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As an important soil cover in deserts, biological crusts play a central role in ecosystem function such as nutrient cycling, nitrogen fixation, and carbon sequestration. Many biological crust organisms could fix CO 2 through photosynthesis to improve soil organic carbon content. There is a knowledge gap in the origin of soil organic carbon (SOC) from biological crusts on a regional level, which restricts the prediction of soil carbon pool. Based on 45 plots in the Mu Us sandy land (42200 km 2 ), we measured the SOC content and soil organic carbon density (SOCD) of two types of typical biological crusts (moss crusts, algal crusts) and their underlying soils, and analyzed together with the climate data, soil and vegetation factors to investigate the spatial distribution characteristics and controlling factors of organic carbon of biological crusts at the regional scale. The results showed that: 1) biological crusts significantly increased SOC and SOCD compared with bare ground. Moss crusts and the underlying SOC (4.93 g·kg -1 ) and SOCD (0.41 kg·m -2 ) were higher than those of algal crusts (1.89 g·kg -1 , 0.18 kg·m -2 ). 2) On the regional scale, the SOC and SOCD of biological crusts had clear spatial distribution characteristics, demonstrating a banded distribution and block mosaic from northeast to central and west to southeast. 3) The SOC and SOCD of biological crusts and their underlying soils were mainly affected by climate, soil and vegetation conditions, while the main controlling factors depended on the types of biological crusts. The SOC and SOCD of moss crust were controlled by annual maximum temperature and potential evapotranspiration, whereas those of algal crusts were controlled by water vapor pressure.

We selected typical croplands in the black soil region of Northeast China to analyze the characteristics of biocrusts during the growing season, including species composition, thickness, coverage, and biomass (chlorophyll content). We collected bareground soil and biocrusts samples with chlorophyll content of 5-15, 15-25, 25-35, and 35-50 mg·g -1 , and measured the soil disintegration rate and soil maximum disintegration ratio of each sample using a force gauge in the laboratory. The results showed that: 1) biocrusts dominated by algae and moss were frequently developed in the croplands, with Stigeoclonium and Bryum capillare as the most common species, respectively. The thickness and biomass of algal crusts were significantly lower than moss crusts, with a successional trend from algal crusts to moss crusts. 2) The coverage, thickness, and biomass of biocrusts in croplands were negatively correlated with the frequency and intensity of tillage disturbance. For instance, the values of those characterisitics were only 27.8%, 1.52 mm, and 6.49 mg·g -1 on average, respectively, in traditional tillage croplands, and increased to 83.5%, 2.74 mm, and 34.16 mg·g -1 , respectively, in the croplands with conservational tillage. 3) Biocrusts considerably reduced the disintegration of surface soil, particularly in the layer of biocrusts. Compared to the bareground soil, the soil disintegration rate of biocrusts, with four levels of biomass (with chlorophyll content of 5-15, 15-25, 25-35, and 35-50 mg·g -1 ), was reduced by 43.1%, 50.1%, 55.5%, and 59.8%, respectively, while the soil maximum disintegration ratios were reduced by 11.4%, 17.7%, 33.2%, and 36.6%, respectively. 4) Soil disintegration rate and maximum disintegration ratio were significantly and negatively correlated with the biomass and thickness of biocrusts, indicating that the impacts of biocrusts on soil disintegration were primarily caused by the improvements in physical properties of surface soil. In conclusion, biocrusts were frequently deve-loped in croplands in the black soil region of Northeast China, owing to less disturbance following the conversion from traditional tillage to conservational tillage. They had the potential to protect surface soil against disintegration and improve soil anti-scourability, which was critical for soil conservation in croplands in this region.

We examined the effects of moss-dominated biocrusts on soil infiltration properties in Wangjiaqiao watershed of the Three Gorges Reservoir Area. Five levels of coverage (1%-20%, 20%-40%, 40%-60%, 60%-80% and 80%-100%) were set, with a nearby bare land as the control. We collected soil samples and conducted infiltration process observation by double cutting ring method. The results showed that biocrusts could appreciably increase soil cohesion, porosity, clay content, water-stable aggregates and organic carbon of topsoil, but significantly reduce soil bulk density and sand content. Biocrusts promoted soil water infiltration, with the initial infiltration rate ( I i ), stable infiltration rate ( I f ), average infiltration rate and cumulative infiltration amount being two times or more of that in bare land. Withbiocrust coverage increasing, soil infiltration properties firstly increased and then decreased, and peaked at 40%-60% coverage level. Results of path analysis indicated that I i was mainly affected by biocrust coverage, soil bulk density, and organic carbon content, while I f was mainly affected by biocrust coverage and soil bulk density. The simulation results of four infiltration models demonstrated that Horton model was the best fitting on the water infiltration process of biocrusted soil in the Three Gorges Reservoir Area.

Biological soil crusts (biocrusts) are the mixed community composed of different ratios of cyanobacteria, mosses, and lichens at the slope scale in the Hilly Loess Plateau region. Biocrusts significantly affect water infiltration in this area. The relationship between infiltration rate and community structure of mixed biocrusts is unknown, which would hinder the assessment of soil permeability of biocrusts at the slope scale. We measured the stable infiltration rate of cyanobacteria, moss, and mixed biocrusts with different proportions of cyanobacteria and moss including moss coverage of <15%, 15%-30%, 30%-45%, 45%-60% and >60%, respectively. The principal component analysis and path analysis were used to understand the influencing factors of stable infiltration rate of mixed biocrusts, and to clarify the relationship between the stable infiltration rate and the community structure of mixed biocrusts. The results showed that the saturated hydraulic conductivity of cyanobacteria and moss crusts was 0.66 mm·min -1 and 2.40 mm·min -1 , respectively. The stable infiltration rates of mixed biocrusts with moss coverage <15% to >60% were 0.80-2.30 mm·min -1 . The stable infiltration rate of mixed biocrusts at the slope scale depended on moss coverage and its improvement on soil pore structure, with the correlation coefficients being 0.636 ( P =0.011) and 0.835 ( P =0.000) respectively. Herein, the saturated hydraulic conductivity and coverage of cyanobacteria and moss confirmed the weighted prediction of water infiltration volume ( y ) i.e ., a significant correlation ( r =0.945) with the measured water infiltration volume ( x ) of mixed biocrusts. The linear fitting of measured and predicted water infiltration volume of mixed biocrusts was y =0.85 x ( R 2 =0.98, P <0.05). This study clarified the relationship between water infiltration of mixed biocrust community composition and individual biocrust composition, which provided a scientific basis for accurately evaluating the hydrological process of biocrusts in this area.

Bryophytes have been used in many fields, such as landscaping and soil and water conservation. How-ever, few studies focused on moss regeneration and its influencing factors, which retards the application study. Three common desiccation-tolerant mosses ( Barbula unguiculata , Didymodon vinealis , and Didymodon tectorum ) in the Loess Plateau region were collected across four seasons. We measured vigor index of new shoots, representing vegetative regeneration capacity, and physiological indices, which included the contents of chlorophyll, soluble sugar, soluble protein and malondialdehyde (MDA), to determine seasonal differences in regeneration capacity and physiological characteristics of the three mosses, as well as their relationships. The results showed that, 1) vegetative regeneration of mosses showed significantly seasonal differences. The vigor index of the three mosses in summer were the lowest, averagely decreased by 56.1%, 48.4%, and 10.1% compared with that in autumn, winter, and spring, respectively. The vigor index of the three mosses in the same season showed interannual variation. 2) There were considerable differences in the regeneration capacity of mosses across species. D. tectorum and B. unguiculata had the highest and lowest vigor indices, respectively. In terms of regeneration capacity, B. unguiculata had the largest seasonal and interannual variations. 3) The physiological characteristics of mosses had seaonal variations. The mosses collected in the summer had the highest MDA content but the lowest contents of soluble sugar and soluble protein. 4) Seasonal variations in desiccation-tolerant moss regeneration were mostly due to soluble sugar. Our results showed seasonal variations of vegetative propagation capacity, and highlighted the role of soluble sugar as a critical factor influencing vegetative propagation of mosses. These findings could provide scientific support for moss protection and artificial cultivation.

Microhabitat factors play an important role in regulating bryophyte species distribution and the development of bryophyte-dominated biological soil crusts (hereafter bryophyte crusts). We investigated the distribution and development of bryophytes in eight microhabitats in the water-wind erosion crisscross region of the Loess Pla-teau. We used the line intercept transects to explore and quantify the influencing pathways of microhabitat factors on bryophyte diversity and analyzed the influencing pathways of plant cover, slope aspect, and slope gradient by using structural equation model to quantify influencing coefficients. Our results showed that: 1) The Patrick, Shannon, Pielou, and Simpson indcies of bryophytes under plant canopy were 63.4%, 66.6%, 91.0%, and 68.3% lower than that without plant canopy, respectively, while the thickness, biomass, and chlorophyll content of bryophyte crusts were 0.5, 0.2, and 1.3 times higher than that without plant canopy, respectively. 2) The Patrick, Shannon, Pielou, and Simpson indexes of bryophytes on the north slope were 0.6, 0.9, 5.6, and 0.9 times higher than those on the south slope, while the thickness, biomass, and chlorophyll content of bryophyte crusts were 0.3, 0.3, and 0.6 times higher than those on the south slope, respectively. 3) As the slope increasing from 14° to 34°, the Patrick, Shannon, Pielou, and Simpson indexes of bryophyte were decreased by 59.8%, 84.1%, 57.3% and 68.0%, and the thickness, biomass, and chlorophyll content of bryophyte crusts were decreased by 15.2%, 25.0%, and 16.5%, respectively. 4) The importance of the three microhabitat factors on bryophyte diversity and the development of bryophyte crusts followed an order of plant canopy cover > slope aspect > slope gradient. The primary influencing pathway varied among the microhabitat factors. In conclusion, plant cover, slope aspect, and slope gradient significantly affected the distribution of bryophytes species and developmental level of bryophyte crusts through direct and indirect pathways. Therefore, full consideration should be given to microhabitat conditions when using bryophyte crusts to control desertification.

Biological crusts (Biocrusts) are important surface active coverings in arid and semi-arid regions, which affect the content of soil organic carbon (SOC), SOC labile fractions and stability of SOC through photosynthetic carbon fixation. At present, studies on the variation characteristics of SOC, SOC labile fractions and the stability of SOC in biocrusts are rather limited. In this study, two types of typical biocrusts (moss crusts and algae crusts) were selected along a precipitation gradient from northwest to southeast in the Mu Us Sandland (straight line distance 188 km) by measuring soil organic carbon (SOC), soil microbial biomass carbon (MBC), water soluble carbon (DOC), particulate carbon (POC), easily oxidizable carbon (ROC). We aimed to explore the effects of biocrusts on the stability of SOC and carbon decomposition across the precipitation gradient. Results showed that:1) Two types of biocrusts significantly increased the contents of SOC, MBC, DOC, POC, ROC and stability of SOC. Moss crusts increased SOC contents by 1.6 to 2.6 times as that of algae crusts. 2) The lowest SOC contents of the two types of biocrusts were 6.43 g·kg -1 and 14.50 g·kg -1 respectively, which showed an increasing trend with increasing precipitation along the gradient. 3) With the increases of precipitation, the decomposition time of moss litters gradually decreased. The decomposition coefficient of moss litters during the study period (From July to Feb-ruary of the next year) ranged from 0.010 to 0.014, which was significantly lower than that of vascular plants. The carbon release of moss litters from northwest to southeast was 8.09, 10.89, 12.88 g·kg -1 , respectively. 4) Results of canonical correspondence analysis showed that water vapor partial pressure, actual evapotranspiration, annual average temperature, subsurface short-wave radiation, potential evapotranspiration and vapor pressure difference were the key climate factors affecting the content of SOC and its active components. Silt content was the main soil factor affecting the content of SOC and its active components.

The formation and development of biological soil crusts (biocrusts) potentially affect the cycles and stoichiometric characteristics of soil carbon (C), nitrogen (N), and phosphorus (P). However, it is still unclear how soil microbes adapt to such changes. In this study, we examined the effects of moss-dominated biocrusts coverage (0, 1%-20%, 20%-40%, 40%-60%, 60%-80%, and 80%-100%) on soil physicochemical properties, soil microbial biomass, and ectoenzyme activities [β-1, 4-glucosidase (BG), β-1, 4-N-acetyl glucosidase (NAG), acid phosphatase (AP)] in two soil layers (0-5 and 5-10 cm) in the Three Gorges Reservoir area, as well as the covariations of soil-microbe-ectoenzyme C:N:P stoichiometry. The results showed that biocrust development significantly increased soil clay content, water stable aggregates, soil C, N, P contents, and significantly decreased soil bulk density and sand content. Microbial biomass C, N, P and ectoenzyme activities were significantly increased with increasing biocrust coverage. Soil depth did not affect soil physicochemical properties and C:N:P, but significantly affected microbial biomass, ectoenzyme activities, BG:AP and NAG:AP. Soil C, N and P contents were significantly positively correlated with microbial biomass and ectoenzyme activities, negatively correlated with BG:NAG, while positively correlated with NAG:AP, but had no significant correlation with microbial biomass C:N:P. There was no significant correlation between soil-microbe and microbial-ectoenzyme C:N:P. BG:NAG:AP decreased gradually with the increase of C:N:P stoichiometric imbalance between microbe and soil. This study indicated that the microbial metabolism was co-limited by N and P and with stronger P limitation. Microbes could maintain homeostasis by adjusting their own biomass and ectoenzyme C:N:P to adapt to changes in soil ecological stoichiometry driven by biocrust development.

Biological soil crusts (biocrusts) are the common cover in arid and semiarid areas. Together with plants, biocrusts affect runoff and flow velocity. However, few studies have focused on the effects of the co-covering of plant and biocrust (plant+biocrust) on the flow velocity, with a knowledge gap in the study of driving factors for slope erosion in arid and semiarid areas. In this study, simulated rainfall experiments were used to investigate the effects of biocrust and three types of biocrusts (more cyanobacteria less moss, more moss less cyanobacteria, and moss) on the flow velocity of revegetated grassland in the hilly Loess Plateau. The results showed that plant and plant+biocrust significantly reduced flow velocity, with that of plants and plant+biocrust being 70.7% and 83.1% lower than bare soil. The reduction benefits of plant and biocrust on flow velocity were 70.7% and 12.4%, respectively, when they were co-covered. Biocrust composition under plant cover affected flow velocity. The reduction benefits of more cyanobacteria less moss, more moss less cyanobacteria, and moss crust on flow velocity were 11.5%, 12.4%, and 19.4%, respectively. There was a significant negative correlation between flow velocity and moss coverage and a significant positive correlation between flow velocity and cyanobacteria coverage. The relationship between moss cove-rage ( x ) and flow velocity ( y ) was y =-2.081 x +0.03 ( R 2 =0.469). The moss coverage was a key factor affecting the flow velocity of co-covering of plant and biocrust slope with similar plant coverage (40%±10%). In conclusion, biocrusts under plant cover significantly slowed flow velocity, and the effect magnitude was related to its composition, implying that the role of biocrusts should be considered in understanding the mechanism underlying slope erosion in revegetated grassland.

The changes in soil properties caused by grazing and trampling are important reasons for the changes in soil respiration rates, carbon fixation, and emission. However, the effects of different intensities of grazing and trampling on biocrusts respiration rate are unclear. In this study, we simulated grazing and trampling disturbances of 10%, 30%, 50%, and 70% intensity on moss biocrusts developed in aeolian sandy soil on Loess Plateau, with undisturbed moss biocrusts serving as a control. The changes in respiration rate of moss biocrusts were monitored continuously, and its responses to different disturbance intensities were analyzed. The results showed that: 1) moderate disturbance stimulated moss biocrusts respiration, while heavy disturbance inhibited that. The respiration rate of moss biocrusts was increased by 41.1% and 22.2% at disturbance intensities of 10% and 30%, but was decreased by 8.9% and 15.3% at disturbance intensities of 50% and 70%, respectively. 2) The trampling disturbance significantly changed soil temperature but did not affect soil water content. In comparison to the control, soil temperature of biocrusts was decreased by 0.4 and 1.2 ℃ at disturbance intensities of 10% and 30%, but it was increased by 1.1 and 1.0 ℃ at disturbance intensities of 50% and 70%, respectively. 3) The respiration rate of moss biocrusts showed a significant exponential relationship with soil temperature and a linear positive relationship with soil water content under different disturbance intensities. However, the correlation between respiration rate of moss biocrust and the characteristics of moss biocrust was not significant. Soil temperature and water content could explain 70.6%-96.3% and 49.1%-70.0% of the total variation of respiration rate of moss biocrusts, respectively. In conclusion, grazing and trampling affected the respiration rate of moss biocrusts, with short-term moderate grazing and trampling would have positive effects. On the other hand, excessive grazing and trampling would reduce the rate of moss biocrust respiration. As a result, future studies on soil carbon balance of the Loess Plateau should consider the effects of grazing and trampling on biocrust respiration.

Mosses are poikilohydric plants. The duration of leaf spreading time is a key factor affecting their growth and development in the field. The dynamics of field growth and development and influencing factors of mosses in arid and semi-arid areas are largely unknown. In the study, we examined leaf spreading situation under natural conditions from September 5th to November 25th for Didymodon vinealis and Barbula unguiculata , two common species in hilly Loess Plateau. The results showed that the leaves of both species showed a regular diurnal dynamic change of 'spreading-closing-spreading' from September to October, and that the average leaf closing time of D. vinealis in the morning was 0.68 hours earlier than that of B. unguiculata , while leaf spreading time was delayed by 1.79 hours in the afternoon. Both species spread their leaves for longer time in the rainy season. The average leaf spreading duration of D. vinealis was 251 min, which was 30.4% lower than B. unguiculata of 361 min. The relative humidity near the surface was the key factor affecting leaf spreading duration. The morphological structure of moss species would affect leaf spreading duration. Compared with D. vinealis , B. unguiculata was relatively short, with a large proportion of costa in leaves, and the mosaic structure of stem cortex cells was more prominent. The humidity threshold during leaf spreading of B. unguiculata (54.3%) was lower than that of D. vinealis (60.1%). The leaf spreading duration was mainly affected by humidity. B. unguiculata was more adaptable to the environment than D. vinealis .

Exploring and quantifying the impacts of biological soil crusts on soil hydrological processes and soil water budget in semi-arid ecosystems can provide a theoretical basis for vegetation restoration and reconstruction in deserts. Based on continuous observation of soil water content in different types of areas covered by biological soil crusts ( e.g. , algae, moss) and bare sand in the Mu Us sandy land during the growing season (May to October) from 2018 to 2020, we examined the effects of biological soil crusts on soil water budget at a depth of 0-40 cm. Results showed that algae and moss crusts significantly reduced soil water supplement below 40 cm by rainfall and increased soil water evaporation loss, compared with that under bare sand. In the relatively wet year (2018), the amount of soil water expenditure (seepage+evaporation) covered by bare sand and the various types of biological soil crusts was less than that of rainfall, resulting in net soil water income. In the relative dry years (2019 and 2020), the amount of soil water expenditure covered by dominant algae and moss crusts was higher than that of rainfall, causing net soil water deficit, but opposite for bare sand. Biological soil crusts led to the imbalance of soil water budget of 0-40 cm depth and even soil water deficit in relatively dry years, which may lead to the succession of plant communities to be dominated by shallow-rooted plants in this area.

The distribution characteristics of exogenous carbon (C) in the C fractions of biocrusts-covered soil are critical for understanding the geochemical cycling of C with biocrusts in drylands. A 13 C pulse labeling experiment was conducted for moss-dominated biocrusts-covered soil and bare soil on the Loess Plateau of China with semiarid climate, with the content of 13 C in different C fractions being continuously measured to determine the biocrust effects on the distribution of exogenous C in each C fraction. Our results showed that, 1) the 13 C abundance of each C fraction in the biocrusts-covered soil was steadily changed with time, due to the relatively low rate of nutrient cycling in the biocrusts-covered soil and also to the relatively low biomass of moss in the biocrusts-covered soil as compared with vascular plants. 2) The 13 C content of each C fraction in the biocrusts-covered soil was significantly higher than that in the bare soil. Specifically, the 13 C content of total organic C (TOC), microbial biomass C (MBC), and dissolved organic C (DOC) in the biocrusts-covered soil was 0.258, 0.078, and 0.004 mg·kg -1 , respectively, which was 3.1, 18.5, and 2.6 times higher than that in the bare soil. Moreover, the 13 C content in the moss of the biocrusts-covered soil was 1.45 mg·kg -1 . 3) The presence of biocrusts changed the distribution characteristics of each C fraction, with the newly assimilated C being mainly distributed in active organic C and biological components of the biocrusts-covered soil. In the biocrusts-covered soil, the 13 C distribution in MBC (30.6%) was higher than that in DOC (1.7%), and the 13 C distribution in the C of moss was 20.3%. 4) The transferred amount and storage capacity of MB 13 C in the biocrusts-covered soil were 15.7 and 19.5 times of that in the bare soil, respectively. The turnover rate of MB 13 C in the biocrusts-covered soil and bare soil was 2.94 and 3.30 times per month, respectively, implying that the turnover time of MB 13 C in the biocrusts-covered soil was 1.1 times longer than that in the bare soil. In conclusion, biocrusts could greatly change the distribution characteristics of each C fraction and increase C turnover rate, highlighting its important roles in C cycling in dryland ecosystems.

Biological soil crusts (BSCs) greatly change surface soil structure and nutrient enrichment processes in arid and semiarid regions. However, their impacts on solute transport characteristics and nutrient loss are still not clear. In this study, the solute (Cl - and Ca 2+ ) transport experiments were conducted on soils covered by moss-dominated BSCs and uncrusted soil on sandy and loessal soils on the Loess Plateau, respectively. We analyzed the solute transport characteristics of the BSCs covered soil and uncrusted soil in different soil depths (0-5 cm and 5-10 cm). The BSCs mulching generated delay effects on the solute breakthrough process of 0-5 cm soils. The breakthrough time of Cl - in the BSCs covered soil was 3.83 (sandy soil) and 2.09 times (loessal soil) longer than that in the uncrusted soil. The breakthrough time of Ca 2+ in the BSCs covered soil was 2.50 and 2.73 times longer than that in the uncrusted soil. Due to the strong influence of BSCs mulching, the pore volume number of the complete solute breakthrough at 0-5 cm depth was higher than that at 5-10 cm depth in the BSCs covered soils. The breakthrough time of Cl - at 0-5 cm depth was increased by 67.3% (sandy soil) and 51.8% (loessal soil) by the BSCs as compared with that at 5-10 cm depth. The breakthrough time of Ca 2+ at 0-5 cm depth was increased by 8.0% and 33.7% by the BSCs. The BSCs reduced soil pore water flow velocity by 37.5%-70.2% compared with the uncrusted soil. Except for the sandy soil at 5-10 cm depth, the BSCs increased the solute dispersion coefficient by 1.73-6.29 times and the degree of dispersion by 2.77-20.95 times compared with the uncrusted soils. After the complete breakthrough of solute, the content of Ca 2+ in the BSCs layer (0-2 cm) was 4.14 and 2.58 times higher than that in the uncrusted sandy and loessal soils, respectively. In conclusion, our results indicated that BSCs could reduce the deep percolation and loss of nutrients accumulated in surface soil through improving their solute adsorption and retention abilities, which is of great significance for the improvement of soil fertility and vegetation restoration on degraded land in arid and semiarid regions.

Nitrogen (N) labeled with 15 N was evenly added into plots of moss-dominated biological soil crusts (BSCs) and bare soil on the Chinese Loess Plateau. After that, the surface BSCs and bare soil samples were continuously collected within 1-30 days. The 15 N content of each N fraction in soil, microorganisms, and mosses was measured for each sample. The effects of BSCs on soil N fate and cycling was determined through analyzing the differences in the distribution of 15 N fractions between the BSCs and bare soil. Our results showed that: 1) The 15 N content of total N (TN), microbial biomass N (MBN), and dissolved organic N (DON) in the BSCs was 2.9, 17.5, and 9.0 times higher than that in the bare soil, respectively. The 15 N content of moss plants in the BSCs was 4.73 mg kg -1 . 2) The residual rate of 15 N in the BSCs and bare soil was 13.0% and 3.3%, respectively, indicating that the N fixing and holding ability of BSCs was four times higher than that of bare soil. The percentage of each 15 N fraction in T 15 N in the BSCs was in the order of MBN (54.3%)&gt;moss plant N (22.5%)&gt;DON (6.2%), while that in the bare soil was in the order of MBN (11.5%)&gt;DON (2.6%). Over all, microorganisms and mosses in the BSCs had 65.3% higher capacity of N fixation as compared with the bare soil. 3) The transferred amount and storage capacity of MB 15 N in the BSCs were 17.2 and 20.5 times higher than that in the bare soil, respectively. Accordingly, the turnover rate of MB 15 N in the BSCs and bare soil was 5.8 and 7.2 times per month, respectively, with the turnover time of MB 15 N in the BSCs being 1.2 times longer than that in bare soil. In conclusion, BSCs fix and hold more N than bare soil and change the distribution of each N fraction, implying that BSCs play a critical role in N cycling in dryland ecosystems.

Three monitoring plots were established in the Yangming Mountains mossy dwarf forest concentration zone in the Yangmingshan National Nature Reserve in Hunan Province in 2005. The changes of species composition and diversity during 2005-2016 were analyzed from species numerical characteristics, species importance value, diameter class structure, height structure, and species diversity. The results showed that from 2005 to 2016, the individual number for ≥1.0 cm diameter at ground height (DGH) in tree layer decreased by 30.9%. Viburnum dilatatum decreased greatest with 98.6%, and Rhododendron fortunei decreased least with 1.8%. Species richness was increased, and evenness was decreased. Except for R. fortunei, the importance values of all the species decreased significantly after ten years, indicating that R. fortunei was the dominant species in the dwarf forest. The diameter-class distribution of species in the dwarf forest was arranged at 1 cm level. The density of small-diameter (1 cm≤DGH＜6 cm) species was higher, but that of medium-diameter (6 cm≤DGH＜15 cm) and large-diameter (15 cm≤DGH) was lower. The diameter-class distribution of R. fortunei exhibited a reversed 'J' shape, and the height of community increased from 2005 to 2016, indicating the population had a stable and well-developing trend. The other species were poorly renewed and their diameter-class structure was irregular. Generally, the Patrick and Menhinick richness indices and the Simpson index were increased, and the Shannon and Pielou indices were decreased during the ten years.

Bryophytes are morphologically special higher plants, with unique ornamental values and a wide application prospect. Its species richness was just secondary to angiosperms. To more effectively utilize bryophytes for greening and landscape construction, 77 plots in 10 green spaces were investigated in urban area of Nanjing. 55 species belonging to 36 genera and 21 families were recorded, among which Pottiaceae and Thuidiaceae were widely distributed. The species richness in green space gradually reduced from the center to the surrounding areas of the city. The 10 green spaces could be clustered into three groups based on bryophyte diversity, habitats, human distur-bance frequency, green space areas. Results from Canonical correspondence analysis (CCA) showed that canopy density, humidity, and substrate types were the major environmental factors the distribution of bryophytes. In Nanjing, most bryophytes preferred to grow in moderate humidity and open areas.

The experiment was conducted in the plots that had been enclosed for nearly 20 years in the hilly Loess Plateau region. The effects of trampling disturbance on the biological soil crust (biocrust) coverage, soil organic carbon (SOC), soil easily oxidizable carbon (SEOC), SOC mineralization amount and mineralization rate were investigated. The biocrust SOC mineralization potential after disturbance in different soil layers were simulated by a first-order kinetic equation. The results showed that the coverage of cyanobacteria biocrust and moss biocrust significantly decreased with the increases of disturbance intensity. Compared to no disturbance, the cyanobacteria coverage declined by 264%-339% and moss coverage declined by 46%-127%. Compared to no disturbance, SOC content in biocrust layer significantly decreased by 211%-300%. No significant difference was found among the five disturbance intensities. Disturbance increased SEOC content in biocrust layer, and the variation amounted to 1.5-3.4 g·kg -1 , with 30%, 40% and 50% distur-bance differed significantly to no disturbance. Disturbance significantly increased biocrust SOC mineralization cumulative amount. However, SOC mineralization rate did not show any significant change. The SOC mineralization cumulative amount under 40% disturbance intensity increased by 77% compared to that under no disturbance. Disturbance significantly increased SOC mineralization potential in biocrust layer, but with no influence in 0-2 cm and 2-5 cm soil layers. The 40% disturbance intensity significantly increased SOC mineralization potential by 4.7 g·kg -1 . The results of principal component analysis showed that SOC, SEOC, SMC and mineralization rate explained 76.7% of the variation of SOC mineralization potential in biocrust layer. Disturbance might be a main factor driving the increases of biocrust SOC mineralization potential in this area.

Based on the measurements of the fluxes of CO 2 , CH 4 and N 2 O from the soil covered by two types of biocrusts dominated separately by moss and algae-lichen, followed by 0 (control), 1 (shallow) and 10 (deep) mm depths of sand burial treatments, we studied the effects of sand burial on greenhouse gases fluxes and their relationships with soil temperature and moisture at Shapotou, southeastern edge of the Tengger Desert. The results showed that sand burial had significantly positive effects on CO 2 emission fluxes and CH 4 uptake fluxes of the soil covered by the two types of biocrusts, but imposed differential effects on N 2 O fluxes depending on the type of biocrust and the depth of burial. Deep burial (10 mm) dramatically increased the N 2 O uptake fluxes of the soil co-vered by the two types of biocrusts, while shallow burial (1 mm) decreased the N 2 O uptake flux of the soil co-vered by moss crust only and had no significant effects on N 2 O uptake flux of the soil covered by algae-lichen crust. In addition, CO 2 fluxes of the two biocrusts were closely related to the soil temperature and soil moisture, thereby increasing with the raised soil surface temperature and soil moisture caused by sand burial. However, the relationships of burial-induced changes of soil temperature and moisture with the changes in the other two greenhouse gases fluxes were not evident, indicating that the variations of soil temperature and moisture caused by sand burial were not the key factors affecting the fluxes of CH 4 and N 2 O of the soil covered by the two types of biocrusts.

Seasonal variations of soil inorganic nitrogen (N) pool and net N transformation rate in moss-covered soil and in the bare soil were comparatively observed by incubating intact soil columns with parafilm capping in the field in a natural vegetation area of Shapotou, southeastern fringe of the Tengger Desert. We found pronounced seasonal variations in soil available N content and net N transformation rate in both moss-covered soil and bare soil, with significant differences among different months. In non-growing season, soil available N content and net N transformation rate were significantly higher in March and October than in other months. Furthermore, immobilization was the dominant form of N mineralization, and no significant difference in net soil N mineralization rate was found between the two sampling soils. In growing season, soil available N content and net N transformation rate markedly increased and reached their peak values during June to August (17.18 mg x kg(-1) and 0.11 mg x kg(-1) x d(-1), respectively). Both soil net nitrification and N mineralization rates in moss-covered soil were significantly higher than in bare soil. Soil ammonium and nitrate N content in April and May were higher in moss-covered soil (2.66 and 3.16 mg x kg(-1), respectively) than in bare soil (1.02 and 2.37 mg x kg(-1), respectively); while the tendency was the converse in June and September, with 7.01 mg x kg(-1) for soil ammonium content and 7.40 mg x kg(-1) for nitrate N content in bare soil, and they were 6.39 and 6.36 mg x kg(-1) in moss-covered soil, respectively. Therefore, the existence and succession of moss crusts could be considered as one of the important biological factors affecting soil N cycling through regulating soil available N content and promoting soil N mineralization process.

Two types of soil covered by biological soil crusts (BSCs) , i.e. moss and algae, and moving sand in the natural vegetation area at the southeast fringe of the Tengger Desert were collected intactly. They were incubated continuously for 20 days under two different temperatures (15 degrees C and 25 degrees C) and moistures (10% and 25%) condition in the laboratory, and soil NO3(-)-N contents were measured after 1, 2, 5, 8, 12, 20 days of incubation and net nitrification rate was evaluated during dehydration. The results showed that NO3(-)-N content of the moss-covered soil (2.29 mg x kg(-1)) was higher than that of the algae-covered soil (1.84 mg x kg(-1)) and sand (1.59 mg x kg(-1)). Net nitrification rate of the three soil types ranged from -3.47 to 2.97 mg x kg(-1) x d(-1). For the moss-covered soil and algae-covered soil at 10% and 25% moisture levels, the net nitrification rates at 15 degrees C were 75.1%, 0.7% and 99.1%, 21.3% higher than those at 25 degrees C, respectively. Also, the net nitrification rates at 15 degrees C and 10% moisture levels were 193.4% and 107.3% higher than those at 25 degrees C and 25% moisture levels, respectively. The results suggested that regardless of soil moisture increasing or decreasing under the global warming senior, the net nitrification rate of BSCs-soil system in the desert would probably be limited to some extent during drought process.

Biological soil crusts play an important role in increasing the accumulation of organic matter and nitrogen in re-vegetated mining wastelands. The diversity of nitrogen-fixing microorganisms in three types of biological soil crusts (algal crust, moss crust and algal-moss crust) from two wastelands of copper mine tailings were investigated by polymerase chain reaction-denaturing gradient gel electrophoresis, based on the nifH gene of diazotrophs, to investigate: The diversity of nifH gene in the crusts of mine wastelands, and whether and how the nifH gene diversity in the crusts could be affected by the development of plant communities. The algal crust on the barren area displayed the highest nifH gene diversity, followed by the algal-moss crusts within vascular plant communities, and the moss crust displayed the lowest nifH gene diversity. The diversity of diazotrophs in algal-moss crust within vascular plant communities decreased with the increase of height and cover of vascular plant communities. No significant relationship was found between wasteland properties (pH, water content, contents of organic matter, nitrogen and phosphorus and heavy metal concentration) and nifH gene diversity in the crusts. Sequencing and phylogenetic analysis indicated that most nitrogen-fixing taxa in the crusts of mine wastelands belonged to Cyanobacteria, especially nonheterocystous filamentous Cyanobacteria.

Soil surface roughness exerts profound influence on runoff and sediments yield from slopes. Biological soil crusts (BSCs) are ubiquitous living cover in the open spaces between plants in arid and semiarid ecosystems, which considerably impact the surface roughness. Aimed to determine the effect of biological crusts on the surface roughness and its influence factors, this study investigated the surface roughness of soil with BSCs at their different successional stages by using the chain method. Besides, the impacts of slope aspects, soil water content and freezing and thawing on surface roughness were also investigated. Then, the correlations between roughness and soil chemical and physical properties were evaluated. The results showed that soil surface roughness decreased at the initial successional stage of BSCs but increased along with BSCs' development in the region. Soil roughness tended to be stabile since BSCs developed ten years later. The roughness was reduced by 47.0% by the light cyanobacteria-dominated crust and 20.4% by the dark cyanobacteria-dominated crust (moss coverage < 20%), compared with the bare soil. Soil moisture significantly impacted surface roughness. The surface roughness of BSCs changed obviously with soil moisture in the early development, but only slightly in their later succession. Freezing-thawing also affected surface roughness with BSCs. Roughness was more susceptible to freezing-thawing in the early de- velopment of BSCs, with an increase of 29.7% compared with the bare soil. The surface roughness of late successional BSCs exhibited a slight change only after repeated freezing-thawing. The surface roughness of BSCs showed a statistically significant relationship with the coverage of moss (P < 0.1).

By using water drop penetration time (WDPT) and molarity of ethanol droplet (MED) methods, the soil water repellence of undisturbed biological soil crusts (biocrusts) in five successional stages, from the hilly Loess Plateau region of China was tested. The five stages of biocrusts were light cyanobacterial crust, dark cyanobacterial crust, cyanobacterial with sparse moss crust, moss and tiny cyanobacteria patches crust and moss dominated crust. The results showed that 1) the soil water repellence was markedly increased both in the intensity and persistence since the formation of biocrusts. 2) The soil water repellence showed a decrease trend along with the successional stages of biocrusts. The soil water repellence of the biocrusts with the moss coverage above 20% was significantly lower than that of the cyanobacterial crusts. 3) The soil water repellence of the biocrusts was closely related to soil moisture and the dominant organism. The soil water repellence increased with the decrease of soil water content for the moss dominated biocrusts, while changed in a bimodal curve with the decrease of soil water content for the cyanobacterial biocrusts.

Undisturbed soil cores were incubated in laboratory at different temperatures (-10, 5, 15, 25, 35 and 40 degrees C) and moistures (29%, 58%, 85% and 170% of field water content, FWC) to analyze the effect of moss crusts, which were sampled from the natural vegetation area of Shapotou, on soil nitrogen transformation and their responses to hydrothermic factors. Results showed that immobilization was the dominant form of nitrogen transformation at the lower temperatures (<15 degrees C), and when the temperature surpassed 25 degrees C, the nitrogen transformation rate significantly increased. The nitrogen transformation rate in the moss-covered soil was more sensitive to temperature variation than in the bare soil, and the highest temperature sensitivity was at 85% FWC. It indicated that the existence and succession of moss crusts facilitated nitrogen transformation. In addition, the nitrogen transformation rates of two microhabitats increased initially and then declined with the increasing moisture, and the maximum nitrogen transformation rate was observed at 85% FWC. Significant interactive effects were found between temperature and moisture in the moss-covered soil. While, the greater enhancement in nitrogen transformation rate appeared at higher temperatures (25-40 degrees C ) and moderate moisture levels (58% FWC and 85% FWC). It was concluded that the existence and succession of moss crusts would increase the ability of soil nitrogen supply, promote nitrogen cycling and even contribute to the restoration of soil ecosystem.

Uncertainties still existed for evaluating greenhouse gases fluxes (GHGs), including carbon dioxide (CO2), methane (CH4) and nitrous oxide (N2O) at the regional scale for desert ecosystem because available GHGs data about biological soil crusts (BSCs) was very scarce. In 2011 and 2012, soil ecosystem covered by various types of BSCs and BSCs at different succession stages in an artificial sand-fixing vegetation region established in various periods at southeast of the Shapotou area in Tengger Desert was selected to measure fluxes of CO2, CH4 and N2O using static chamber and gas chromatography. The results showed that curst type, recovery time and their interactions with sampling date significantly affected CO2 flux. Recovery time and interaction of crust type and sampling date significantly affected CH4 flux. Sampling date significantly affected the fluxes of CO2, CH4 and N2O. The mean annual flux of CO2 for moss crust (105.1 mg x m(-2) x h(-1)) was significantly higher than that of algae crust (37.7 mg x m(-2) x h(-1)) at the same succession stage. Annual mean CH4 and N2O consumption was 19.9 and 3.4 microg x m(-2) x h(-1), respectively. Mean annual consumption of CH4 and N2O for algae crust was slightly higher than that of moss crust, however, significant difference was not found. Ecosystem respiration (Re) of desert soil covered by BSCs increased with the recovery process of desert ecosystem, in contrast, consumption of CH4 and N2O decreased. Re of moss crust was more sensitive to temperature and moisture variation than algae crust and Re sensitivity of temperature and moisture gradually increased with the development and succession of BSCs. Both soil temperature and moisture were not the main factor to determine CH4 and N2O fluxes of BSCs-soil in desert ecosystem.

Vegetation indexes were the most common and the most important parameters to characterizing large-scale terrestrial ecosystems. It is vital to get precise vegetation indexes for running land surface process models and computation of NPP change, moisture and heat fluxes over surface. Biological soil crusts (BSC) are widely distributed in arid and semi-arid, polar and sub-polar regions. The spectral characteristics of dry and wet BSCs were quite different, which could produce much higher vegetation indexes value for the wet BSC than for the dry BSC as reported. But no research was reported about whether the BSC would impact on regional vegetation indexes and how much dry and wet BSC had impact on regional vegetation indexes. In the present paper, the most common vegetation index NDVI were used to analyze how the moss soil crusts (MSC) dry and wet changes affect regional NDVI values. It was showed that 100% coverage of the wet MSC have a much higher NDVI value (0.657) than the dry MSC NDVI value (0.320), with increased 0.337. Dry and wet MSC NDVI value reached significant difference between the levels of 0.000. In the study area, MSC, which had the average coverage of 12.25%, would have a great contribution to the composition of vegetation index. Linear mixed model was employed to analyze how the NDVI would change in regional scale as wet MSC become dry MSC inversion. The impact of wet moss crust than the dry moss crust in the study area can make the regional NDVI increasing by 0.04 (14.3%). Due to the MSC existence and rainfall variation in arid and semi-arid zones, it was bound to result in NDVI change instability in a short time in the region. For the wet MSC's spectral reflectance curve is similar to those of the higher plants, misinterpretation of the vegetation dynamics could be more severe due to the "maximum value composite" (MVC) technique used to compose the global vegetation maps in the study of vegetation dynamics. The researches would be useful for detecting and mapping MSC from remote sensing imagery. It also is to the advantage to employing vegetation index wisely.