Succession of the herbaceous layer in Haloxylon ammodendron woodland in a desert-oasis ecotone of the Hexi Corridor

doi:10.7522/j.issn.1000-694X.2025.00331

Journal of Desert Research ›› 2026, Vol. 46 ›› Issue (1): 184-198.DOI: 10.7522/j.issn.1000-694X.2025.00331

Succession of the herbaceous layer in Haloxylon ammodendron woodland in a desert-oasis ecotone of the Hexi Corridor

Guohua Wang¹^,²(), Xiaoying Zhang¹, Jiaqi Wang¹, Changsheng Shen¹

^1.College of Geographical Sciences，Shanxi Normal University，Taiyuan 030000，China
^2.Linze Inland River Basin Research Station，Northwest Institute of Eco-Environment and Resources，Chinese Academy of Sciences，Lanzhou 730000，China

Received:2025-11-23 Revised:2025-12-23 Online:2026-01-20 Published:2026-03-09

Abstract

Abstract:

The Herbaceous layer in artificial forest ecosystems plays a crucial role in maintaining ecological stability and regulating energy and matter cycling in the desert-oasis ecotone. In this study， we investigated succession of herbaceous plant communities and changes in physicochemical properties of soils at depths of 0-10 cm and 10-20 cm under artificial Haloxylon ammodendron forests of different ages （0， 5， 10， 20， 30 and 40 years） in the desert-oasis ecotone of the Hexi Corridor. The results showed that：（1） With increasing forest age， soil electrical conductivity and nutrient contents （organic matter， total nitrogen and total phosphorus） significantly increased in both soil layers， while soil pH remained alkaline （7.78-8.86）. Soil moisture content in the 0-10 cm layer first increased and then decreased with increasing forest age， reaching its maximum at 30 years. （2） The dominant herbaceous species shifted from Agriophyllum squarrosum on mobile dunes （0 years） to Bassia dasyphylla and A. squarrosum in the early stage （5-10 years）， and to B. dasyphylla in the mid-stage （20-30 years） and Halogeton arachnoideus in the late stage （40 years）. The dominant species in early- and mid-successional stages had strong drought tolerance， with soil-moisture ecological thresholds of 0.437%-1.245% for A. squarrosum and 0.153%-1.560% for B. dasyphylla， whereas the late-successional dominant H. arachnoideus displayed high salt tolerance， with an electrical conductivity threshold of 53.003-179.131 μS·cm^-1. （3） The diversity of herbaceous layer followed an “initial increase followed by decline” pattern， with Margalef， Shannon-Wiener， Simpson， and Pielou reached to the highest level at 30 years. Soil moisture content and electrical conductivity were key factors influencing the successional processes. These findings provide important reference for the restoration and sustainable development of herbaceous layer under desert artificial forests with annual precipitation around 100 mm.

Key words: desert-oasis ecotone of Hexi Corridor, annual herbaceous layer, dominant species, species diversity, soil water-salt ecological threshold

CLC Number:

Guohua Wang, Xiaoying Zhang, Jiaqi Wang, Changsheng Shen. Succession of the herbaceous layer in Haloxylon ammodendron woodland in a desert-oasis ecotone of the Hexi Corridor[J]. Journal of Desert Research, 2026, 46(1): 184-198.

Figures/Tables 16

Fig.1 Sample areas of three dominant annual herbaceous species of Haloxylon ammodendron woodland with different ages

Table 1 Composition of soil particles and salt ions in different ages of Haloxylon ammodendron woodland

深度 /cm	林龄 /a	土壤粒径组成/%			土壤盐离子含量/(g·kg^-1)
深度 /cm	林龄 /a	砂粒(>0.05 mm)	粉粒(0.002~0.05 mm)	黏粒(<0.002 mm)	Na⁺	Ca²⁺	Cl^-	HCO $3 -$
0~10	CK(0)	92.76±0.55^ab	3.56±0.31^c	3.70±0.27^c	0.01±0.00^c	0.03±0.00^b	0.28±0.06^c	0.17±0.00^e
	5	92.20±0.79^ab	2.72±0.22^c	5.07±0.57^b	0.24±0.01^a	0.05±0.02^b	0.87±0.16^b	0.61±0.08^b
	10	93.86±0.09^a	2.87±0.26^c	3.27±0.17^c	0.21±0.02^a	0.05±0.01^b	0.78±0.09^b	0.44±0.02^c
	20	90.25±2.22^bc	5.86±1.53^b	3.89±0.69^c	0.25±0.03^a	0.04±0.01^b	0.69±0.04^b	0.72±0.01^a
	30	87.40±1.54^bc	6.70±0.33^b	5.91±0.20^b	0.13±0.02^b	0.10±0.02^a	0.87±0.17^b	0.31±0.04^d
	40	76.74±2.74^d	11.57±1.90^a	11.68±0.84^a	0.22±0.03^a	0.08±0.01^a	1.12±0.18^a	0.74±0.05^a
10~20	CK(0)	94.90±0.44^a	1.84±0.34^c	3.25±0.28^b	0.04±0.02^b	0.03±0.00^b	0.26±0.15^c	0.18±0.01^b
	5	93.10±0.50^ab	3.23±1.17^bc	3.67±1.57^b	0.20±0.07^b	0.05±0.02^b	0.57±0.20^ab	0.35±0.08^ab
	10	92.52±0.24^b	2.68±0.20^bc	4.80±0.31^b	0.08±0.04^b	0.07±0.02^b	0.41±0.07^bc	0.28±0.06^ab
	20	91.63±2.76^b	3.10±0.19^bc	4.64±3.73^b	0.21±0.05^b	0.07±0.01^b	0.56±0.18^ab	0.40±0.18^a
	30	91.56±0.11^b	3.73±1.79^b	5.34±0.11^b	0.16±0.07^b	0.06±0.01^b	0.43±0.17^bc	0.29±0.01^ab
	40	81.98±0.79^c	7.11±0.53^a	10.91±0.31^a	0.65±0.50^a	0.15±0.09^a	0.78±0.13^a	0.38±0.11^a

Table 1 Composition of soil particles and salt ions in different ages of Haloxylon ammodendron woodland

深度 /cm	林龄 /a	土壤粒径组成/%			土壤盐离子含量/(g·kg^-1)
深度 /cm	林龄 /a	砂粒(>0.05 mm)	粉粒(0.002~0.05 mm)	黏粒(<0.002 mm)	Na⁺	Ca²⁺	Cl^-	HCO $3 -$
0~10	CK(0)	92.76±0.55^ab	3.56±0.31^c	3.70±0.27^c	0.01±0.00^c	0.03±0.00^b	0.28±0.06^c	0.17±0.00^e
	5	92.20±0.79^ab	2.72±0.22^c	5.07±0.57^b	0.24±0.01^a	0.05±0.02^b	0.87±0.16^b	0.61±0.08^b
	10	93.86±0.09^a	2.87±0.26^c	3.27±0.17^c	0.21±0.02^a	0.05±0.01^b	0.78±0.09^b	0.44±0.02^c
	20	90.25±2.22^bc	5.86±1.53^b	3.89±0.69^c	0.25±0.03^a	0.04±0.01^b	0.69±0.04^b	0.72±0.01^a
	30	87.40±1.54^bc	6.70±0.33^b	5.91±0.20^b	0.13±0.02^b	0.10±0.02^a	0.87±0.17^b	0.31±0.04^d
	40	76.74±2.74^d	11.57±1.90^a	11.68±0.84^a	0.22±0.03^a	0.08±0.01^a	1.12±0.18^a	0.74±0.05^a
10~20	CK(0)	94.90±0.44^a	1.84±0.34^c	3.25±0.28^b	0.04±0.02^b	0.03±0.00^b	0.26±0.15^c	0.18±0.01^b
	5	93.10±0.50^ab	3.23±1.17^bc	3.67±1.57^b	0.20±0.07^b	0.05±0.02^b	0.57±0.20^ab	0.35±0.08^ab
	10	92.52±0.24^b	2.68±0.20^bc	4.80±0.31^b	0.08±0.04^b	0.07±0.02^b	0.41±0.07^bc	0.28±0.06^ab
	20	91.63±2.76^b	3.10±0.19^bc	4.64±3.73^b	0.21±0.05^b	0.07±0.01^b	0.56±0.18^ab	0.40±0.18^a
	30	91.56±0.11^b	3.73±1.79^b	5.34±0.11^b	0.16±0.07^b	0.06±0.01^b	0.43±0.17^bc	0.29±0.01^ab
	40	81.98±0.79^c	7.11±0.53^a	10.91±0.31^a	0.65±0.50^a	0.15±0.09^a	0.78±0.13^a	0.38±0.11^a

Fig.2 Physicochemical properties of soil in artificial Haloxylon ammodendron forests with different ages

Fig.3 PCoA analysis of differences in plant community structure among sites of Haloxylonammodendron woodland

Fig.4 Relative abundance of understory herbaceous plants in Haloxylon ammodendron woodland of different ages

Table 2 Species composition and importance values in Haloxylon ammodendron woodland of different ages

物种名	生活型	林龄/a
		0（CK）			5			10			20			30			40
		D	F	IV	D	F	IV	D	F	IV	D	F	IV	D	F	IV	D	F	IV
泡泡刺(Nitraria sphaerocarpa)	S	5.00	0.02	1.71	—	—	—	—	—	—	—	—	—	—	—	—	—	—	—
沙拐枣(Calligonum mongolicunl)	S	3.81	0.69	8.55	2.00	0.04	1.88	1.00	0.03	0.85	1.00	0.07	1.35	—	—	—	3.60	0.17	4.47
梭梭(Haloxylon ammodendron)	S	—	—	—	1.50	0.04	18.61	—	—		1.00	0.04	7.32	—	—		—	—
骆驼蹄瓣(Zygophyllum fabago)	P	—	—	—	—	—	—	—	—	—	7.00	0.02	2.71	22.38	0.27	7.92	—	—	—
雾冰藜(Bassia dasyphylla)	A	62.83^a	0.93^a	32.93^a	32.75^a	0.98^a	36.85^a	98.13^a	1.00^a	61.93^a	87.22^a	0.91^a	38.73^a	45.50^b	1.00^a	21.46^a	88.95^a	0.7^a	57.73^a
沙蓬(Agriophyllum Squarrosum)	A	102.21^a	0.84^a	37.57^a	13.11^a	0.62^a	17.58^a	1.00^b	0.03^b	0.79^b	4.33^b	0.07^b	4.38^b	124.00^a	0.07^b	18.89^a	5.57^b	0.23^b	6.60^b
白茎盐生草(Halogeton arachnoideus)	A	3.00^b	0.24^b	3.93^b	2.80^b	0.11^b	7.54^b	18.55^a	0.67^a	21.52^a	20.50^a	0.76^a	18.65^a	20.89^b	0.63^a	12.33^b	9.40^b	0.33^a	16.29^a
刺沙蓬(Salsola ruthenica)	A	9.47^b	0.33^b	11.47^b	4.50^b	0.04^b	3.58^b	—	—	—	—	—	—	2.0^b	0.10^b	2.07^b	3.0^a	0.03^b	1.86^b
画眉草(Eragrostis pilosa)	A	—	—	—	5.0^b	0.02^b	10.34^b	5.0^b	0.20^b	4.55^b	21.0^a	0.07^b	8.61^b	32.33^b	0.20^b	7.43^b	12.60^b	0.17^b	10.52^b
猪毛菜(Salsola collina）	A	—	—	—	1.63^b	0.16^b	3.63^b	1.36^b	0.37^b	7.71^b	2.60^b	0.11^b	6.02^b	3.43^b	0.23^b	5.63^b	1.00^b	0.07^b	2.53^b
虎尾草（Chloris virgata）	A	3.50^b	0.04^b	3.85^b	—	—	—	2.00^b	0.09^b	1.26^b	2.50^b	0.09^b	9.72^b	123.00^a	0.03^b	17.87^a	—	—	—
虫实（Corispermum macrocarpum）	A	—	—	—	—	—	—	1.0^b	0.03^b	1.39^b	4.33^b	0.07^b	2.51^b	16.00^b	0.10^b	6.41^b	—	—	—
群落密度		189.82^b			63.28^b			128.05^b			151.19^b			389.53^a			124.12^b
灌木层密度		8.81^a			3.50^b			1.00^b			2.00^b			—			3.60^b
草本层密度		181.01^b			59.78^b			127.05^b			149.49^b			389.53^a			120.52^b
灌木层物种数		2			2			1			2			0			1
草本层物种数		5			6			7			8			9			7
草本层优势种		沙蓬+雾冰藜			雾冰藜+沙蓬			雾冰藜+白茎盐生草			雾冰藜+白茎盐生草			雾冰藜+沙蓬			雾冰藜+白茎盐生草

Fig.5 Understory herbaceous plant diversity in Haloxylon ammodendron woodland of different ages

Fig.6 Changes in herbaceous plant diversity index under Haloxylon ammodendron woodland with different ages

Table 3 Correlation analysis of biological traits among three dominant annual herb species under artificial Haloxylon ammodendron forests

	密度	地上生物量	地下生物量	总生物量
密度	1
地上生物量	0.517^**	1
地下生物量	0.641^**	0.908^**	1
总生物量	0.538^**	0.999^**	0.929^**	1

Table 4 Correlations between biomass and soil physicochemical properties in the three dominant annual herbs

物种	土壤理化指标（x）	拟合曲线方程	R²	P
沙蓬	电导率	y=-0.002x²+0.336x-13.049	0.589	0.045
	含水量	y=-23.924x²+39.424x-14.142	0.637	0.029
	pH	y=1.07x-6.994	0.018	0.715
	无机碳	y=-1.257x²+10.198x-19.011	0.332	0.244
	有机碳	y=-0.131x²+1.188x-1.293	0.175	0.510
	全氮	y=10.91x²-10.637x+3.424	0.358	0.212
	全磷	y=598.543x²-314.178x+41.986	0.149	0.568
	全钾	y=0.192x²-6.799x+61.127	0.033	0.889
	水解性氮	y=0.001x²+0.020x-0.083	0.317	0.264
	有效磷	y=0.008x²-0.238x+2.310	0.204	0.451
	速效钾	y=0.003x²-0.520x+25.418	0.270	0.332
	Cl^-	y=-0.002x²+0.556x-39.621	0.200	0.458
	HCO $3 -$	y=-0.065x+5.815	0.098	0.698
	Na⁺	y=0.00005667x²-0.009x+1.383	0.005	0.982
	Ca²⁺	y=-0.042x²+0.521x-0.444	0.001	0.996
雾冰藜	电导率	y=-0.001x²+0.174x-6.967	0.622	0.033
	含水量	y=-3.101x²+5.13x-1.002	0.613	0.036
	pH	y=0.598x-3.832	0.008	0.817
	无机碳	y=0.987x²-8.813x+20.069	0.736	0.018
	有机碳	y=0.963x²-5.892x+9.399	0.768	0.012
	全氮	y=-5.467x²+3.755x+0.264	0.170	0.571
	全磷	y=-33.807x²+29.221x-4.956	0.302	0.341
	全钾	y=0.061x²-2.432x+24.419	0.716	0.023
	水解性氮	y=0.001x²+0.009x+0.408	0.218	0.478
	有效磷	y=-0.011x²+0.238x-0.185	0.331	0.300
	速效钾	y=0.092x-4.532	0.241	0.437
	Cl^-	y=0.036x-0.699	0.265	0.397
	HCO $3 -$	y=-0.036x+2.57	0.286	0.363
	Na⁺	y=-0.038x+2.434	0.508	0.119
	Ca²⁺	y=-0.026x²+0.166x+0.638	0.198	0.516
白茎盐生草	电导率	y=-0.001x²+0.160x-8.201	0.680	0.019
	含水量	y=-9.129x²+15.328x-5.229	0.584	0.046
	pH	y=-3.281x+25.425	0.225	0.197
	无机碳	y=1.543x²-13.975x+32.159	0.506	0.121
	有机碳	y=-0.133x²+0.943x-0.789	0.041	0.882
	全氮	y=-6.963x²+5.108x-0.053	0.028	0.919
	全磷	y=303.944x²-177.117x+26.423	0.100	0.729
	全钾	y=-0.244x²+8.774x-78.036	0.232	0.453
	水解性氮	y=0.003x²-0.125x+1.916	0.029	0.915
	有效磷	y=0.007x²-0.164x+1.14	0.404	0.212
	速效钾	y=0.027x-0.352	0.127	0.665
	Cl^-	y=-0.00004078x²-0.01x+3.366	0.229	0.458
	HCO $3 -$	y=-0.084x+4.098	0.429	0.186
	Na⁺	y=-0.00008733x²+0.023x-0.405	0.196	0.519
	Ca²⁺	y=0.029x²-0.723x+4.911	0.140	0.637

Table 4 Correlations between biomass and soil physicochemical properties in the three dominant annual herbs

物种	土壤理化指标（x）	拟合曲线方程	R²	P
沙蓬	电导率	y=-0.002x²+0.336x-13.049	0.589	0.045
	含水量	y=-23.924x²+39.424x-14.142	0.637	0.029
	pH	y=1.07x-6.994	0.018	0.715
	无机碳	y=-1.257x²+10.198x-19.011	0.332	0.244
	有机碳	y=-0.131x²+1.188x-1.293	0.175	0.510
	全氮	y=10.91x²-10.637x+3.424	0.358	0.212
	全磷	y=598.543x²-314.178x+41.986	0.149	0.568
	全钾	y=0.192x²-6.799x+61.127	0.033	0.889
	水解性氮	y=0.001x²+0.020x-0.083	0.317	0.264
	有效磷	y=0.008x²-0.238x+2.310	0.204	0.451
	速效钾	y=0.003x²-0.520x+25.418	0.270	0.332
	Cl^-	y=-0.002x²+0.556x-39.621	0.200	0.458
	HCO $3 -$	y=-0.065x+5.815	0.098	0.698
	Na⁺	y=0.00005667x²-0.009x+1.383	0.005	0.982
	Ca²⁺	y=-0.042x²+0.521x-0.444	0.001	0.996
雾冰藜	电导率	y=-0.001x²+0.174x-6.967	0.622	0.033
	含水量	y=-3.101x²+5.13x-1.002	0.613	0.036
	pH	y=0.598x-3.832	0.008	0.817
	无机碳	y=0.987x²-8.813x+20.069	0.736	0.018
	有机碳	y=0.963x²-5.892x+9.399	0.768	0.012
	全氮	y=-5.467x²+3.755x+0.264	0.170	0.571
	全磷	y=-33.807x²+29.221x-4.956	0.302	0.341
	全钾	y=0.061x²-2.432x+24.419	0.716	0.023
	水解性氮	y=0.001x²+0.009x+0.408	0.218	0.478
	有效磷	y=-0.011x²+0.238x-0.185	0.331	0.300
	速效钾	y=0.092x-4.532	0.241	0.437
	Cl^-	y=0.036x-0.699	0.265	0.397
	HCO $3 -$	y=-0.036x+2.57	0.286	0.363
	Na⁺	y=-0.038x+2.434	0.508	0.119
	Ca²⁺	y=-0.026x²+0.166x+0.638	0.198	0.516
白茎盐生草	电导率	y=-0.001x²+0.160x-8.201	0.680	0.019
	含水量	y=-9.129x²+15.328x-5.229	0.584	0.046
	pH	y=-3.281x+25.425	0.225	0.197
	无机碳	y=1.543x²-13.975x+32.159	0.506	0.121
	有机碳	y=-0.133x²+0.943x-0.789	0.041	0.882
	全氮	y=-6.963x²+5.108x-0.053	0.028	0.919
	全磷	y=303.944x²-177.117x+26.423	0.100	0.729
	全钾	y=-0.244x²+8.774x-78.036	0.232	0.453
	水解性氮	y=0.003x²-0.125x+1.916	0.029	0.915
	有效磷	y=0.007x²-0.164x+1.14	0.404	0.212
	速效钾	y=0.027x-0.352	0.127	0.665
	Cl^-	y=-0.00004078x²-0.01x+3.366	0.229	0.458
	HCO $3 -$	y=-0.084x+4.098	0.429	0.186
	Na⁺	y=-0.00008733x²+0.023x-0.405	0.196	0.519
	Ca²⁺	y=0.029x²-0.723x+4.911	0.140	0.637

Fig.7 Water-salt response and model fitting results for three dominant annual herbaceous plants

Table 5 Soil water and salt ecological thresholds of three dominant annual herbaceous plants Gaussian regression equation

物种	拟合结果
物种	土壤含水量	土壤电导率
沙蓬	$y = 17.916 e x p - 12 x - 0.841 2 / 0.2022$	$y = 17.916 e x p - 12 x - 84.975 2 / 22.3482$
雾冰藜	$y = 4.312 e x p - 12 x - 0.856 2 / 0.3522$	$y = 4.312 e x p - 12 x - 84.967 2 / 20.9862$
白茎盐生草	$y = 7.166 e x p - 12 x - 0.807 2 / 0.2042$	$y = 7.166 e x p - 12 x - 116.067 2 / 31.5322$

Table 5 Soil water and salt ecological thresholds of three dominant annual herbaceous plants Gaussian regression equation

物种	拟合结果
物种	土壤含水量	土壤电导率
沙蓬	$y = 17.916 e x p - 12 x - 0.841 2 / 0.2022$	$y = 17.916 e x p - 12 x - 84.975 2 / 22.3482$
雾冰藜	$y = 4.312 e x p - 12 x - 0.856 2 / 0.3522$	$y = 4.312 e x p - 12 x - 84.967 2 / 20.9862$
白茎盐生草	$y = 7.166 e x p - 12 x - 0.807 2 / 0.2042$	$y = 7.166 e x p - 12 x - 116.067 2 / 31.5322$

Fig.8 Water-salt ecological thresholds of three dominant annual herbaceous plants

Fig.9 Correlation analysis between diversity indices of understory herbaceous plants and soil environmental factors

Fig.10 RDA ordination plot of species diversity in understory vegetation and soil environmental factors

Fig.11 Conceptual diagram of dominant species in different evolutionary stages

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Succession of the herbaceous layer in Haloxylon ammodendron woodland in a desert-oasis ecotone of the Hexi Corridor

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