Kou Xinchang (Orcid ID: 0000-0002-6434-6161)

Type of contribution: Research paper

Soil meso-fauna community composition predicts ecosystem

multifunctionality along a coastal-inland gradient of the Bohai Bay

Xinchang Kou a, Yan Tao , Shengzhong Wang

a, b*
, Zhengfang Wu a,
a, c

Haitao Wu d

a
Key Laboratory of Geographical Processes and Ecological Security in Changbai
Mountains, Ministry of Education, School of Geographical Sciences, Northeast
Normal University, Changchun, Jilin Province, China;
b
Jilin Provincial Key Laboratory of Animal Resource Conservation and Utilization,
Changchun, Jilin Province, China;
c
State Environmental Protection Key Laboratory of Wetland Ecology and Vegetation
Restoration, Institute for Peat and Mire Research, Northeast Normal University,
Changchun, Jilin Province, China
d
Key Laboratory of Wetland Ecology and Environment, Northeast Institute of
Geography and Agroecology, Chinese Academy of Sciences, Changchun, Jilin
Province, China

*Corresponding authors: Prof. Yan Tao

School of Geographical Sciences,
Northeast Normal University,
Changchun 130024, China
E-mail address: taoy431@nenu.edu.cn;

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cite this article as doi: 10.1002/ldr.4053
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Abstract

Previous studies have determined the importance of soil biodiversity for maintaining

ecosystem services and multifunctionality. However, the relationship between soil

meso-fauna biodiversity and ecosystem multifunctionality in the real world has yet to

be evaluated. In this study, the relationships between soil meso-fauna community

compositions and ecosystem multifunctionality along a coastal-inland distance

gradient were investigated, including 1 km (1T), 15 km (2T), 30 km (3T), 45 km (4T),

and 60 km (5T) locations. Our results showed that the soil properties were

significantly indicated a decreasing trend after the 3T line. However, the soil

meso-fauna biodiversity indices were observed to have all reached the peak value at

the 4T line. Additionally, it was found that the soil meso-fauna biodiversity indices

were highly correlated with the ecosystem multifunctionality. These findings further

confirmed the fact that the contributions of the soil meso-fauna community

biodiversity indices (59.34%) to ecosystem multifunctionality were as important as, or

even more important than, those of the soil environmental factors (40.66%). Therefore,

it was concluded that the variations in ecosystem multifunctionality were closely

related to the changes in the soil meso-fauna community biodiversity. In other words,

soil meso-fauna community compositions can be effectively used to predict

ecosystem multifunctionality along coastal-inland ecosystems. These findings, are the

first study to provide empirical evidence linking soil meso-fauna community

compositions to ecosystem multifunctionality in real-world coastal-inland ecosystems.

Certainly, future studies which combine multi-trophic levels and consider their
interactions will be able clarify the relationships between the below-ground soil fauna

communities and ecosystem multifunctionality.

Keywords: Multifunctionality; Soil meso-fauna; Biodiversity; Coastal-inland gradient;

Bohai Bay.

1. Introduction

Coastal ecosystems cover only 0.4% of the Earth’s surface area (Walsh, 1988), yet are

critically important environments for soil biodiversity as they provide habitat for

many species of concern (Balakrishnan et al., 2014). In recent years, rising sea levels

have been expected to impact estuarine habitats and push seawater-freshwater

interfaces further inland. Consequently, increasingly salinization levels have been

recognized as serious emerging environmental problems (Morrissey et al., 2014;

Zhang et al., 2019). Soil salinization seems certain to drive changes in soil fauna

(Zheng et al., 2020; Neubauer et al., 2013), as well as plant community compositions

(Sharpe and Baldwin, 2012), which will ultimately alter ecosystem multifunctionality.

At the present time, there are many measurement methods which are commonly

used to access variations in plant/fauna communities and further predict ecosystem

functions (Sechi et al., 2018; Wagg et al., 2018). The earlier studies had mainly

focused on the descriptions of above-ground or below-ground community indices,

including Shannon-Wiener diversity, community richness, community synchrony, and

the population coefficients of variation (CV) (Lehman et al., 2000; Gross et al., 2014;

Wagg et al., 2018). All things considered, there is now a pressing need for moving
beyond mere descriptions of plant/fauna community compositions. Therefore,

ecosystem multifunctionality has been recently highlighted (Wagg et al., 2019;

Delgado-Baquerizo et al., 2020; Jochum et al., 2020). The links between plant and

fauna compositions and ecosystem multifunctionality have been increasing explored.

For example, Wagg et al. (2019) observed that the diversity and complexity of

microorganisms positively influenced multiple ecosystem functions within grassland

ecosystems. Similar results were found by Delgado-Baquerizo et al. (2020), in which

evidence was revealed that multiple ecosystem functions were maintained by soil

biodiversity, and further suggested that the positive association between plant

diversity and multifunctionality across biomes was indirectly driven by soil

biodiversity combining a global observational data. Further, Jochum et al. (2020)

compared the data from two biodiversity experiments (Jena-Germany and

BioDIV-United States), which demonstrated that the discussions drawn from the

experiments of soil biodiversity were robust. Thus, these studies added a new

dimension to earlier observations that the soil biodiversity was matching the multiple

ecosystem functions and predicted the ecosystem multifunctionality. However, the

evidence from the real-world ecosystems continues to be rare or absent, especially for

soil meso-faunas under ongoing global environmental changes.

Soil meso-fauna drive several processes required for plant growth and soil health

(Darby et al., 2011; Yin et al., 2017; Thakur and Geisen, 2019), which are known to

be sensitive to the changes in environmental conditions due to their weak tolerance to

disturbances (Ma et al., 2019). In particular, in the processes of large-scaled

environmental gradients, soil meso-fauna seem to be more representative of the

variations of ecosystem functions (Mitchell et al., 2016). In these regards, from that

perspective, soil meso-fauna are considered to potentially play critical roles in the

accurate predictions of ecosystem multifunctionality. However, none of the

aforementioned studies have attempted to establish the relationships between soil

meso-fauna community compositions and ecosystem multifunctionality, particularly

in real-world coastal-inland ecosystems which tend to accumulate redundant salt in

the soil.

Therefore, in order to close the knowledge gap, this study experimented with

different distance gradients in a coastal-inland ecosystem for the purpose of exploring

the relationships between the compositions of soil meso-fauna communities and

ecosystem multifunctionality. We hypothesized that the changes in soil meso-fauna

community compositions across the coastal-inland ecosystem were consistent with the

ecosystem multifunctionality. Specially, we hypothesized that the indices of the soil

meso-fauna communities, such as the Shannon-Wiener diversity, richness, and

community complexity levels, could be utilized to predict the ecosystem

multifunctionality. The main objective of this study was to investigate whether the soil

meso-fauna community compositions could be effectively used to predict ecosystem

multifunctionality along coastal-inland distance gradients.

2. Materials and methods

2.1 Site description and experimental design

This study’s experimental processes were completed in the coastal areas of Bohai Bay,

Huanghua, Hebei Province, China (38°09′-38°39′N; 117°05′-117°40′E), as detailed in

Fig. A1. The conditions of the experimental site were described in detail in our

previous study (Zheng et al., 2020).

The soil meso-faunal samples examined in this study were collected in July

(summer) of 2019. Five sampling lines were selected across the coastal area toward

the inland region. The sampling sites were identified as natural ecosystems, with the

dominate species determined to be Phragmites australis. The ranges of distance were

as follows: 1 km (1T); 15 km (2T); 30 km (3T); 45 km (4T); and 60 km (5T),

respectively. In that regard, this study preferred to assume that the coastal-inland

gradients were the main driving force affecting the ecosystem functions, which was

determined by the degrees of soil salinization within each range.

2.2 Soil sampling and analysis

In each sampling line, four sampling plots (20 × 20 m) were randomly selected and

each of the plots were consistently spaced at least 1 km apart (Fig. A1). Then, four

subplots (20 cm × 10 cm × 10 cm, (length × width × depth)) were randomly collected

in each sampling plot. In total, 80 fresh samples (5 lines × 4 plots × 4 subplots) were

collected and stored under 4°C refrigerated conditions until required for further

analysis. The samples were divided into two parts, which corresponded to analyses of

the soil properties and the extractions of the soil meso-fauna communities,

respectively. The measurement methods for the above-ground plant biomass (PB),
plant coverage (PC), and soil properties were described in detail in our previous study

(Zheng et al., 2020).

2.3 Soil meso-fauna extraction and identification

For the purpose of extracting the soil meso-fauna communities, the selected soil

samples (10 cm × 10 cm × 10 cm) were placed in Tullgren funnel extractors for 48

hours. Then, a 75% ethanol solution was used to preserve the extracted meso-fauna.

The abundance of meso-fauna was determined using a stereoscopic microscope

(OLYMPUS SZX16) and then identified at the family (or suborder) level (Yin, 2000a;

Yin, 2000b).

2.4 Statistical analysis

In the present research investigation, SPSS19 statistical software was used for the data

analysis processes, and the Shapiro-Wilk Method was adopted to test for normality.

Differences at the P < 0.05 level were considered to be statistically significant. In

addition, a Tukey’s HSD test was used if the main effects were found to be significant.

Also, a Mantel test (Spearman's rank correlation) was performed for the purpose of

estimating the effects on the soil meso-fauna communities of the above-ground and

below-ground environmental factors. Then, aggregated boosted tree (ABT) analysis

was performed in order to quantify the relative effects of the environmental factors on

ecosystem multifunctionality using the “gbmplus” package in R v.2.7.1 software

(De'ath, 2007; Wang et al., 2020).

 The soil meso-fauna richness (number of families), abundance (number of

individuals), and Shannon-Wiener diversity (H′) were calculated for each plot. The

calculations of the communities’ synchrony and population CV were based on the

studies conducted by Gross et al. (2014) and Wagg et al. (2018), respectively. The

following equations were utilized in this study:

𝑃𝑃

𝐻𝐻′ = − � 𝑛𝑛𝑖𝑖 ⁄𝑁𝑁 × ln(𝑛𝑛𝑖𝑖 ⁄𝑁𝑁)

𝑖𝑖=1

𝑆𝑆𝑆𝑆𝑆𝑆𝑆𝑆ℎ𝑟𝑟𝑟𝑟𝑟𝑟𝑟𝑟 (𝜂𝜂) = (1/𝑛𝑛) � 𝑐𝑐𝑐𝑐𝑐𝑐𝑐𝑐 �𝑌𝑌𝑖𝑖 , � 𝑌𝑌𝑗𝑗 �

𝑖𝑖 𝑗𝑗≠1

Where H′ indicates the Shannon-Wiener diversity; P represents the number of

families; ni is the number of each family’s individuals within a sample; N denotes the

total number of individuals in a sample; and Yi represents the abundance of family i in

the meso-fauna community of the n family.

In addition, in order to determine the effects of the coastal-inland gradients on the

soil meso-fauna associations, the Spearman’s rank correlation matrix based on the

abundance data of the meso-fauna was depicted through network analysis using

Cytoscape software, version 3.7.1 (Shannon et al., 2003; Morriën et al., 2017). The

complexity index was calculated as the linkage density among the meso-fauna taxa

(Wagg et al., 2019). The network topological features were then calculated in the

“igraph” package (Zhang et al., 2018).

In order to obtain a quantitative index of the ecosystem multifunctionality for each

line, this study standardized each of the 16 measured functions (for example, PC, PB,

SM, SOM, NO3-N, pH, AP, AK, TN, PPT, CEC, Na+, SO42-, K+, Cl-, and Mg2+) using
a Z-score transformation method. Then, these standardized functions were averaged in

order to calculate the multifunctionality index. The aforementioned method had been

widely used in previous reports regarding multifunctionality research

(Delgado-Baquerizo et al., 2016; Wagg et al., 2019).

A structural equation model (SEM) was used to explore how the soil meso-fauna

community affected the ecosystem multifunctionality. The SEM was based on this

study’s predictions and reviews of relevant literature (Grace, 2006). In this study,

Amos 17.0 software was used to construct the SEM (Arbuckle, 2006).

3. Results

3.1 Soil physiochemical properties

The soil moisture (SM) and pH results showed an obvious decreasing tendency from

the coastal areas to the inland region, and the highest and lowest values were found at

the 1T and 5T lines, respectively. The total nitrogen (TN), soil available K (AK), soil

salinity (PPT), and soil cation exchange capacity (CEC) were found to be

significantly higher at the 1T, 2T, and 3T lines, when compared with the content

levels at the 4T and 5T lines. Moreover, among the different sampling lines, the soil

organic matter (SOM) and the plant biomass (PB) were significantly higher at the 3T

line than at the other lines (P < 0.05). Also, higher values of plant coverage (PC) were

found at the 3T and 4T lines when compared with those observed at the 1T, 2T, and

5T lines, as detailed in Table 1.

3.2 Soil meso-fauna community compositions

Generally speaking, the dominant taxa of the soil meso-fauna in the examined

coastal-inland gradients were Actinedida (36.35%), Gamasida (12.61%), and

Oribatida (12.11%). The coastline gradients had significant effects on the total soil

meso-fauna abundance, richness, Shannon-Wiener diversity, synchrony, and

population CV (P < 0.05). This study found that the total richness of the soil

meso-fauna was significantly higher at the 2T, 3T, and 4T lines, when compared with

the 1T and 5T lines (P < 0.05). However, the complete opposite trend was observed in

the community synchrony (Fig. 1). In addition, the soil meso-fauna abundance and

Shannon-Wiener diversity were observed to be significantly influenced by the

distance gradients, with the highest value found at the 4T line (Fig. 1). The population

CV of the soil meso-fauna communities was significantly lower at the 4T line when

compared with that at the 1T line (P < 0.05). Furthermore, the results of this study’s

network analysis showed that the community complexity and key topological features

(for example, nodes, edges, average degree, and average clustering coefficient) had

increased with the distance gradients from the coastal areas. However, these had

sharply declined at the 5T line, as detailed in Fig. 2 and Table A1. The analysis results

of the mantel test indicated that the variations in the soil meso-fauna community

compositions were significantly correlated with the SOM, TN, AP, AK, NO3-N, PPT,

Na+, SO42-, PB, and PC (P < 0.05) (Table 2).

3.3 Correlations between the soil environment, meso-fauna community indices and
multifunctionality

This study’s experimental results indicated that the soil multifunctionality was

positively correlated with the richness, abundance, Shannon-Wiener diversity, and

community complexity of the soil meso-fauna, and negatively correlated with

population CV (P < 0.05). However, no significant correlations were observed

between the multifunctionality and meso-fauna community synchrony (Fig. 3). The

ABT analysis results were employed to interpret the relative importance of the

above-ground and below-ground factors with regard to the ecosystem

multifunctionality. The community complexity (17.89%), richness (15.04%), PC

(12.63%), and Shannon-Wiener diversity (12.18%) were identified as the primary

factors affecting the ecosystem multifunctionality (Fig. 4). The SEM revealed how the

soil meso-fauna communities effected the ecosystem multifunctionality (χ2 = 7.055; df

= 6; GFI = 0.906; CFI = 0.975; TLI = 0.912; SRMR = 0.007; RMSEA = 0.016; and P

= 0.316) (Fig. 5). The P-value was higher than 0.05 and we considered the model to

be acceptable. Specifically, the soil multifunctionality was determined to be positively

related to the diversity of the soil meso-fauna communities. This was followed by the

community abundance and complexity (P < 0.05; Fig. 5). The soil meso-fauna

abundance was found to be significantly negatively associated with the population

synchrony and population CV. Overall, this study found that the coastal distance

gradients had indirectly increased the soil multifunctionality by increasing the

abundance, diversity, and complexity of the soil meso-fauna, as shown in Fig. 5.

4. Discussions

4.1 Effects of the coastal distances on the soil properties and meso-fauna community

compositions

The results obtained in this study demonstrated that the soil properties had

significantly changed along the coastal gradients. In particular, there was an obvious

trend observed at the 3T line (highest or lowest value). However, the soil meso-fauna

biodiversity indices, including richness, abundance, and Shannon-Wiener diversity,

were all observed to be the highest at the 4T line. In those cases, and perhaps

counter-intuitively, the results seemed to contradict the theory that the soil

environmental conditions impacted the soil biodiversity (Yin et al., 2017; Wagg et al.,

2018). At the very least, there was some inconsistency in the acquired results. For

example, in our previous study, the results of the same experiment suggested that the

soil environment variations along the coastal gradients could potentially affect the soil

macro-faunal communities (Zheng et al., 2020). In other words, the soil macro-faunal

diversity had matched the soil environment variations, among which the peak values

were all observed at the 3T line (Zheng et al., 2020). This study suggested the

inconsistencies in the experimental results may have been largely due to the different

sensitivities of the soil macro- or meso-fauna to the soil environmental conditions. In

addition, previous related studies have also indicated that the growth of soil

meso-fauna may require more suitable soil environmental conditions (Wu et al., 2014;

Mitchell et al., 2016). It was important to note that the mantel-test analysis results

assisted our conjecture, revealing that not all of the environmental variables had
significant effects on the soil meso-fauna communities (Table 2). Meanwhile, the soil

properties which were found to have significant influencing effects, such as the PC,

PB, SOM, PPT, Na+, and SO42-, also made major contributions to the ecosystem

multifunctionality (Fig. A2). Therefore, it was concluded that the 4T line with its

moderate environment, particularly due to the fact that it had a lower PPT than the 3T

line, may have benefited the increases in the soil meso-fauna biodiversity (Morrissey

et al., 2014). This study’s network visualization of the interactions within the soil

meso-fauna communities showed that such network topology characteristics as nodes,

edges, complexity levels, and average degrees, were all highest at the 4T line, thereby

making major contributions to the ecosystem multifunctionality (Fig. 2). Similar

results were found in the study conducted by Wagg et al. (2019), in which it was

demonstrated that greater complexity among the bacterial and fungal communities

also supported multifunctionality. This study proposed that the higher complexity

levels of soil meso-fauna communities will potentially predict higher ecosystem

multifunctionality. Additionally, it was found that the community synchrony and

population CV showed opposite trends to the meso-fauna biodiversity and complexity.

Therefore, this study suggested that in all likelihood, increased community

complexity will contribute to the connectivity and complementarity of soil

meso-fauna communities, but may also result in decreases in synchrony and

population CV (Loreau and de Mazancourt, 2008; Gross et al., 2014).

4.2 Exploring the effects of the soil meso-fauna community compositions on the
ecosystem multifunctionality

In the present study, the results revealed that ecosystem multifunctionality was

significantly positively correlated with meso-fauna richness, abundance,

Shannon-Wiener diversity, and community complexity (Fig. 3). The ABT analysis

results had further indicated that the soil meso-fauna community indices were as

important as, or more important than, the other multifunctionality predictors, such as

community complexity and richness (Fig. 4). Consequently, this study suggested that

the soil meso-fauna community compositions could be used to accurately predict

multifunctionality along coastal-inland ecosystems. This was determined to be in line

with earlier studies which revealed that the diversity of soil fauna communities

demonstrated the importance of biodiversity for multi-functions, as well as to

maintain multifunctionality (Cardinale et al., 2011; Isbell et al., 2011). Furthermore,

this study’s SEM results further extended the aforementioned viewpoint, since it was

found that the distance gradients had no detectable direct effects on the ecosystem

multifunctionality. However, since the meso-fauna abundance increased with the

distance gradients, that factor was considered to have positive effects on the

multifunctionality (Fig. 5). It has been certainly confirmed that variations in

abundance will definitely influence community compositions via competitive or

compensatory effects (Lehman and Tilman., 2000; Hallett et al., 2014). In other words,

higher abundance levels will provide greater insurance that some meso-fauna will

benefit over others through environmental variations in a compensatory manner,

thereby maintaining ecosystem multifunctionality. In support of this theory, this study

found that the soil meso-fauna abundance was negatively related to the population

synchrony and population CV. However, it was also positively related to the

community complexity and diversity, which are known to have indirect effects on

multifunctionality (Fig. 5). Therefore, this study proposed that on one hand,

compensatory dynamics had occurred within the meso-fauna communities which

resulted in the variability reductions with increased abundance. On the other hand, the

increases in abundance had led to higher diversity and complex networks, thereby

ultimately contributing to the multifunctionality. Although previous studies have

linked diversity and complexity to multifunctionality (Maestre et al., 2012;

Delgado-Baquerizo et al., 2016; Wagg et al., 2019), the task of relating soil

meso-fauna community compositions to ecosystem multi-functions is a non-trivial

one, largely due to the unclear regulatory mechanisms.

The results obtained in this experimental study provided, to the best of our

knowledge, the first evidence linking soil meso-fauna community compositions to

ecosystem multifunctionality. Therefore, it was not surprising that those results were

consistent with previous studies which proposed that microbial diversity drives

multifunctionality in terrestrial ecosystems (Wagg et al., 2014; Soliveres et al., 2016).

To some extent, the below-ground soil fauna community compositions should be

considered as critical for maintaining the multifunctionality of natural ecosystems

(Hector and Bagchi, 2007). However, the empirical evidence regarding the soil fauna

which occupy important trophic levels in soil food webs, such as nematodes, protozoa

or even macro-fauna, require further exploration.

5. Conclusions

The results of this study revealed that, similar to what has been determined in

microbial studies, soil meso-fauna community compositions can be utilized to predict

ecosystem multifunctionality in the real-world ecosystems of coastal-inland regions.

The experimental results demonstrated that the predictors of soil meso-fauna

community richness, abundance, and complexity were as important as, or even more

important than, those of the soil environmental factors. In particular, this study’s

proven results should be used to provide a new perspective for further studies

regarding soil biodiversity and multifunctionality. However, in order to advance this

field, the relationships between different trophic groups should also be acknowledged

and their functional traits carefully considered. Overall, the finding obtained in this

research study will advance key ecological topics, such as biodiversity-ecosystem

multifunction relationships in soil meso-fauna communities.

Acknowledgements

This work was supported by the National Key Joint Fund of National Natural Science

Foundation of China (U20A2083, U19A2023), the Fundamental Research Funds for

the Central Universities (2412021QD017) and the Jilin Provincial Key Laboratory of

Animal Resource Conservation and Utilization (130028687).

Declaration of competing interest

The authors declare that they have no known competing financial interests or personal

relationships that could have appeared to influence the work reported in this paper.

Data available on request from the authors.

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Tables
Table 1 Soil properties and vegetation characteristics along coast-inland lines (mean ± SE).

Sampling lines

1T 2T 3T 4T 5T

SM (%) 27 ± 2.97 a 21 ± 2.34 b 20 ± 1.71 b 19 ± 0.27 b 15 ± 0.96 c

SOM (g/kg) 13.29 ± 0.29 b 14.06 ± 0.15 b 17.07 ± 0.62 a 11.99 ± 0.45 c 10.95 ± 0.12 c

pH 8.26 ± 0.05 a 8.21 ± 0.21 ab 8.17 ± 0.01 ab 8.11 ± 0.02 b 8.09 ± 0.02 b

TN (mg/kg) 610.11 ± 7.47 a 625.51 ± 3.08 a 645.37 ± 9.16 a 552.12 ± 2.29 b 517.89 ± 1.50 b

AP (mg/kg) 13.63 ± 0.71 b 19.53 ± 0.77 a 23.78 ± 0.46 a 15.07 ± 0.58 b 7.31 ± 0.49 c

AK (mg/kg) 272.03 ± 3.10 a 291.40 ± 1.88 a 297.51 ± 1.92 a 233.04 ± 3.08 b 211.64 ± 2.47 b

CEC (c mol/kg) 14.57 ± 1.86 a 14.22 ± 1.03 a 14.16 ± 1.36 a 13.20 ± 1.32 b 13.19 ± 0.31 b

PPT 0.82 ± 0.01 a 0.81 ± 0.02 a 0.80 ± 0.01 a 0.75 ± 0.01 b 0.74 ± 0.02 b

PB (kg/m2) 84.92 ± 2.95 c 109.15 ± 2.23 b 171.84 ± 2.72 a 147.32 ± 1.79 b 145.73 ± 7.62 b

PC (%) 60 ± 3.00 c 65 ± 2.50 c 86 ± 4.25 a 81 ± 4.00 a 76 ± 4.20 b

Notes: SM, soil moisture; SOM, soil organic matter; TN, soil total N; AP, soil available P; AK, soil available K; CEC, soil cation exchange capacity; PB, plant
biomass; PC, plant coverage. Different superscript lower-case letters represent significant differences at P < 0.05 among treatments as determined by a Tukey’s
post-hoc test.
Table 2 Mantel-test of environmental variables on soil meso-fauna communities.
Environmental variable Meso-fauna community
r P

SM 0.0574 0.336

SOM 0.3791 0.002**

TN 0.3624 0.001**

AP 0.3745 0.002**

AK 0.3082 0.001**

CEC 0.0641 0.240

NO3-N 0.127 0.048*

pH -0.1902 0.900

PPT 0.2085 0.050*

+
Na 0.2963 0.009**
+
K 0.0926 0.110
2+
Mg 0.0374 0.228
2+
Ca 0.0725 0.116
-
Cl 0.0453 0.243

SO4
2- 0.1381 0.046*

PC 0.1416 0.045*

PB 0.1532 0.035*

Notes: SM, soil moisture; SOM, soil organic matter; TN, soil total N; AP, soil available P; AK,

soil available K; CEC, soil cation exchange capacity; NO3-N, nitrate nitrogen; PPT, soil salinity;

PC, plant cover; PB, plant biomass. Significant: *, P < 0.05; **, P < 0.01; ***, P < 0.001.
Figure Captions:

Figure.1 The richness, abundance, Shannon-Wiener diversity, complexity,

asynchrony and population CV of soil meso-fauna communities across coast-inland

gradient. Different lower-case letters represent significant differences at P < 0.05

among treatments as determined by a Tukey’s post-hoc test.

Figure.2 Network visualization of the interaction strengths within soil meso-fauna

community. The size of the circles represents the abundance of meso-faunas, and the

different filled colors represent different groups.

Figure.3 Linear regressions between ecosystem multifunctionality and richness,

abundance, Shannon-Wiener diversity, complexity, asynchrony and population CV of

soil meso-fauna communities.

Figure.4 Aggregated boosted tree (ABT) analysis showed the relative effect of

environmental factor and soil meso-fauna community indices on ecosystem

multifunctionality. PC, plant cover; PB, plant, biomass; SM, soil moisture; PPT, soil

salinity;

Figure.5 Structural equation modeling of soil meso-fauna community indices

contribution to ecosystem multifunctionality (χ2 = 7.055; df = 6; GFI = 0.906; CFI =

0.975; TLI = 0.912; SRMR = 0.007; RMSEA = 0.016; P = 0.316). Numbers next to

the arrows are the standardized path coefficients. The width of the arrows indicates

the strength of the causal influence.

Figure 1.
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Figure 5.