorus ratio; LC : LP, Leaf carbon to phosphorus ratio; D1, Semi-fixed dune; D2, Fixed dune; D3, Fixed dune with biological soil crusts; D4, Fixed dune with abundant herbaceous plants. Arrows indicate soil environmental factors; circles denote plant leaf traits.
| 土壤环境因子 Soil environmental factors | 禾本科 Poaceae | 非禾本科 Non-Poaceae | 灌木 Shrub | |||
|---|---|---|---|---|---|---|
| R2 | P | R2 | P | R2 | P | |
| 土壤碳氮比 Soil carbon to nitrogen ratio (SC : SN) | 0.54 | < 0.001 | 0.30 | < 0.001 | 0.47 | < 0.001 |
| 土壤容重 Soil bulk density (BD) | 0.31 | < 0.001 | 0.26 | < 0.001 | 0.21 | < 0.001 |
| 土壤有机碳含量 Soil organic carbon content (SOC) | 0.30 | < 0.001 | 0.19 | < 0.001 | 0.34 | < 0.001 |
| 土壤全氮含量 Soil total nitrogen content (TN) | 0.19 | < 0.001 | 0.45 | < 0.001 | 0.27 | < 0.001 |
| 土壤全磷含量 Soil total phosphorus content (TP) | 0.20 | < 0.001 | 0.08 | < 0.005 | 0.18 | < 0.001 |
| 土壤含水量 Soil water content (SWC) | 0.20 | < 0.001 | 0.17 | < 0.001 | 0.21 | < 0.001 |
| 土壤氮磷比 Soil nitrogen to phosphorus ratio (SN : SP) | 0.19 | < 0.001 | 0.12 | < 0.001 | 0.27 | < 0.001 |
| 土壤温度 Soil temperature (ST) | 0.07 | 0.054 | 0.12 | < 0.001 | 0.03 | 0.264 |
| 土壤碳磷比 Soil carbon to phosphorus ratio (SC : SP) | 0.02 | 0.389 | 0.00 | 0.907 | 0.00 | 0.911 |
Table 4 Variance in leaf traits across three plant functional groups explained by soil environmental factors: a redundancy analysis (RDA)
| 土壤环境因子 Soil environmental factors | 禾本科 Poaceae | 非禾本科 Non-Poaceae | 灌木 Shrub | |||
|---|---|---|---|---|---|---|
| R2 | P | R2 | P | R2 | P | |
| 土壤碳氮比 Soil carbon to nitrogen ratio (SC : SN) | 0.54 | < 0.001 | 0.30 | < 0.001 | 0.47 | < 0.001 |
| 土壤容重 Soil bulk density (BD) | 0.31 | < 0.001 | 0.26 | < 0.001 | 0.21 | < 0.001 |
| 土壤有机碳含量 Soil organic carbon content (SOC) | 0.30 | < 0.001 | 0.19 | < 0.001 | 0.34 | < 0.001 |
| 土壤全氮含量 Soil total nitrogen content (TN) | 0.19 | < 0.001 | 0.45 | < 0.001 | 0.27 | < 0.001 |
| 土壤全磷含量 Soil total phosphorus content (TP) | 0.20 | < 0.001 | 0.08 | < 0.005 | 0.18 | < 0.001 |
| 土壤含水量 Soil water content (SWC) | 0.20 | < 0.001 | 0.17 | < 0.001 | 0.21 | < 0.001 |
| 土壤氮磷比 Soil nitrogen to phosphorus ratio (SN : SP) | 0.19 | < 0.001 | 0.12 | < 0.001 | 0.27 | < 0.001 |
| 土壤温度 Soil temperature (ST) | 0.07 | 0.054 | 0.12 | < 0.001 | 0.03 | 0.264 |
| 土壤碳磷比 Soil carbon to phosphorus ratio (SC : SP) | 0.02 | 0.389 | 0.00 | 0.907 | 0.00 | 0.911 |
Fig. 3 UpSet matrix layout of variation partitioning and hierarchical partitioning analysis results to show the relative importance of soil environmental factors on variation of plant traits. (A) Percentage of variation explained by each component (individual effects and common effects of environmental factors). (B) In the dot-matrix plot on the right, each row corresponds to an environmental factor. For each column, the isolated black dot represents the marginal effect of each environmental factor, lines connecting multiple dots represent the common effect among these corresponding environmental factors. Column diagram on the left shows individual effect of each environmental factor (from hierarchical partitioning), its value is equal to its marginal effect plus its average shared common effect with other environmental factors. BD, Soil bulk density; SOC, Soil organic carbon content; TN, Soil total nitrogen content; TP, Soil total phosphorus content; SC : SN, Soil carbon to nitrogen ratio; SN : SP, Soil nitrogen to phosphorus ratio; SC : SP, Soil carbon to phosphorus ratio; SWC, Soil water content; ST, Soil temperature. ***, P < 0.001.
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