Habitat zonation shapes ecosystem multifunctionality and functional trade-offs in farmland across an oasis-desert ecotone in Xinjiang, China

doi:10.1016/j.jaridl.2026.09.003

Journal of Arid Land

2026, Vol. 18

Issue (9): 1524-1550 DOI: 10.1016/j.jaridl.2026.09.003

Research article

Habitat zonation shapes ecosystem multifunctionality and functional trade-offs in farmland across an oasis-desert ecotone in Xinjiang, China

YANG Yan¹, SHEN Liuji², ZHOU Wenjie¹, JIANG Chenfeng², QIN Linfeng¹, WU Yangyang², ZHOU Zhengli²^,^*(

)

¹ School of Life Science and Technology, Tarim University, Alar 843300, China
² School of Horticulture and Forestry, Tarim University, Alar 843300, China

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Abstract

Farmland ecosystems within the oasis-desert ecotone of arid regions play a crucial role in sustaining regional ecological stability and environmental functioning. However, the spatial distribution patterns of ecosystem multifunctionality (EMF) and the key factors associated with its variation across different crop types and habitat zones remain inadequately understood. This study investigated jujube orchards and cotton fields across two contrasting habitat zones, namely the oasis core and the oasis-desert transition zone, in the Alar region of southern Xinjiang Uygur Autonomous Region, China, during the growing season from June to September 2024. Eight ecosystem functional indicators, including three soil nutrient pools, soil water-holding capacity, three herbaceous α-diversity indices, and dust retention per unit leaf area, were quantified and standardized to calculate EMF and subsequently integrated into five ecosystem functional dimensions representing carbon sequestration, nutrient cycling, water storage, plant diversity, and air purification. An integrated analytical framework combining random forest analysis, two-stage partial least squares structural equation modeling (PLS-SEM), and trade-off-synergy analysis was employed to identify dominant predictors associated with EMF, clarify potential pathways linking habitat zone and crop type to EMF, and characterize functional interactions across different habitat zone-crop type systems. Along the gradient from the oasis core to the oasis-desert transition zone, soil organic carbon, total nitrogen, and total phosphorus stocks in both jujube and cotton systems declined by 14.8%-37.2%, 25.0%-51.0%, and 16.9%-24.9%, respectively. Soil water-holding capacity also decreased by 5.3-20.8 mm, whereas dust retention per unit leaf area increased markedly by 16.0%-73.0%. Accordingly, EMF was approximately 10.0% lower in the oasis-desert transition zone than in the oasis core, but remained consistently higher in jujube orchards than in cotton fields within the same habitat zone. Random forest analysis and two-stage PLS-SEM consistently showed that habitat zone regulates EMF primarily through a soil fertility-biodiversity pathway, whereas crop type exerted a weaker effect mainly through soil water dynamics. Trade-off-synergy analysis revealed predominantly synergistic functional relationships in the oasis core, with all pairwise relationships among the five ecosystem functional dimensions in oasis-core cotton fields exhibiting synergy. In contrast, air purification in oasis-core jujube orchards showed clear trade-offs with the other four functional dimensions. Trade-offs became more frequent and pronounced in the oasis-desert transition zone, particularly in transition-zone cotton fields, where reductions in carbon sequestration and water storage are accompanied by enhancements in plant diversity, nutrient cycling, or air purification. Overall, this study elucidates the mechanistic basis of EMF decline and the intensification of functional trade-offs under conditions of farmland marginalization in oasis-desert ecotones. These results provide important scientific support for zone-specific management strategies that emphasize multifunctional co-benefit optimization in the oasis core, while prioritizing the conservation of carbon sequestration and water storage functions in environmentally vulnerable oasis-desert transition zones.

Key words： ecosystem multifunctionality ecosystem functions functional trade-offs habitat zone partial least squares structural equation modeling (PLS-SEM) oasis-desert ecotone

Received: 15 December 2025 Published: 30 September 2026

Corresponding Authors: ^*ZHOU Zhengli (E-mail: zzlzkytd@163.com)

About author: First author contact:

The first and second authors contributed equally to this work. Author contributions

Conceptualization: YANG Yan, SHEN Liuji; Methodology: YANG Yan, SHEN Liuji; Formal analysis: YANG Yan, SHEN Liuji, ZHOU Wenjie, JIANG Chenfeng, QIN Linfeng, WU Yangyang; Writing - original draft preparation: YANG Yan; Writing - review and editing: ZHOU Zhengli, YANG Yan; Funding acquisition: ZHOU Zhengli; Resources: ZHOU Zhengli; Supervision: ZHOU Zhengli. All authors approved the manuscript.

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Cite this article:

YANG Yan, SHEN Liuji, ZHOU Wenjie, JIANG Chenfeng, QIN Linfeng, WU Yangyang, ZHOU Zhengli. Habitat zonation shapes ecosystem multifunctionality and functional trade-offs in farmland across an oasis-desert ecotone in Xinjiang, China. Journal of Arid Land, 2026, 18(9): 1524-1550.

URL:

http://jal.xjegi.com/10.1016/j.jaridl.2026.09.003 OR http://jal.xjegi.com/Y2026/V18/I9/1524

Fig. 1 Overview of the study area and distribution of sampling plots in jujube orchards and cotton fields within the oasis core and the oasis-desert transition zone, overlaid on a fractional vegetation cover (FVC) map. The FVC map was derived from the normalized difference vegetation index (NDVI) using the dimidiate pixel model and was used to visualize the spatial gradient in vegetation cover from the oasis farmland area toward the desert margin. Higher FVC values mainly correspond to vegetation- or farmland-dominated surfaces, whereas lower values mainly correspond to sparsely vegetated or desert surfaces. Regular rectangular patches with high FVC values mainly represent farmland parcels. The remote sensing imagery is based on 30 m spatial resolution data from the Landsat 8 Operational Land Imager (OLI), captured on 12 May 2023 and downloaded in March 2024.

Table 1 Basic characteristics of sampling plots in jujube orchards and cotton fields within the oasis core and the oasis-desert transition zone

Table 2 Ecosystem functional dimensions, component indicators, and formulas used for ecosystem multifunctionality (EMF) calculation

Fig. 2 Herbaceous α-diversity indices and community composition across different habitat zone-crop type systems. (a), species richness; (b), Margalef richness index; (c), Shannon-Wiener diversity index; (d), Pielou's evenness index; (e), relative abundance of dominant herbaceous species. OJ, oasis-core jujube orchards; OC, oasis-core cotton fields; TJ, transition-zone jujube orchards; TC, transition-zone cotton fields. In the bar charts (Fig. 2a), bars represent the standard errors. In the boxplots (Fig. 2b-d), the horizontal line within each box indicates the median; the box boundaries represent the interquartile ranges; the whiskers denote the data range within 1.5 times the interquartile range; and the points indicate the individual observations. Different lowercase letters indicate significant differences in α-diversity indices among habitat zone-crop type systems (P<0.050). In the stacked bar chart (Fig. 2e), different colors correspond to different dominant herbaceous species.

Fig. 3 Individual ecosystem functional indicators across different habitat zone-crop type systems. (a), soil water-holding capacity (SWHC); (b), dust retention per unit leaf area (DRA); (c), soil organic carbon stock (SOCS); (d), soil total nitrogen stock (STNS); (e), soil total phosphorus stock (STPS). In Figure 3a and b, bars represent standard errors, while vertical dotted lines separate the different habitat zone-crop type systems. In Figure 3c-e, bars represent standard errors, and small circles denote individual replicate observations. Different lowercase letters indicate significant differences of ecosystem functional indicators among habitat zone-crop type systems (P<0.050).

Fig. 4 Ecosystem multifunctionality (EMF) and its key predictors and pathways across different habitat zone-crop type systems. (a), EMF across the four habitat zone-crop type systems; (b), relative predictor importance derived from the random forest analysis, expressed as normalized relative importance percentages calculated from the original percentage increase in mean squared error (%IncMSE) values; (c), linear relationships between ln(EMF) and ln-transformed herbaceous α-diversity indices (Shannon-Wiener diversity index, Pielou's evenness index, and Margalef richness index); (d), two-stage partial least squares structural equation model (PLS-SEM) illustrating the pathways through which habitat zone and crop type influence EMF via four lower-order constructs (soil fertility, soil water storage capacity, biodiversity, and air quality regulation). In Figure 4a, points and bars represent the mean and standard errors across replicate plots, respectively. Different lowercase letters indicate significant differences of EMF among habitat zone-crop type systems (P<0.050), and percentage values denote relative changes in EMF between paired habitat zone-crop type systems. In Figure 4b, the percentages shown beside the predictors indicate normalized relative importance values calculated from the original %IncMSE values, and the circles are scaled in proportion to these normalized values, with larger circles corresponding to greater contributions to EMF prediction. R² value indicates the proportion of explained variance for each endogenous construct. In Figure 4c, points, fitted regression lines, and shaded regions represent observed values, linear regression fits, and 95.0% bootstrap confidence intervals (CIs), respectively. R² value indicates the proportion of variance in ln(EMF) explained by each ln-transformed herbaceous α-diversity index. In Figure 4d, values on arrows from habitat zone and crop type to the four lower-order constructs indicate the standardized structural path coefficient (β) data, whereas values on arrows from the four lower-order constructs to EMF represent the second-stage formative weight (w) data. DRA, dust retention per unit leaf area; SRMR, standardized root mean square residual; RMSEA, root mean square error of approximation; CFI, comparative fit index; NFI, normed fit index; NNFI, non-normed fit index; χ²/df, chi-square divided by degrees of freedom; f², effect size.

Fig. 5 Trade-offs and synergies among ecosystem functions across different habitat zone-crop type systems in farmland ecosystems. The diagonal panels display the five ecosystem functional dimensions: plant diversity function (PDF), nutrient cycling function (NCF), water storage function (WSF), carbon sequestration function (CSF), and air purification function (APF). The lower-left panels illustrate the pairwise relationships among standardized ecosystem functional dimensions using the root mean square deviation (RMSD) approach. Points and bars represent the mean functional values and standard errors among replicate plots for each habitat zone-crop type system, respectively. The gray dashed line indicates the equal standardized values of the paired ecosystem functional dimensions. Points closer to the 1:1 dashed line indicate greater balance between paired ecosystem functional dimensions, whereas points farther from the 1:1 dashed line indicate stronger functional imbalance or trade-off intensity. The upper-right panels present the trade-off index (TOI) for each pair of ecosystem functional dimensions. Each 2×2 colored matrix represents the TOI values for a specific ecosystem functional dimension pair across the four habitat zone-crop type systems, with colors corresponding to the system types shown in the legend. Larger absolute TOI values denote stronger trade-offs. Arrows indicate the dominant ecosystem function within each pairwise TOI comparison, defined as the function with the higher standardized value relative to its paired function. To avoid over-interpretation of negligible contrasts, only functional pairs with |TOI|≥0.20 are labeled, whereas cells without numerical values or arrows indicate |TOI|<0.20 and therefore no clear trade-off relationship.

First-stage β values linking habitat zone and crop type to the four LOCs of EMF
Predictor	Response	Estimate type	Estimate	SE	95.0% bootstrap CI	P	Sign stability
Habitat zone	Soil fertility	β	-0.87	0.025	-0.91 to -0.82	<0.001	1.0000
Habitat zone	Soil water storage capacity	β	-0.30	0.048	-0.39 to -0.20	<0.001	1.0000
Habitat zone	Biodiversity	β	-0.90	0.022	-0.93 to -0.85	<0.001	1.0000
Habitat zone	Air quality regulation	β	0.69	0.041	0.60 to 0.77	<0.001	1.0000
Crop type	Soil fertility	β	-0.16	0.059	-0.28 to -0.05	0.007	0.9965
Crop type	Soil water storage capacity	β	-0.80	0.034	-0.87 to -0.73	<0.001	1.0000
Crop type	Biodiversity	β	-0.20	0.049	-0.29 to -0.10	<0.001	1.0000
Crop type	Air quality regulation	β	-0.03	0.073	-0.18 to 0.11	0.618	0.6910
Second-stage w values of the four LOCs in the EMF higher-order construct
Functional dimension	Higher-order construct	Estimate type	Estimate	SE	95.0% bootstrap CI	P	Sign stability
Soil fertility	EMF	w	0.51	0.063	0.38 to 0.62	<0.001	1.0000
Soil water storage capacity	EMF	w	-0.11	0.038	-0.19 to -0.04	<0.001	1.0000
Biodiversity	EMF	w	0.40	0.072	0.27 to 0.55	<0.001	1.0000
Air quality regulation	EMF	w	-0.27	0.044	-0.35 to -0.18	<0.001	1.0000
Pathway-specific indirect effect on EMF through each mediating dimension (calculated as β×w)
Exogenous variable	Mediating dimension	Pathway	Indirect effect	SE	95.0% bootstrap CI	P	Sign stability
Habitat zone	Soil fertility	Habitat zone→soil fertility→EMF	-0.44	0.062	-0.55 to -0.32	<0.001	1.0000
Habitat zone	Soil water storage capacity	Habitat zone→soil water storage capacity→EMF	0.03	0.013	0.01 to 0.06	<0.001	1.0000
Habitat zone	Biodiversity	Habitat zone→ biodiversity→EMF	-0.36	0.068	-0.50 to -0.24	<0.001	1.0000
Habitat zone	Air quality regulation	Habitat zone→air quality regulation→EMF	-0.19	0.036	-0.26 to -0.12	<0.001	1.0000
Crop type	Soil fertility	Crop type→soil fertility→EMF	-0.08	0.028	-0.14 to -0.02	0.007	0.9965
Crop type	Soil water storage capacity	Crop type→soil water storage capacity→EMF	0.09	0.031	0.04 to 0.16	<0.001	1.0000
Crop type	Biodiversity	Crop type→ biodiversity→EMF	-0.08	0.021	-0.12 to -0.04	<0.001	1.0000
Crop type	Air quality regulation	Crop type→air quality regulation→EMF	0.01	0.020	-0.03 to 0.05	0.618	0.6910

Table S1 Bootstrap estimates for the two-stage partial least squares structural equation modeling (PLS-SEM) linking habitat zone and crop type to ecosystem multifunctionality (EMF)


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