Catena

Multifaceted impacts of moss-dominated biocrusts on dryland soils: From soil pore

structure to aeration and water infiltration
--Manuscript Draft--

Manuscript Number: CATENA22400

Article Type: Research Paper

Keywords: Biological soil crust; Pore characteristics; Air permeability; Gas diffusivity; Soil
infiltrability; Chinese Loess Plateau

Abstract: Biocrusts are a crucial living cover of dryland soils that alter near-surface soil structure
and properties. To date, it is still unclear how biocrusts affect soil pore structure, and
subsequently alter gas and water transport processes. In this study, we investigated
aeolian sand and loess soil colonized with moss-dominated biocrusts and in bare state
of the Chinese Loess Plateau. X-ray computed tomography (CT) was used to quantify
and visualize the pore characteristics for all considered scenarios. In addition, the soil
air permeabilities and relative gas diffusivities were measured together with infiltration
rates. CT imaging and reconstruction showed 102%–107%, 56%–87%, and
72%–203% higher macroporosities, surface area densities, and mean pore volumes,
respectively, for soils colonized with biocrusts when compared to their bare
counterparts. The biocrusts also increased the ratio of connected and isolated
porosities as well as pore node densities, while decreasing Euler numbers and
tortuosities when compared to the bare soils. This suggests that the pore networks in
biocrust-colonized soils exhibit higher continuity and less tortuosity. In addition, the
biocrusts increased air permeability by 277%–301% and relative gas diffusivity by
47%–59%, which is attributable to more air-filled pores and better pore connectivity.
However, when compared with bare soils, the biocrusts reduced steady state infiltration
rates by 41%–50% and the 60 min cumulative infiltration amount by 33%–54%. The
decrease in infiltrability is attributable to the higher fine particle content and the
enhanced water-holding capacity of biocrusts. In summary, the biocrusts improved soil
pore structure and soil aeration, and reduced infiltrability when compared to bare soils.
This implies that biocrusts play a multifaceted and complex role in the transport of
gases and water, and as a consequence significantly impact near-surface hydrological
and biochemical processes, such as soil evaporation and respiration in drylands.

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Graphical Abstract
Highlights (for review)

Highlights

 Moss biocrusts averagely increase soil macroporosity by 102%–107%

 Biocrusts have higher pore continuity and less tortuosity as compared with bare soil

 Biocrusts significantly increase air permeability and relative gas diffusivity

 Biocrusts results in 41%–50% lower steady state infiltration rate than bare soil

 Biocrusts induce a complex influence on soil gas and water transport in drylands
Responses to Technical Check Results

Response to Technical Check Results

Dear Editor,

We give our thanks to you for the valuable comments and suggestions on our
manuscript. According to the comments and suggestions, we have thoroughly revised
our manuscript. After careful revision, we would like to resubmit our revised
manuscript and ask for your reconsideration of publication in your journal. We hope
that our revised manuscript would satisfy your requirements and meet the standard of
your journal.

We list our revisions and responses to the comments in the Response to Technical Check
Results.

Thanks for your attention and consideration.

Sincerely yours,

Bo Xiao

On behalf of all the co-authors.

College of Land Science and Technology

China Agricultural University

Address: No. 2, Yuanmingyuan West Road, Haidian District, Beijing 100193, China

Tel.: +86 10 62734558; Fax: +86 10 62734558

E-mail addresses: xiaobo@cau.edu.cn, xiaoboxb@gmail.com

May 26, 2023

Comments:

Q: Abstract should not exceed 300 words. Please revise.

A: Thanks for the comments and suggestions.

According to the comments and suggestions, in the revised files we have

reorganized the abstract within 300 words.
Manuscript including Abstract & Keywords & Title page with
author details

1 Multifaceted impacts of moss-dominated biocrusts on dryland soils:

2 From soil pore structure to aeration and water infiltration

3 Fuhai Suna, Bo Xiaob,c,*, Markus Tullerd

a
4 Key Laboratory of Arable Land Conservation in North China, Ministry of Agriculture and

5 Rural Affairs/ College of Land Science and Technology, China Agricultural University, Beijing

6 100193, China

b
7 State Key Laboratory of Soil Erosion and Dryland Farming on the Loess Plateau, Institute of

8 Soil and Water Conservation, Chinese Academy of Sciences and Ministry of Water Resources,

9 Yangling, Shaanxi 712100, China

c
10 Breeding Base for State Key Laboratory of Land Degradation and Ecological Restoration in

11 Northwestern China/ Key Laboratory of Restoration and Reconstruction of Degraded

12 Ecosystems in Northwestern China of Ministry of Education, Ningxia University, Yinchuan

13 750021

d
14 Department of Environmental Science, The University of Arizona, Tucson, AZ 85721, USA

15

*
Corresponding author at: Key Laboratory of Arable Land Conservation in North China,

Ministry of Agriculture and Rural Affairs/ College of Land Science and Technology, China

Agricultural University, No. 2, Yuanmingyuan West Road, Haidian District, Beijing 100193,

China.

E-mail addresses: xiaobo@cau.edu.cn, xiaoboxb@gmail.com (B. Xiao).

1
16 ABSTRACT

17 Biocrusts are a crucial living cover of dryland soils that alter near-surface soil structure and

18 properties. To date, it is still unclear how biocrusts affect soil pore structure, and subsequently

19 alter gas and water transport processes. In this study, we investigated aeolian sand and loess

20 soil colonized with moss-dominated biocrusts and in bare state of the Chinese Loess Plateau.

21 X-ray computed tomography (CT) was used to quantify and visualize the pore characteristics

22 for all considered scenarios. In addition, the soil air permeabilities and relative gas diffusivities

23 were measured together with infiltration rates. CT imaging and reconstruction showed 102%–

24 107%, 56%–87%, and 72%–203% higher macroporosities, surface area densities, and mean

25 pore volumes, respectively, for soils colonized with biocrusts when compared to their bare

26 counterparts. The biocrusts also increased the ratio of connected and isolated porosities as well

27 as pore node densities, while decreasing Euler numbers and tortuosities when compared to the

28 bare soils. This suggests that the pore networks in biocrust-colonized soils exhibit higher

29 continuity and less tortuosity. In addition, the biocrusts increased air permeability by 277%–

30 301% and relative gas diffusivity by 47%–59%, which is attributable to more air-filled pores

31 and better pore connectivity. However, when compared with bare soils, the biocrusts reduced

32 steady state infiltration rates by 41%–50% and the 60 min cumulative infiltration amount by

33 33%–54%. The decrease in infiltrability is attributable to the higher fine particle content and

34 the enhanced water-holding capacity of biocrusts. In summary, the biocrusts improved soil pore

35 structure and soil aeration, and reduced infiltrability when compared to bare soils. This implies

36 that biocrusts play a multifaceted and complex role in the transport of gases and water, and as

37 a consequence significantly impact near-surface hydrological and biochemical processes, such

38 as soil evaporation and respiration in drylands.

39 Keywords: Biological soil crust; Pore characteristics; Air permeability; Gas diffusivity; Soil

40 infiltrability; Chinese Loess Plateau

2
41 1. Introduction

42 Drylands make up 41% of the Earth’s terrestrial surface and support more than 38% of the

43 world’s population, forming the largest terrestrial biome on Earth (Reynolds et al., 2007; Huang

44 et al., 2015). They are characterized by sparse vegetation with low annual net productivity and

45 are especially vulnerable to climate change and land desertification and degradation. It is

46 predicted that the increasing aridity in the late twenty-first century will globally increase the

47 total area of drylands and exacerbate their desiccation and degradation (Dai, 2013; Huang et al.,

48 2017). This will have negative consequences for multiple structural and functional attributes of

49 dryland ecosystems, such as soil structure, nutrient cycling, plant productivity, and microbial

50 community richness (Berdugo et al., 2020). Biocrusts are particularly prevalent in vegetation

51 interspaces of drylands due to specific adaptation mechanisms, which allow them to cope with

52 insolation, limited precipitation, and severe drought. As a crucial layer between soil and

53 atmosphere, biocrusts cover about 12% of the global terrestrial surface and about 30% of

54 dryland soils. They provide critical functions in dryland ecosystems such as modifying soil

55 structure and thereby influencing hydrologic and nutrient cycles, soil stability, and vascular

56 plant establishment (Rodríguez-Caballero et al., 2018) and provide a barrier against land

57 degradation and desertification (Xiao and Bowker, 2020).

58 Biocrusts are photoautotrophic communities of cyanobacteria, algae, liverworts, liches, and

59 bryophytes that coexist with heterotrophic bacteria, archaea, and fungi, and form an encrusted

60 layer within or on top of the uppermost few millimeters or even centimeters of soil (Li et al.,

61 2022a; Weber et al., 2022). Biocrusts act as ecosystem engineers and regulate various soil

62 processes and functions. They accelerate soil formation (Pointing and Belnap, 2012), reduce

63 wind and water erosion (Bu et al., 2015; Faist et al., 2017), stabilize the soil surface (Felde et

64 al., 2018), govern water and nutrient cycles (Kidron and Büdel, 2014; Heindel et al., 2018),

65 fixate atmospheric carbon and nitrogen (Su et al., 2013; Liu et al., 2016), and promote microbial

3
66 diversity (Maier et al., 2018). The modification of surface soil processes due to biocrusts is

67 mediated via the special physical structure and the biotic components of the biocrust layer.

68 Although biocrusts only occupy a minor proportion of the vertical soil profile, they generate a

69 distinct physical structure that differs from the underlying soil by tightly integrating soil

70 particles and biotic components thereby creating a structure that affects soil geomorphologic,

71 hydrologic, and biological dryland processes (Xiao et al., 2022).

72 Soil structure controls many essential processes occurring in soils, such as gas transport,

73 water infiltration and retention, and solute transport. Structure is also an important factor for

74 root penetrability (Rabot et al., 2018). The quantification of biocrust effects on structure is

75 crucial for comprehending soil functions in drylands. Although there are several studies about

76 biocrust impacts on soil structure, they only analyze bulk density, penetration resistance, and

77 aggregate size distribution to evaluate total porosity, stability, and solid phase arrangement of

78 biocrusts (Felde et al., 2018; Li et al., 2021a; Wang et al., 2021). However, the spatial

79 distribution, morphological and topological soil pore characteristics (e.g., pore size distribution,

80 pore shape, and connectivity) induced by biocrusts are important factors for soil structure

81 development and evolution. Currently, the impact of biocrusts on soil pore characteristics is

82 mainly measured indirectly through soil water retention curves and mercury porosimetry (Felde

83 et al., 2014; Sun et al., 2021). These methods assume idealized regular pore shapes to interpret

84 results, exhibit hysteresis (Rabot et al., 2018), and are not suitable to authentically characterize

85 soil structural pores of biocrusts. X-ray computed tomography (CT) is an established technique

86 capable of rapidly and non-destructively imaging the soil pore system in various directions

87 (Zhou et al., 2017; Tian et al., 2022) to directly visualize and quantify soil structural pores. Yet,

88 X-ray CT has not been fully exploited for soils colonized with biocrusts.

89 The impact of biocrusts on soil pore structure is also crucial for soil gas and water transport

90 processes. The most important soil gas transport parameters, namely air permeability (Ka) and

4
 91 relative gas diffusivity (Dp/D0), the ratio of gas diffusion coefficients of soil and free air, are

92 governed by soil structure, but the pore characteristics affect both in different ways (Martínez

93 et al., 2016). Both Ka and Dp/D0 are also impacted by the soil water content (Hamamoto et al.,

94 2009). The soil gas transport parameters can be used to characterize aeration, which is important

95 for O2 intake and CO2 efflux by soil microorganisms, plant roots, and soil fauna (Hillel, 1998),

96 yet the impact of biocrusts on gas transport parameters is not fully understood.

97 The impact of biocrusts on water transport in soils (e.g., infiltration) has drawn more

98 attention due to limited water resources in dryland ecosystems. The infiltration of water into

99 soils is determined by a combination of soil texture and structure, particle size distribution, pore

100 continuity, and land surface conditions (Sun et al., 2022a). The role of biocrusts in regulating

101 water infiltration varies widely for different regions of the world and related studies generated

102 conflicting results (Kidron, 2021; Yang et al., 2022). This is why much more research is needed

103 to entirely understand how biocrusts influence soil pore structure and consequently gas and

104 water transport processes in dryland ecosystems.

105 We hypothesize that biocrusts significantly alter soil pore structure and as a consequence

106 soil aeration and water infiltration at the soil-atmosphere interface. Within this context, the

107 specific objectives of this study are: (ⅰ) to quantify the effects of biocrusts on soil pore

108 geometrical and morphological characteristics; (ⅱ) to determine the impact of biocrusts on soil

109 gas transport properties (Ka and Dp/D0) for varying soil moisture regimes; (ⅲ) to relate water

110 infiltrability of biocrust-colonized soils to the infiltrability of bare soils; and (ⅳ) to analyze the

111 connection between biocrust-induced changes in soil pore structure, aeration, and infiltrability.

112 To achieve these objectives, we sampled and analyzed the pore structure of both biocrust-

113 colonized and bare aeolian sand and loess soil on the northern Loess Plateau of China by means

114 of X-ray CT and various laboratory and in-situ soil aeration and infiltration experiments. The

115 obtained results provide new insights on how biocrusts alter soil structure and reshape soil gas

5
116 and water balances in dryland ecosystems.

117

118 2. Materials and Methods

119 2.1. Study site

120 This study was conducted in the Liudaogou watershed (38°46′–38°51′N, 110°21′–110°21′E;

121 see Fig. 1A) located on the Chinese Loess Plateau. The watershed covers an area of 6.89 km2

122 and is characterized by a typical semiarid continental monsoon climate with a mean annual

123 precipitation of 454 mm, ~80% of which occurs from June to September (Xiao and Bowker,

124 2020). The average annual potential evaporation is 1337 mm, which is approximately three

125 times the mean annual precipitation. The annual average temperature is 8.4℃, with a mean

126 temperature of 23.7℃ in summer (June to August) and −9.7℃ in winter (December to February)

127 (Xiao et al., 2014). The watershed exhibits a hilly topology with numerous gullies, which is

128 typical for the loess region, and the elevation ranges from 1081 to 1274 m above sea level. It is

129 situated close to the center of the “Water-Wind Erosion Crisscross Region” of the Loess Plateau

130 that exhibits a very fragile ecosystem and previously endured severe soil erosion (i.e., up to

131 10,000 t km−2 yr−1) due to the sparse vegetation cover (Bu et al., 2015).

132 In this area, intensive ecological restoration programs have been implemented in the past

133 decades to recover vegetation by planting a large number of native shrubs to reduce soil erosion

134 (Deng et al., 2017). Common plants in this area include Bothriochloa ischaemum (Linn.) Keng.,

135 Medicago sativa Linn., Artemisia ordosica Krasch., Caragana Korshinskii Kom., Cotoneaster

136 horizontalis Dcne., and Lespedeza davurica (Laxm.) Schindl (Sun et al., 2022b). Consequently,

137 biocrusts naturally developed on the stabilized soil surfaces, and now natural moss-dominated

138 biocrusts are widely distributed across fallow land, grassland, shrubland, and woodland

139 covering ~30% and sometimes up to 70%–80% of the area (Xiao et al., 2019a). The dominant

6
140 moss species are Didymodon vinealis (Brid.) Zander., Bryum argenteum Hedw., and Barbula

141 vinealis Brid. (Sun et al., 2022b).

142 2.2. Experimental design and soil sampling

143 Four treatments in five replicates were established for this study. The treatments include:

144 (ⅰ) bare aeolian sand (Fig. 1B); (ⅱ) biocrust-colonized aeolian sand (i.e., the moss covers about

145 90% of the surface) (Fig. 1C); (ⅲ) bare loess soil (Fig. 1D); and (ⅳ) biocrust-colonized loess

146 soil (i.e., the moss cover is about 90%) (Fig. 1E).

147 For each of the four treatments, five experiments were involved: (ⅰ) biocrust and bare soil

148 properties measurements; (ⅱ) X-ray CT image acquisition; (ⅲ) in-situ measurements of Ka and

149 surface soil moisture; (ⅳ) laboratory measurements of Dp/D0 from saturation to dry; and (ⅴ)

150 in-situ measurement of soil water infiltration.

151 All sampling and in-situ measurements within the experimental site (~0.5 km2) took place

152 in areas with almost flat surface (i.e., slope < 5°) to minimize variations in biocrust

153 characteristics, slope gradients and orientations, and shrub types and densities. Four sampling

154 plots (~5 m × 5 m) were randomly selected to collect undisturbed soil samples in 5-cm tall (100

155 cm3 volume) stainless steel cylinders for bulk density, total porosity, saturated water content,

156 filed capacity, saturated hydraulic conductivity, and relative gas diffusivity (Dp/D0)

157 measurements in the laboratory. In addition, 9-cm diameter and 2-cm high Petri dishes were

158 used to sample the biocrust layers for measurement of biocrust thickness, moss density, total

159 biomass, and chlorophyll a content. Furthermore, 5-cm diameter and 5-cm tall polyvinyl

160 chloride (PVC) cylinders were carefully pushed into the soil and excavated to collect

161 undisturbed soil cores for X-ray CT imaging. Lastly, bulk soil samples were collected from 0–

162 5 cm depth, air-dried at room temperature, and passed through 2 mm and 0.149 mm mesh sieves

163 for soil physicochemical analysis that included soil particle size distribution, soil pH, and

7
164 organic matter content.

165 2.3. Measurement of biocrust characteristics and soil properties

166 Prior to sampling, all plots were photographed to determine the moss-covered area. The

167 obtained images were analyzed with ImageJ (Schneider et al., 2012), as described in Sun et al.

168 (2022a). The surface roughness of the plots was measured with the chain method (Jester and

169 Klik, 2005). In the laboratory, the biocrust layer was carefully separated from the underlying

170 soil, and its thickness was measured with a digital Vernier caliper. The moss was separated from

171 the soil by washing the samples through a 2-mm sieve with water and then dried at 65 ℃ for 24

172 h to measure moss biomass (Xiao et al., 2019b). The moss density was calculated from the total

173 moss gametophytes in a 4 cm2 sample (Xiao et al., 2019a). The chlorophyll a content was

174 determined with an DR 5000TM UV–Vis Spectrophotometer (Hach Company, Loveland, Co.,

175 USA).

176 The particle size distribution of the samples was measured with a Mastersizer 2000 laser

177 diffraction particle size analyzer (Malvern Panalytical Ltd, Malvern, UK), and the organic

178 matter content was determined with the dichromate redox titration method (Carter and

179 Gregorich, 2006). The soil pH was potentiometrically measured in 1/2.5 soil/water supernatant

180 with a PXSJ-216F pH meter (Shanghai INESA Instrument Co., Ltd., Shanghai, China) (Gentili

181 et al., 2018). The determine the saturated water content (θs) the samples were first saturated and

182 then oven-dried at 105 ℃. The field capacity (θf) was measured with a pressure plate apparatus

183 at −33 kPa soil water potential (Li et al., 2022b). The saturated hydraulic conductivity (Ks) was

184 determined with the constant head method (Reynolds and Elrick, 2002).

185 2.4. X-ray computed tomography analysis

186 2.4.1. X-ray computed tomography scanning and image processing

187 A Phoenix V|tome|x M® microfocus X-ray computed tomography (CT) scanner (Fig. 1F;
8
188 GE Sensing and Inspection Technologies GmbH, Pforzheim, Germany) was used to scan the

189 collected soil samples. Reconstruction of the X-ray CT images was performed with the General

190 Electric (GE) Datos|x CT software (version 2.0). A stack of 1000 image slices with a voxel

191 resolution of 50 μm was acquired with the CT scanner for each soil sample and stored in 8-bit

192 Tiff format for further processing.

193 After reconstruction, the image slices were imported into the Avizo™ software package

194 (Thermo Fisher Scientific Inc., Waltham, MA, USA) and a region of interest (ROI) with a size

195 of 20.0 × 20.0 × 50.0 mm (400 × 400 × 1000 voxels) was cropped for further analysis. The ROI

196 was selected from the central part of the sample to avoid edge effects. To reduce image noise,

197 a 3D median filter (2 × 2 × 2) was first applied, and then indicator kriging method was used for

198 image segmentation (Oh and Lindquist, 1999; Peth et al., 2008).

199 2.4.2. Pore parameters

200 After segmentation, the images were used to quantify macropore characteristics, including

201 macroporosity, surface area density, pore volume, node density, macropore pore size

202 distribution, shape factor, and morphological indicators (i.e., degree of anisotropy (DA), fractal

203 dimension (FD), Euler number (EN), and tortuosity). Moreover, the macropores were separated

204 into two classes, connected pores and isolated pores. Connected pores refer to all pores in the

205 system that are connected to at least one of the neighboring pores, whereas the isolated pores

206 have no connection to neighboring pores.

207 Macroporosity is defined as the macropore volume divided by the total volume of ROI. The

208 connected porosity is defined as the percentage of connected macropore volume to the total

209 volume of ROI. The isolated porosity was calculated as macroporosity minus connected

210 porosity. The surface area density was calculated as the total macropore surface area divided by

211 the total volume of ROI. The node density, which is the number of nodes where at least two

9
212 macropore branches connect per unit volume was used to quantify the interconnectivity of

213 macropores. A high node density represents an extensive and well-connected macropore

214 network. The macropores were separated into four size classes according to the equivalent

215 diameters of < 200 μm, 200−500 μm, 500−1000 μm, and > 1000 μm. The macropore shape

216 factor was calculated as (Zhou et al., 2012):

Ae
F (1)
A
217 where Ae is a sphere’s surface area with a volume equal to the measured pore volume (μm2),

218 and A is the measured pore surface area (μm2). The shape of a pore with F = 1 represents a

219 perfect sphere, whereas F values smaller than 1 represent pores with irregular or elongated

220 shapes. Based on the F value, the pores were classified as regular pores (F ≥ 0.5), irregular

221 pores (0.2 < F < 0.5), and elongated pores (F ≤ 0.2) (Pagliai et al., 2004).

222 The DA is an indicator of the degree of dissimilarity in the orientation of substructures

223 within a volume, and it varies between 0 (perfect isotropic structure) and 1 (anisotropic

224 structure). The FD quantifies self-similarity, and it increases with structural complexity (Dal

225 Ferro et al., 2013). The values for 3D image FD that are between 2 and 3 were estimated with

226 the box-counting method (Perret et al., 2003). The EN was utilized to estimate the degree of

227 connectivity. The smaller (more negative) the EN, the higher is the pore connectivity

228 (Wildenschild and Sheppard, 2013). The tortuosity was calculated as the ratio of the total actual

229 length of all macropores and the sum of the shortest distance between two ends of all

230 macropores (Katuwal et al., 2015).

231 2.5. Measurement of air permeability (Ka) and volumetric water content (θ)

232 For Ka measurements, a 11-cm diameter and 5-cm high PVC cylinders (Fig. 1G) were used

233 as in previous studies (Sun et al., 2022b; Sun et al., 2023). The PVC cylinders were carefully

234 pushed into the soil to ensure a tight fit. After allowing several days for soil stabilization, the

10
235 Ka of each plot was continuously measured every morning from 7:00 AM to 8:00 AM via the

236 steady-state method described in Iversen et al. (2001). An air compressor with a pressure

237 regulator and flow meter was employed to control the pressure applied to the top of the soil

238 enclosed by the cylinders to establish a pneumatic pressure difference of ~2.0 hPa. After steady

239 state air flow was established, the flow rate through the soil surface was recorded. Concurrently,

240 the near-surface volumetric soil water content (θ) was measured with TEROS 11 sensors

241 (METER Group, Inc., Pullman, WA, USA) to enable calculation of air-filled porosity (ɛa). The

242 Ka and θ measurements were repeated daily from August 1st to September 31st, 2022. The ɛa

243 (cm3 cm−3) and Ka (μm2) values were calculated according to Jalbert and Dane (2003):

b
a  1  (2)
s
Q
Ka  1012 (3)
DGP

 D  D D
G     ln(1  ) (1  ) (4)
4 H H H

  (1717  4.8Ta ) 108 (5)

244 where ρb is the soil bulk density (g cm–3), ρs is soil particle density (assumed to be 2.65 g cm−3),

245 θ is the volumetric soil water content (cm3 cm−3), μ is the dynamic viscosity of air (Pa s), Q is

246 the volumetric air flow rate (m3 s−1), D is the radius of the PVC cylinder (i.e., 0.055 m), G is a

247 geometric shape factor related to the dimensions of the inserted PVC cylinder, H is the height

248 of the cylinder (i.e., 0.05 m), ΔP is pneumatic pressure difference between the air inside the

249 cylinder (above the soil surface) and the free atmosphere (Pa), and T is the air temperature (℃).

250 2.6. Measurement of relative gas diffusivity (Dp/D0)

251 The soil samples collected in stainless steel rings for the Dp/D0 laboratory measurements

252 were first saturated with water for 48 h and then covered with a plastic film while allowing

253 excess water to drain. Then, the soil samples were gradually dried in an oven at 70 ℃ to obtain
11
254 a series of measurement points from saturation to dry. At each measurement point, the samples

255 were allowed to equilibrate at room temperature before weighing and subsequent measurement

256 of relative gas diffusivity with a transient method as described in Taylor (1949). A one-chamber

257 apparatus as discussed in Schjønning (1985) was used with O2 as the diffusing gas. Airtight

258 contact between the soil samples and the diffusion chamber was established and the O2

259 concentration within the chamber was measured with KE-25F3 Maxell oxygen sensor (Figaro

260 Engineering Inc., Osaka, Japan) and recorded in 5 second intervals with a CR1000 datalogger

261 (Campbell Scientific Inc., Logan, Utah, USA). Prior to the O2 measurements the chamber was

262 flushed with N2 until the O2 concentration in the chamber reached zero. In addition, the ambient

263 temperature and barometric pressure were measured throughout the experiment to account for

264 their effects gas diffusivity (Bakker and Hidding, 1970). Assuming steady state diffusion, the

265 O2 diffusion coefficient was determined from the relation between time and the logarithm of

266 the O2 concentration difference between the chamber and ambient air (Schjønning et al., 2013).

267 The standard gas diffusivity (Dp) at 20 ℃ and 101 kPa was calculated according to Bakker and

268 Hidding (1970):

 A   Ct 
Dp'    hs  hc  c  ln( ) t (6)
 As   C0 
1.75
 T  273   P0 
Dp  D  0
'
   (7)
 Ta  273 
p
 Pa 

269 where Dp' is the gas diffusivity at ambient temperature and barometric pressure (cm2 s−1), ΔCt

270 is the difference in O2 concentration between the chamber and ambient air, ΔC0 is the initial

271 difference in O2 concentration between the chamber and ambient air, hs is the height of the soil

272 sample (cm), hc is the height of the diffusion chamber (cm), As is the cross-sectional area of the

273 soil sample (cm2), Ac is the cross-sectional area of the diffusion chamber (cm2), t is time (s), T0

274 is the standard temperature (i.e., 20 ℃), Ta is the ambient temperature (℃), P0 is the standard

275 pressure (i.e., 101 kPa), and Pa is the ambient air pressure (kPa).
12
276 The relative gas diffusivity (Dp/D0) was calculated as the ratio of Dp and the gas diffusion

277 coefficient of O2 in free air (D0) at standard temperature and pressure. For D0 a value of 0.205

278 cm2 s−1 at 20 ℃ and 101 kPa was used (Currie, 1960; Hamamoto et al., 2009).

279 2.7. Measurement of water infiltration

280 Within each plot, five locations were selected to measure infiltration rates. A double-ring

281 infiltrometer, which consists of a 25-cm diameter and 30-cm high inner ring and a 45-cm

282 diameter and 30-cm high outer ring and two constant head devices (Fig. 1H) was used for the

283 measurements. The concentric rings were gently tapped into the soil to a depth of 15 cm with a

284 rubber hammer and the gaps between the rings and soil was sealed with wet soil to prevent

285 potential water leakage and preferential flow (Janssen and Lennartz, 2008). The inner and outer

286 rings were filled with water and a constant water level of 3 cm was maintained in both

287 throughout the measurement with the constant head devices. The decline in water height in the

288 constant head device connected to the inner ring was recorded in constant time intervals

289 throughout the measurements that lasted about 90 min. After the water decline per time interval

290 remained constant for 5 consecutive intervals steady state flow conditions were attained. The

291 average infiltration rate (ia) was calculated as the mean value of the entire 90 min. The initial

292 infiltration rate (i0) was calculated as the mean value of the first 3 min. Similarly, the steady-

293 state infiltration rate (is) was calculated using the last three values. The standard soil infiltration

294 rate at 10 ℃ (i) was calculated as (Suwardji and Eberbach, 1998):

r 2 h
i (8)
R 2 t (0.7  0.03T )

295 r is the radius of the constant head device (cm) (i.e., 7.0 cm), Δh is the decline in water height

296 per measurement interval of the constant head device connected to the inner ring (cm), R is the

297 radius of the inner ring (cm) (i.e., 12.5 cm), Δt is the constant measurement time interval (s),

298 and T is the water temperature (℃).

13
299 2.8. Data analysis

300 After normality and equality of variance tests, a one-way ANOVA with a least significant

301 different post hoc test at 5% probability level was performed to investigate the differences in

302 measured soil properties, water and gas transport parameters, and pore structure parameters

303 between biocrust-colonized and bare soils. The presented results represent the mean values of

304 all replicate measurements and are expressed as the mean ± standard error.

305 A structural equation model was established to explore causal relationships between surface

306 cover, soil properties, pore characteristics, aeration, and water infiltration. To evaluate our

307 hypothesis that biocrusts significantly alter soil pore structure and as a consequence soil

308 aeration and water infiltration, we determined the absolute fit of the best models via the

309 maximum likelihood χ2/df, goodness of fit index (GFI), root mean square error of

310 approximation (RMSEA), and P index. All path analyses were performed with version 26.0 of

311 the IBM SPSS AMOS software package (Amos Development Corporation, Chicago, IL, USA).

312

313 3. Results

314 3.1. Biocrust effects on near-surface soil properties

315 As shown in Table 1, all measured biocrust parameters were larger for aeolian sand than for

316 loess soil. The moss biomass, biocrust thickness, moss cover, moss density, and chlorophyll a

317 content averaged 0.22 g cm−2, 16.2 mm, 92.6%, 84.2 gametophyte cm−2, and 0.09 mg g−1 for

318 aeolian sand and were 29%, 26%, 6%, 4% and 125% higher than for loess soil, respectively.

319 When compared to the bare soils, the biocrust-colonized soils exhibited 52%–60% higher clay

320 contents, 36%–279% higher silt contents, but 12%–20% lower sand contents. In contrast to

321 bare soils, the biocrusts decreased bulk density and pH by 10%–19% and 4%–8%, respectively.

322 On average, the biocrusts also induced 70%–174% higher organic matter content and 139%–
14
323 154% higher surface roughness.

324 The biocrusts significantly modified hydraulic parameters. While the saturated hydraulic

325 conductivities (Ks) of biocrust-colonized soils were 39%–44% lower than for bare soils (Figs.

326 2A and 2B), the saturated water contents and field capacities were 16%–37% and 43%–173%

327 higher than for bare soils (Figs. 2C, 2D, 2E, and 2F).

328 3.2. Biocrust effects on pore characteristics

329 In general, the biocrusts significantly modified the characteristics of the soil pore system

330 (Fig. 3). In contrast to the bare soils, the biocrusts established on aeolian sand and loess soil

331 increased macroporosity by 102% and 107%, respectively. Correspondingly, the biocrusts also

332 increased connected porosity (i.e., 178% for aeolian sand and 107% for loess soil) and

333 connected/isolated porosity ratio (i.e., 787% for aeolian sand and 7% for loess soil). Specifically,

334 pores < 500 μm accounted for more than 62% of the macroporosity for all investigated scenarios

335 (Fig. 4). There were obvious differences in pore size distributions between biocrust-colonized

336 and bare soils. With regard to pores < 200 μm, the macroporosity of biocrust-colonized aeolian

337 sand was 32% higher than of bare aeolian sand (12.4% vs. 9.4%). Similarly, the macroporosity

338 of pores< 200 μm of biocrust-colonized loess soil was 12% higher than of bare loess soil (4.6%

339 vs. 4.1%). For the 200–1000 μm and > 1000 μm pore classes, the biocrusts increased

340 macroporosity by 156%–273% and 90%–8538% for biocrust-colonized aeolian sand and loess

341 soil relative to their bare counterparts.

342 Relative to the bare soils, the biocrust-colonized soils exhibited higher surface area density,

343 mean pore volume, and node density (Table 2). Specifically, the surface area density, mean pore

344 volume, and node density of aeolian sand colonized with biocrusts were increased by 56%,

345 203%, and 82%, respectively, when compared to bare aeolian sand. For the loess soil the

346 biocrusts increased the surface area density, mean pore volume, and node density by 87%, 72%,

15
347 and 178%, respectively, relative to bare loess soil.

348 As shown in Table 2, the biocrusts also had a significant effect on pore morphology

349 characteristics. For aeolian sand colonized with biocrusts, the degree of anisotropy (DA), Euler

350 number (EN), and mean tortuosity were 7%, 269%, and 7%, respectively, lower than for bare

351 aeolian sand. Compared to bare loess soil, the DA, EN, and mean tortuosity of biocrust-

352 colonized loess soil decreased by 12%, 2150%, and 2%, respectively. However, the fractal

353 dimension (FD) of biocrust-colonized aeolian sand and loess soil increased by 4% and 6% when

354 compared to their bare counterparts. The results above imply that pore structure is more

355 complex and stable because of the biocrusts. The elongated pores contributing the most to

356 macroporosity, and macroporosity of elongated pores of biocrust-colonized soils was 108%–

357 167% higher than the bare soils (Fig. 5).

358 3.3. Biocrust effects on soil aeration

359 The in-situ air permeability (Ka) measurements revealed that the Ka of biocrust-colonized

360 soils is significantly higher than Ka for the bare soils (Fig. 6). After precipitation, Ka markedly

361 decreased and then gradually increased until reaching a stable state (e.g., from August 28 to

362 September 18). The mean Ka of biocrust-colonized aeolian sand and loess soil was 50.24 and

363 24.00 μm2, which is 277% and 301% (F ≥ 93.70, P < 0.001) higher than for the corresponding

364 bare soils (13.33 and 5.98 μm2) (Figs. 7A and 7B). Furthermore, the biocrusts increased air-

365 filled porosity (ɛa) by 34% for aeolian sand (0.39 cm3 cm−3 vs. 0.29 cm3 cm−3) and by 59% for

366 loess soil (0.27 cm3 cm−3 vs. 0.43 cm3 cm−3) relative to the bare soils (Figs. 7C and 7D).

367 As shown in Fig. 8, the relative gas diffusivity (Dp/D0) was highest at low water contents

368 (θ) for all investigated scenarios. It initially rapidly decreased with increasing θ slowed down

369 until remaining constant. For the full θ range, the mean Dp/D0 of biocrust-colonized soils were

370 47%–59% higher than for the bare soils. Specifically, Dp/D0 for biocrust-colonized aeolian sand

16
371 increased by 53% (0.29 vs. 0.19) and by 64% (0.23 vs. 0.14) for biocrust-colonized loess soil

372 at minimum θ when compared to bare soils. However, it should be noted that Dp/D0 of the

373 biocrust-colonized soils were 17%–40% lower than for the bare soils at maximum θ. The Dp/D0

374 measurements indicate the existence of a θ threshold for biocrust-colonized soils close to

375 saturation above which soil aeration is impeded.

376 3.4. Biocrust effects on water infiltration

377 For both the loess soil and aeolian sand, biocrusts significantly reduced infiltration (i.e.,

378 infiltration rates and cumulative infiltration) (Fig. 9). As shown in Table 3, the biocrust-

379 colonized soils exhibited very different hydraulic parameters than the bare soils. The initial (i0),

380 steady state (is), and average (ia) infiltration rates of biocrust-colonized aeolian sand were 16%,

381 41%, and 28% lower than for bare aeolian sand, respectively. For biocrust-colonized loess soil,

382 i0, is, and ia were 50%, 50%, and 53% lower than for the bare loess soil, respectively. For both

383 the biocrust-colonized aeolian sand, the time to attain steady state flow conditions was 29%

384 longer than for the bare soils (Table 3). In addition, the cumulative infiltration amounts until

385 steady state flow for the biocrust-colonized soils were 12%–46% lower than for the bare soils.

386 The cumulative infiltration amounts after 60 min for the biocrust-colonized soils were 33%–

387 54% lower than for the bare soils (Fig. 9).

388 3.5. Effects of soil properties on soil pore characteristics, gas transport properties and

389 infiltrability

390 The structural equation model depicted in Fig. 10 shows that the moss-dominated biocrust

391 soil cover has a significant direct impact on clay particle content, organic matter content, pore

392 tortuosity, Euler number, soil gas transport properties, and soil infiltrability (path coefficient ≥

393 0.11, P < 0.05). The biocrust cover indirectly also impacted the pore morphological

394 characteristics (i.e., pore tortuosity and Euler number) via influencing the clay particle and

17
395 organic matter contents. The pore tortuosity and Euler number negatively impacted the soil gas

396 transport properties, which indicates that higher pore continuity facilitates elevated soil gas

397 exchange. The structural equation model also elucidated 94%–97% of the variance in the soil

398 infiltrability (Fig. 10). Specifically, the clay particle content had a significant direct negative

399 impact on water infiltrability due to the increase in soil water holding capacity. The results of

400 the structural equation model imply that the biocrusts modified the physicochemical soil

401 properties and altered soil pore structure and as a consequence affected soil aeration and

402 infiltrability.

403

404 4. Discussion

405 4.1. Biocrust effects on soil properties and soil macropore characteristics

406 Pore structure is affected by various soil properties, such as soil type and texture, clay

407 mineralogy, cation content, and soil organic matter content (Abiven et al., 2009). The pore

408 characteristics significantly deviated between moss-dominated biocrust-colonized soils and

409 their bare counterparts. Biocrust-colonized soils exhibited higher macroporosity, surface area

410 density, and pore volume than the bare soils, which was mostly due to the effects of biocrusts

411 on the soil physicochemical properties, such as bulk density, organic matter content, and a

412 higher number of clay particles (Table 1). Biocrusts trap and accumulate windblown fine

413 sediments and thereby modify the particle size distribution of the near-surface soil, which also

414 improves the arrangement and connection between soil particles. The enhanced surface

415 roughness of biocrust-colonized soils is effective in reducing sediment detachment and

416 transport by lowering the energy of falling raindrops. Furthermore, the addition of microbial

417 communities induced by biocrusts also promotes biological weathering of the mineral

418 component to generate fine particles (Chen et al., 2009). The enrichment of fine particles in

18
419 biocrust-colonized soils leads to a reduction in bulk density and higher soil porosity when

420 compared to bare soils.

421 In addition, the photoautotrophic component of biocrusts fixes atmospheric CO2 and

422 releases it into the surrounding soil. When the belowground biomass of biocrusts dies, organic

423 matter is accumulated, thereby enhancing soil aggregate formation and stability (Belnap et al.,

424 2016; Adessi et al., 2018). We found that biocrusts increased the clay particle fraction and

425 organic matter content. When compared to coarse soils, fine soils are generally denser and

426 exhibit higher porosities. Besides, as a critical cementing component, clay can increase

427 aggregate stability. Consequently, due to the larger fraction of fine particles and higher organic

428 matter contents of biocrust-colonized soils exhibit an increase of total pore space when

429 compared to bare soils.

430 Soil organic matter plays a pivotal role for soil aggregation and the stability of aggregates

431 (Zhou et al., 2012; Mustafa et al., 2020). A well-aggregated granular structure increases the

432 proportion of larger pores. In our study, biocrusts significantly increased mean pore volume and

433 surface area density, which is attributable to the higher organic matter content and the biotic

434 components of the biocrust layer. Soil organic matter can cohere mineral particles and facilitate

435 aggregate formation and the establishment of structural macropores when decomposed by

436 microbes into various metabolites (Peng et al., 2022). Biocrusts also directly contribute to the

437 formation of aggregates through their filamentous biological components (e.g., moss rhizoids,

438 lichen rhizines, and cyanobacterial filaments), microscopic fungal hyphae, and polysaccharide

439 and organic gels secreted by cyanobacteria, which is key to physically entwining clay particles

440 with each other and to enhance soil aggregation in the biocrust layer (Zhang et al., 2006; Weber

441 et al., 2022). Furthermore, the root-like structures of mosses are beneficial for generating voids

442 and large pores or enlarging smaller pores. Similar results are shown in Zhao et al. (2017), who

443 documented that root biomass was significantly positively correlated with macropores and

19
444 macroporosity.

445 Because of their effects on soil aggregation, biocrusts also modify morphological

446 parameters of the pore system. The connected/isolated pore (C/I) ratio, pore node density, and

447 the Euler number (EN) are indicators for the extent of pore connectivity. The results of the

448 presented study reveal that biocrusts increased the C/I ratio and node density but decreased the

449 Euler number relative to bare soil. This means that the pore system of biocrust-colonized soils

450 is more interconnected (Table 2). In general, macropores exhibited high continuity because the

451 biocrusts enhanced the establishment of macropores because of their positive effects on soil

452 aggregation as discussed above. When compared to bare soils, the enhanced macroporosity and

453 pore connectivity of biocrust-colonized soils is due to a higher number of large pore node

454 junctions and an increase of pore channels (Vogel, 1997). Biocrusts also increase the number

455 of biopores and enhance their connectivity because they promote the activity of microbial and

456 soil fauna communities (Souza-Egipsy et al., 2004; Liu et al., 2017). The lower tortuosity and

457 degree of anisotropy of the pore systems in biocrust-colonized soils suggest a more aligned and

458 isotropic pore system, which is because of an extensive and better connected macropore

459 network (Pires et al., 2019). These results are in agreement with our previous findings (Sun et

460 al., 2022b), which showed that biocrusts increase pore continuity but decrease tortuosity in the

461 upper soil layer.

462 All considered scenarios showed similar pore shape parameters. Elongated pores accounted

463 for the largest fraction of macroporosity. However, the biocrust-colonized soils increased the

464 fraction of elongated pores relative to the bare soils. In general, the large macropores are mostly

465 elongated, whereas regular pores are typically rounded and small (Wang et al., 2019). The

466 increased elongated porosity of biocrust-colonized soils is mainly attributable to the higher

467 organic matter content. Organic matter can cement more microaggregates to generate new large

468 pores and enhance biological activity in the soil to facilitate further development of pores.

20
469 Similar results are reported in Zhao et al. (2017) and Peng et al. (2022), who pointed out that

470 soil organic matter content is positively correlated with elongated pores. Furthermore, an

471 increase in the proportion of elongated pores in the biocrust-colonized soils also has contributed

472 to the increase in macroporosity, and complexity and connectivity of the pore network (Ma et

473 al., 2015; Zhao et al., 2017).

474 4.2. Biocrust effects on gas and water transport

475 The capability of soils to transport gas determines soil aeration and is dependent on various

476 soil properties, such as texture, bulk density, water content, the topology of the pore structure

477 (i.e., pore size and volume, connectivity, and tortuosity) (Sun et al., 2022b; Sun et al., 2023).

478 As a consequence, even minor changes in soil pore structure significantly impact the soil gas

479 transport characteristics. The air permeability (Ka) and relative gas diffusivity (Dp/D0) provide

480 insights into the advective and diffusive migration of gases due to air pressure and gas

481 concentration gradients. Gas transport within the soil and subsequent emission across the

482 soil/atmosphere interface predominantly occurs by means of diffusion without external forces

483 (Deepagoda et al., 2011). However, in the presence of external forces, the near-surface gas

484 movement caused by pressure changes due to temperature fluctuations, near-surface wind, and

485 water table fluctuations is governed by advection (Martínez et al., 2016).

486 The results of the presented study show that biocrust-colonized soils exhibited higher Ka

487 and Dp/D0 than the bare soils, which suggests that the biocrusts improve soil aeration in dryland

488 ecosystems (Figs. 7 and 8). In general, gas transport parameters are strongly affected by soil

489 bulk density, air-filled pore space, pore network connectivity, and tortuosity (Hamamoto et al.,

490 2009). The increase in Ka and Dp/D0 in biocrust-colonized soils is attributable to biocrust

491 induced changes of soil properties, such as a reduction of bulk density and tortuosity, and an

492 increase in air-filled porosity (ɛa) and pore connectivity. A reduction in bulk density caused by

21
493 biocrusts leads to an increase of total pore space within a distinct soil volume (Li et al., 2021b).

494 It is notable that Ka and Dp/D0 are affected differently by soil pore structure. Ka is more sensitive

495 to soil pore size than Dp/D0, because almost all gas transport by advection occurs across a few

496 connected macropores under pressure-induced conditions (Uteau et al., 2013). According to the

497 Hagen-Poiseuille law, the viscous flow through a simple pore is influenced by the fourth power

498 of the pore radius (Ball et al., 1981). Based on the X-ray CT analysis, biocrusts greatly increased

499 the percentage of soil pores > 1000 μm, connected porosity, and mean pore volume by 90%–

500 8538%, 107%–178%, and 72%–203% as well as increased Ka when compared to bare soils.

501 Similar results are also reported by Xiao et al. (2019a) and Sun et al. (2021), who documented

502 that biocrusts significantly increased the microscopic pore radius. However, gas diffusion is a

503 much slower process, because it is driven by concentration gradients (Arthur et al., 2012).

504 Unlike Ka, Dp/D0 is mainly associated with the connectivity and tortuosity of the soil pore

505 network, as well as air-filled pore space at a given matric potential (Masís-Meléndez et al.,

506 2014). Our data show that biocrust-colonized soils exhibit higher connectivity and lower

507 tortuosity values than bare soil (Table 2). Therefore, biocrusts are expected to facilitate Dp/D0.

508 Furthermore, the enhanced fraction of elongated pores in biocrusts also contributes to increased

509 Dp/D0, as elongated pores usually exhibit better continuity (Zhao et al., 2017). Apart from soil

510 properties, soil water content has a direct effect on air-filled pore space, thereby influencing

511 soil gas transport capability. Increasing the soil water content can block air pore space, decrease

512 airflow channels, and increase air pore tortuosity, consequently reducing soil gas transport

513 parameters (Mentges et al., 2016).

514 Our study shows that the Ks and infiltration parameters (e.g., is and cumulative infiltration

515 amount over 60 min) of biocrust-colonized soils are significantly lower than for bare soils,

516 indicating that the colonization with biocrusts impedes infiltration. In general, the hydraulic

517 conductivity is influenced by various soil properties, such as texture and structure, bulk density,

22
518 and organic matter content (Sun et al., 2022b). The impediment of water infiltration under

519 biocrust surfaces is modulated by the soil particle composition and swelling of extracellular

520 polymeric substances (EPS). Fine soils exhibit lower infiltrability than coarse sandy soils, as

521 sand grains do not bond. In the presented study, in comparison to bare soils, the fine particles

522 of biocrust-colonized soils 36%–279% higher (Table 1). The abundant fine particles in biocrusts

523 are tightly bound to and incorporated into the sticky sheaths of some cyanobacteria, thereby

524 generating a denser fine particle enriched soil surface, which is expected to decrease water

525 infiltration (Xiao et al., 2019a). Furthermore, biocrusts also increase the soil water-holding

526 capacity and saturated water content. The swelling of EPS in biocrusts is mainly caused by

527 bacteria, cyanobacteria, and microfungi (Raddadi et al., 2018; Kidron et al., 2022) and can lead

528 to pore clogging, which also reduces water infiltration. When wetted, EPS expands its volume

529 within seconds to up to 70 times of its dry volume (Chenu, 1993), which leads to an increase of

530 water-holding capacity and causes surface sealing and pore clogging. According to Kidron et

531 al. (2022), SEM pictographs show a myriad of loosely-bound and scattered EPS in biocrusts,

532 which inevitably leads to the reduction of available pore space near the soil surface. Moreover,

533 the sheaths of some cyanobacteria also increase their original volume during wetting and block

534 available pore space (Verrecchia et al.,1995; Fischer et al., 2010). The lower initial infiltration

535 rate of biocrusts is attributable to water repellency and lower water sorptivity of the initially

536 dry soil (Sun et al., 2022a). In addition, while the accumulation of organic matter in biocrust-

537 colonized soils improves the soil physical structure (i.e., pore connectivity and water

538 infiltration), pore blocking due to biotic biocrust components can counteract these positive

539 effects. The negative effects of biocrusts on soil water infiltration and positive effects on soil

540 water-holding capacity are consistent with previously obtained results for the same watershed

541 (Xiao et al., 2019a; Sun et al., 2021; Sun et al., 2022a). This implies that biocrusts play a

542 significant role in regulating hydrological processes and the surface soil water balance in

23
543 dryland ecosystems.

544 4.3. Implications of biocrust effects on soil pore structure, aeration, and water infiltration

545 Biocrusts are important engineers of the surface soil architecture and are capable of

546 reshaping various dryland soil properties, functions, and processes (Xiao et al., 2022). In

547 general, the soil structure regulates gas and vapor exchange processes, water storage and flow,

548 and it is also important for the ability of plant roots to penetrate the surface soil (Rabot et al.,

549 2018). The accurate determination of biocrust induced modifications to the soil structure

550 improves the understanding of the multifaceted biocrust effects on soil hydrological, ecological,

551 biological, and chemical processes. Based on the results of this study we conclude that moss-

552 dominated biocrusts significantly increased macroporosity, improved pore connectivity,

553 decreased pore tortuosity, increased soil aeration, and altered the pore shapes in the soils of the

554 Chinese Loess Plateau (Table 2). Higher pore continuity and connectivity of the macropores in

555 biocrust-colonized soils also promote preferential flow of water. This is consistent with findings

556 in Li et al. (2022b), who found based on dye tracer experiments that biocrust-colonized soils

557 exhibit a higher ratio of low and high contact flows than bare soils. Furthermore, soil gas and

558 vapor exchange is primarily governed by connected pores while isolated pores control carbon

559 sequestration (Yu et al., 2018). This is why the ratio of connected and isolated pores (C/I) is an

560 essential indicator for the functioning of the pore system. Because the soil structure provides a

561 habitat for various soil organisms, an increase in C/I ratio in biocrust-colonized soils facilitates

562 soil biological activity and carbon turnover (Yu et al., 2018) with important consequences for

563 dryland ecosystems. Furthermore, the biocrust induced improvement of soil aeration modifies

564 the soil oxygen and redox status, which promotes soil microbial abundance and community

565 diversity (Zhang et al., 2009). For example, Xiao and Veste (2017) found that moss-dominated

566 biocrust-colonized soils exhibited a large number and higher diversity of soil bacteria and fungi

567 than bare sand. It is well documented that soil microbial communities play significant roles in

24
568 nutrient cycling (Zhang et al., 2014), which implicates that rich and diversified microbial

569 communities in biocrust-colonized soils accelerate the rate of decomposition of soil organic

570 matter and sustain and improve soil fertility both of which are crucial for the ecological

571 restoration of degraded dryland ecosystems. In addition, the enhanced microbial activity in the

572 biocrust layer is important for soil respiration (Yao et al., 2020; Dou et al., 2023), which

573 indicates that biocrusts create stable microhabitats for soil microorganisms and thereby alter

574 soil structure and aeration. This is important for various ecological functions of microorganisms,

575 such as enzyme activity, nutrient turnover, and water cycling (Xiao and Veste, 2017).

576 The results of this study also show that biocrusts increase the water holding capacity while

577 they decrease water infiltration (Fig. 2 and Table 3). After medium- and high-intensity

578 precipitation events, the enhanced water retention and reduced water infiltration caused by

579 biocrusts leads to interception of water within the near-surface soil, which promotes the

580 generation of surface runoff and enhances water redistribution (Guan and Cao, 2019; Sun et al.,

581 2021). This is in agreement with Lichner et al. (2010), who reported an extremely high

582 infiltration rate of > 300 mm h−1 in bare sand of the Negev desert, whereas surface runoff of

583 biocrust-covered soils was triggered when the rainfall intensity approached 9–12 mm h−1. The

584 ratio of infiltration to surface runoff also affects the soil water balance. The effects of biocrusts

585 on soil water content are linked to multifaceted ecological processes. Some researchers argue

586 that a high soil water content induced by biocrusts supports vegetation establishment, enriches

587 species diversity, and promotes microbial activity, thereby causing a positive impact on the

588 ecosystem (Jiang et al., 2018; Eldridge et al., 2020). Other researchers argue that the high water

589 holding capacity of biocrusts provides a continuous supply of water close to the soil surface,

590 which enhances evaporation (Xiao and Bowker, 2020; Li and Xiao, 2022) and shortens the

591 pathways for vapor flow to the atmosphere (Li et al., 2022b). This is why biocrusts may cause

592 an overall negative effect on the soil water balance.

25
593

594 5. Conclusions

595 In this study, we analyzed the differences in soil pore structure, soil gas transport properties,

596 and infiltration processes between moss-dominated biocrust-colonized and bare soils on the

597 Chinese Loess Plateau. Our results revealed an obvious improvement in soil structure due to

598 biocrusts. Biocrusts significantly increased macroporosity, surface area density, and mean pore

599 volume by 102%–107%, 56%–87%, and 72%–203%, respectively, when compared to the bare

600 soils. The biocrust induced modification of morphological pore parameters indicates a pore

601 network with high connectivity and less tortuosity. Moreover, a large part of the macroporosity

602 of biocrust-colonized soils was comprised of < 500 μm elongated pores. The results of in-situ

603 and laboratory measurements also revealed that biocrusts increased Ka and Dp/D0 by 277%–

604 301% and 47%–59%, respectively, which is attributable to biocrust induced effects on surface

605 soil air-filled pore space, pore continuity and tortuosity. However, biocrust-colonized soils

606 exhibited a significantly lower soil infiltrability but higher water-holding capacity. The Ks and

607 steady state infiltration rate of biocrust-colonized soils were 39%–44% and 41%–50% lower

608 than for the bare soils. Nevertheless, biocrusts increased the θs and θf by 16%–37% and 43%–

609 173%. The results of this study imply that biocrusts play a multifaceted and complex role in the

610 transport of gases and water, and as a consequence significantly impact near-surface

611 hydrological and biochemical processes, which is vital for sustaining and improving dryland

612 ecosystem functions and services.

613

614 CRediT authorship contribution statement

615 Fuhai Sun: Conceptualization, Methodology, Formal analysis, Investigation, Resources,

616 Writing - original draft. Bo Xiao: Formal analysis, Writing - original draft, Writing - review &

26
617 editing, Supervision, Project administration, Funding acquisition. Markus Tuller: Writing -

618 review & editing.

619

620 Acknowledgments

621 This study was funded by the National Natural Science Foundation of China (grant no.

622 42077010), the “Light of West China” Program of the Chinese Academy of Sciences (grant no.

623 2019), and the Open Fund for Key Laboratory of Land Degradation and Ecological Restoration

624 in Northwestern China of Ningxia University (grant no. LDER2022Z02). The authors are

625 grateful to the Shenmu Experimental Station of Soil Erosion and Environment, CAS & MOE

626 for logistical support. Furthermore, the authors express their gratitude to Yanjia Zheng for

627 assistance with data collection.

628

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923 Zhao, D., Xu, M.X., Liu, G.B., Ma, L.Y., Zhang, S.M., Xiao, T.Q., Peng, G.Y., 2017. Effect of

924 vegetation type on microstructure of soil aggregates on the Loess Plateau, China. Agr.

925 Ecosyst. Environ. 242, 1–8. https://doi.org/10.1016/j.agee.2017.03.014.

926 Zhou, H., Mooney, S.J., Peng, X.H., 2017. Bimodal soil pore structure investigated by a

927 combined soil water retention curve and X-Ray computed tomography approach. Soil Sci.

928 Soc. Am. J. 81, 1270. https://doi.org/10.2136/sssaj2016.10.0338.

929 Zhou, H., Peng, X.H., Peth, S., Xiao, T.Q., 2012. Effects of vegetation restoration on soil

930 aggregate microstructure quantified with synchrotron-based micro-computed tomography.

931 Soil Till. Res. 124, 17–23. https://doi.org/10.1016/j.still.2012.04.006.

41
932 Table 1

933 Properties of biocrust-colonized and bare soils.

Aeolian sand Loess soil

Measurements
Bare soil Biocrusts F P Bare soil Biocrusts F P

Moss biomass (g cm−2) – 0.22 ± 0.02 – – – 0.17 ± 0.02 – –

Biocrust thickness (mm) – 16.21 ± 0.89 – – – 12.87 ± 0.53 – –

Moss cover (%) – 92.59 ± 3.57 – – – 87.44 ± 2.36 – –

Moss density (gametophyte cm−2) – 184.23 ± 15.85 – – – 177.81 ± 18.89 – –

Chlorophyll a content (mg g−1) – 0.09 ± 0.02 – – – 0.04 ± 0.01 – –

Sand content (20‒2000 μm) (%) 94.78 ± 0.61 83.79 ± 0.80 361.86 < 0.001 66.63 ± 1.49 53.33 ± 0.21 234.20 < 0.001

Silt content (2‒20 μm) (%) 3.58 ± 0.42 13.58 ± 0.79 378.12 < 0.001 26.08 ± 1.05 35.57 ± 0.15 239.40 < 0.001

Clay content (< 2 μm) (%) 1.64 ± 0.06 2.63 ± 0.02 732.87 < 0.001 7.29 ± 0.44 11.10 ± 0.06 220.95 < 0.001

Bulk density (g cm−3) 1.56 ± 0.01 1.27 ± 0.02 525.18 < 0.001 1.35 ± 0.02 1.21 ± 0.07 11.80 0.026

Soil pH 8.20 ± 0.01 7.53 ± 0.12 92.48 0.001 8.62 ± 0.09 8.25 ± 0.03 46.20 0.002

Surface roughness 2.22 ± 0.15 5.64 ± 0.26 400.00 < 0.001 3.50 ± 0.15 8.38 ± 0.53 232.07 < 0.001

Organic matter content (g kg−1) 7.26 ± 2.06 19.87 ± 0.78 98.24 0.001 15.61 ± 3.94 26.47 ± 1.14 21.06 0.010
934

42
935 Table 2

936 Pore characteristics of biocrust-colonized and bare soils.

Aeolian sand Loess soil

Parameters
Bare soil Biocrusts F P Bare soil Biocrusts F P

Macroporosity (%) 12.25 ± 0.23 24.75 ± 0.23 1712.90 < 0.001 15.67 ± 1.45 32.42 ± 1.02 267.24 < 0.001

Connected porosity (%) 8.48 ± 0.85 23.59 ± 0.55 670.43 < 0.001 15.25 ± 1.52 31.64 ± 1.05 235.95 < 0.001

Isolated porosity (%) 3.78 ± 0.62 1.16 ± 0.10 51.55 0.002 0.42 ± 0.10 0.78 ± 0.09 22.11 0.009

C/I ratio 2.31 ± 0.60 20.49 ± 2.22 186.96 < 0.001 38.20 ± 13.70 40.73 ± 5.31 0.09 0.781

Surface area density (mm2 mm−3) 2.78 ± 0.09 4.35 ± 0.33 61.99 0.001 1.44 ± 0.17 2.69 ± 0.69 9.19 0.039

Mean pore volume (×10−2) (mm3) 2.04 ± 0.07 6.18 ± 0.22 932.78 < 0.001 11.03 ± 3.23 18.92 ± 4.17 8.25 0.045

Node density (number mm−3) 7.16 ± 0.58 13.00 ± 1.61 34.80 0.004 1.99 ± 0.18 5.53 ± 2.14 8.09 0.047

Degree of anisotropy 0.60 ± 0.02 0.56 ± 0.02 12.14 0.025 0.59 ± 0.04 0.52 ± 0.01 8.13 0.046

Fractal dimension 2.57 ± 0.00 2.67 ± 0.01 316.07 < 0.001 2.48 ± 0.06 2.62 ± 0.05 9.45 0.037

Euler number (×105) 0.70 ± 0.10 −1.18 ± 0.35 80.34 0.001 −0.02 ± 0.05 −0.45 ± 0.27 15.28 0.017

Mean tortuosity 1.22 ± 0.02 1.13 ± 0.00 74.55 0.001 1.13 ± 0.00 1.11 ± 0.00 33.73 0.004

937

43
938 Table 3

939 Infiltration parameters of biocrust-colonized and bare soils.

Aeolian sand Loess soil

Measurements
Bare soil Biocrusts F P Bare soil Biocrusts F P

i0 (cm min−1) † 0.68 ± 0.05 0.57 ± 0.04 17.24 0.003 0.36 ± 0.04 0.18 ± 0.04 49.97 < 0.001

is (cm min−1) 0.46 ± 0.02 0.27 ± 0.02 217.41 < 0.001 0.08 ± 0.01 0.04 ± 0.01 80.78 < 0.001

ia (cm min−1) 0.54 ± 0.02 0.39 ± 0.03 98.18 < 0.001 0.19 ± 0.02 0.09 ± 0.02 57.23 < 0.001

ts (min) 34.76 ± 1.42 44.97 ± 2.00 87.01 < 0.001 41.03 ± 2.76 52.86 ± 3.05 41.35 < 0.001

Ic (cm) 18.90 ± 0.63 16.58 ± 1.52 9.98 0.013 7.16 ± 0.88 3.84 ± 0.98 31.91 < 0.001

I60 (cm) 30.83 ± 0.82 20.71 ± 1.64 151.89 < 0.001 9.08 ± 1.06 4.14 ± 1.05 55.32 < 0.001
940 † i0 is the initial infiltration rate, which was obtained from the average infiltration rate of the first 3 minutes; is is the steady state infiltration rate, which was obtained
941 from the average infiltration rate of the last 10 minutes; ia is the average infiltration rate, which was obtained from the of the average infiltration rate within the 90
942 minutes; ts is time to reach steady-state infiltration; Ic is the cumulative infiltration at the time reached steady-state infiltration; I60 is the cumulative infiltration within
943 60 minutes.

44
944 Figure Captions

945 Fig. 1. (A) Location of the Liudaogou watershed, (B) bare aeolian sand, (C) biocrust-colonized

946 aeolian sand, (D) bare loess soil, (E) biocrust-colonized loess soil, (F) Phoenix V|tome|x M®

947 microfocus X-ray computed tomography (CT) scanner, (G) PVC cylinder used for air

948 permeability measurements, and (H) double-ring infiltrometer experiment.

949 Fig. 2. Hydraulic properties of biocrust-colonized and bare soils. (A) Saturated hydraulic

950 conductivity (Ks) of aeolian sand, (B) Ks of loess soil, (C) saturated water content (θs) of aeolian

951 sand, (D) θs of loess soil, (E) field capacity (θf) of aeolian sand, and (F) θf of loess soil.

952 Fig. 3. 2D and 3D macropore networks of biocrust-colonized and bare soils. (A) Bare aeolian

953 sand, (B) biocrust-colonized aeolian sand, (C) bare loess soil, and (D) biocrust-colonized loess

954 soil.

955 Fig. 4. Pore size distributions of biocrust-colonized and bare soils for aeolian sand (A) and loess

956 soil (B). * and ** indicate a significant difference between bare soil and biocrusts at the 0.05 or

957 0.01 probability levels, respectively.

958 Fig. 5. Pore shapes of biocrust-colonized and bare soils for aeolian sand (A) and loess soil (B).

959 Fig. 6. (A) Daily precipitation and soil moisture (θ) data displayed together with in-situ air

960 permeability measurements for (B) bare aeolian sand, (C) biocrust-colonized aeolian sand, (D)

961 bare loess soil, and (E) biocrust-colonized loess soil.

962 Fig. 7. Mean air permeability (Ka) and air-filled porosity (εa) of biocrust-colonized and bare

963 soils measured in-situ.

964 Fig. 8. Relationship between relative gas diffusivity (Dp/D0) and soil water content (θ) for

965 biocrust-colonized and bare soils for aeolian sand (A) and loess soil (B).

966 Fig. 9. Infiltration rates and cumulative infiltration plotted as a function of time for biocrust-

45
967 colonized and bare soils for aeolian sand and loess soil.

968 Fig. 10. Structural equation model revealing the relationships between soil cover, clay particle

969 content, organic matter content, pore tortuosity, Euler number, soil aeration, soil water holding

970 capacity, and soil infiltrability of biocrust-colonized aeolian sand (A) and loess soil (B). The

971 value next to each arrow represents the standardized path coefficient. The arrows indicate a

972 direct effect of a single one-way causal relationship. The arrow width indicates the strength of

973 causality, which is in accordance with the standardized path coefficients and shown adjacent to

974 the arrows. The solid lines indicate positive path coefficients, and the dashed lines indicate

975 negative path coefficients. The oval represents latent variable and rectangle represents manifest

976 variable. * and **

means the coefficient is significant at the 0.05 or 0.01 level of probability.

977 Goodness-of-fit statistics are shown underneath the modeling frames.

46
Figure 1- Upload high quality Images Click here to access/download;Figure - Upload high quality Images;Figure 1.docx

Fig. 1. (A) Location of the Liudaogou watershed, (B) bare aeolian sand, (C) biocrust-colonized aeolian sand, (D) bare loess soil, (E) biocrust-

colonized loess soil, (F) Phoenix V|tome|x M® microfocus X-ray computed tomography (CT) scanner, (G) PVC cylinder used for air permeability

measurements, and (H) double-ring infiltrometer experiment.

Figure 2- Upload high quality Images Click here to access/download;Figure - Upload high quality
Images;Figure 2.docx

Fig. 2. Hydraulic properties of biocrust-colonized and bare soils. (A) Saturated hydraulic

conductivity (Ks) of aeolian sand, (B) Ks of loess soil, (C) saturated water content (θs) of aeolian

sand, (D) θs of loess soil, (E) field capacity (θf) of aeolian sand, and (F) θf of loess soil.
Figure 3- Upload high quality Images Click here to access/download;Figure - Upload high quality
Images;Figure 3.docx

Fig. 3. 2D and 3D macropore networks of biocrust-colonized and bare soils. (A) Bare

aeolian sand, (B) biocrust-colonized aeolian sand, (C) bare loess soil, and (D) biocrust-

colonized loess soil.

Figure 4- Upload high quality Images Click here to access/download;Figure - Upload high quality Images;Figure 4.docx

100 100
A Bare soil B *
* ** Biocrusts
*
10 10
**
**
Macroporosity (%)

Macroporosity (%)
1 1

0.1 0.1

0.01 0.01

0.001 0.001
<200 200-500 500-1000 >1000 <200 200-500 500-1000 >1000
Pore size (μm) Pore size (μm)

Fig. 4. Pore size distributions of biocrust-colonized and bare soils for aeolian sand (A) and loess soil (B). * and ** indicate a significant difference

between bare soil and biocrusts at the 0.05 or 0.01 probability levels, respectively.
Figure 5- Upload high quality Images Click here to access/download;Figure - Upload high quality Images;Figure 5.docx

100 100
A Bare soil F = 380.55, P = 0.002 B F = 234.68, P < 0.001
Biocrusts
a
a
b
b
Macroporosity (%)

Macroporosity (%)
10 10
F = 85.62, P = 0.001

F = 45.91, P = 0.002
a
F = 24.40, P = 0.008
a
b
1 1
a

b
a a
0.1 0.1
Regular pores Irregular pores Elongated pores Regular pores Irregular pores Elongated pores

Fig. 5. Pore shapes of biocrust-colonized and bare soils for aeolian sand (A) and loess soil (B).
Figure 6- Upload high quality Images Click here to access/download;Figure - Upload high quality
Images;Figure 6.docx

Fig. 6. (A) Daily precipitation and soil moisture (θ) data displayed together with in-situ
air permeability measurements for (B) bare aeolian sand, (C) biocrust-colonized aeolian
sand, (D) bare loess soil, and (E) biocrust-colonized loess soil.
Figure 7- Upload high quality Images Click here to access/download;Figure - Upload high quality
Images;Figure 7.docx

Fig. 7. Mean air permeability (Ka) and air-filled porosity (εa) of biocrust-colonized and bare
soils measured in-situ.
Figure 8- Upload high quality Images Click here to access/download;Figure - Upload high quality Images;Figure 8.docx

Fig. 8. Relationship between relative gas diffusivity (Dp/D0) and soil water content (θ) for biocrust-colonized and bare soils for aeolian sand (A)

and loess soil (B).

Figure 9- Upload high quality Images Click here to access/download;Figure - Upload high quality Images;Figure 9.docx

0.9 0.9
A Aeolian sand Bare soil B Loess soil
Biocrusts

Infiltration rate (cm min−1)

Infiltration rate (cm min−1)

0.6 0.6

0.3 0.3

0.0 0.0
0 20 40 60 80 100 0 20 40 60 80 100
Time (min) Time (min)
50 15
C Aeolian sand D Loess soil

40 12

Cumulative infiltation (cm)

Cumulative infiltation (cm)

30 9

20 6

10 3

0 0
0 20 40 60 80 100 0 20 40 60 80 100
Time (min) Time (min)

Fig. 9. Infiltration rates and cumulative infiltration plotted as a function of time for biocrust-colonized and bare soils for aeolian sand and loess

soil.
Figure 10- Upload high quality Images Click here to access/download;Figure - Upload high quality Images;Figure 10.docx

Fig. 10. Structural equation model revealing the relationships between soil cover, clay particle content, organic matter content, pore tortuosity,
Euler number, soil aeration, soil water holding capacity, and soil infiltrability of biocrust-colonized aeolian sand (A) and loess soil (B). The value
next to each arrow represents the standardized path coefficient. The arrows indicate a direct effect of a single one-way causal relationship. The
arrow width indicates the strength of causality, which is in accordance with the standardized path coefficients and shown adjacent to the arrows.
The solid lines indicate positive path coefficients, and the dashed lines indicate negative path coefficients. The oval represents latent variable and
rectangle represents manifest variable. * and ** means the coefficient is significant at the 0.05 or 0.01 level of probability. Goodness-of-fit statistics
are shown underneath the modeling frames.