Biological soil crusts and their potential applications in the sand land over Qinghai-Tibet Plateau

The Qinghai-Tibet Plateau is now experiencing ecological degradation risks as a result of climate change and human activities.The alpine grassland ecology in permafrost zones is fragile and susceptible to deterioration due to its high altitude,low temperature,and limited oxygen,which complicates the repair of damaged land.Biological soil crusts(BSCs)are crucial for land restoration in plateau regions because they can thrive in harsh conditions and have environmentally beneficial traits.Inoculated biological soil crust(IBSC)has shown success in low-altitude desert regions,but may not be easily duplicated to the plateau environment.Therefore,it is essential to do a comprehensive and multifaceted analysis of the basic theoretical comprehension and practical application of BSCs on the Tibetan Plateau.This review article aims to provide a brief summary of the ecological significance and the mechanisms related to the creation,growth,and progression of BSCs.It discusses the techniques used for cultivating BSCs in laboratories and using them in the field,focusing on the Qinghai-Tibet Plateau circumstance.We thoroughly discussed the potential and the required paths for further studies.This study may be used as a basis for selecting suitable microbial strains and accompanying supplemental actions for implementing IBSCs in the Qinghai-Tibet Plateau.

Responses of microbial activities and soil physical-chemical properties to the successional process of biological soil crusts in the Gurbantunggut Desert,Xinjiang

Biological soil crusts （BSCs） are capable of modifying nutrient availability to favor the establishment of biogeochemical cycles. Microbial activities serve as critical roles for both carbon and nutrient transformation in BSCs. However, little is known about microbial activities and physical-chemical properties of BSCs in the Gurbantunggut Desert, Xinjiang, China. In the present research, a sampling line with 1-m wide and 20-m long was set up in each of five typical interdune areas selected randomly in the Gurbantunggut Desert. Within each sampling line, samples of bare sand sheet, algal crusts, lichen crusts and moss crusts were randomly collected at the depth of 0-2 cm. Varia- tions of microalgal biomass, microbial biomass, enzyme activities and soil physical-chemical properties in different succession of BSCs were analyzed. The relationships between microalgal biomass, microbial biomass, enzymatic activities and soil physical-chemical properties were explored by stepwise regression. Our results indicate that micro- algal biomass, microbial biomass and most of enzyme activities increased as the BSCs developed and their highest values occurred in lichen or moss crusts. Except for total K, the contents of most soil nutrients （organic C, total N, total P, available N, available P and available K） were the lowest in the bare sand sheet and significantly increased with the BSCs development, reaching their highest values in moss crusts. However, pH values significantly decreased as the BSCs developed. Significant and positive correlations were observed between chlorophyll a and microbial biomass C. Total P and N were positively associated with chlorophyll a and microbial biomass C, whereas there was a significant and negative correlation between microbial biomass and available P. The growth of cyanobacteria and microorganism contributed C and N in the soil, which offered substrates for enzyme activities thus increasing enzyme activities. Probably, improvement in enzyme activities increased soil fertility and promoted the growth of cyanobacteria, eukary- otic algae and heterotrophic microorganism, with the accelerating succession of BSCs. The present research found that microalgal-microbial biomass and enzyme activities played important roles on the contents of nutrients in the successional stages of BSCs and helped us to understand developmental mechanism in the succession of BSCs.

Biological soil crust distribution in Artemisia ordosica communities along a grazing pressure gradient in Mu Us Sandy Land, Northern China

This study investigated the distribution pattern of biological soil crust （BSC） in Artemisia ordosica communities in Mu Us Sandy Land. Three experimental sites were selected according to grazing pressure gradient. In each experimental site, the total vegetation cover, A. ordosica cover, BSC cover, litter-fall cover, BSC degree of fragmentation, BSC thickness and soil properties were investigated in both fixed and semi-fixed sand dunes and simultaneously analyzed in the laboratory. The results showed that at the same grazing pressure, BSC cover and composition were significantly affected by the fixation degree of sand dunes. In addition, BSC cover in the fixed sand dunes was 83.74% on average, whereas it is proportionally dominated by 28% mosses, 21% lichens, and 51% algae. Meanwhile, BSC cover in the semi-fixed sand dunes was 23.54% on average, which is proportionally domi- nated by 6.3% mosses, 2.5% lichens, and 91.2% algae. Fine sand, organic matter, and total nitrogen （N） contents in the fixed sand dunes were all significantly higher than those in the semi-fixed sand dunes. Litter-fall cover de- creased along the grazing pressure gradient, whereas BSC fragmentation degree increased. Fine sand content decreased along with the increase of grazing pressure, whereas medium sand content increased in both fixed and semi-fixed dunes. The organic matter and total N contents in the no grazing site were significantly higher than those in light and normal grazing sites. However, there were no significant differences between the light and normal grazing sites. In addition, there were also no significant differences in BSC thickness between the light and normal grazing sites in the fixed sand dunes. However, a significant decrease was observed in both BSC cover and thick- ness in the normal grazing site. The BSC in the semi-fixed dunes was more sensitive to disturbance.

Effects of artificially cultivated biological soil crusts on soil nutrients and biological activities in the Loess Plateau

Biological soil crusts （BSCs） play an important role in the early succession of vegetation restoration in the Loess Plateau, China. To evaluate the effects of artificially cultivated BSCs on the soil surface micro-envir- onment, we obtained natural moss crusts and moss-lichen crusts from the Loess Plateau of Shaanxi province, and subsequently inoculated and cultivated on horizontal and sloping surfaces of loess soil in a greenhouse. The chemical and biological properties of the subsoil under cultivated BSCs were determined after 10 weeks of cul- tivation. The results indicated that BSCs coverage was more than 65% after 10 weeks of cultivation. Moss crust coverage reached 40% after 5 weeks of cultivation. Compared with the control, soil organic matter and available nitrogen contents in moss crust with the horizontal treatments increased by 100.87% and 48.23%, respectively; increased by 67.56% and 52.17% with the sloping treatments, respectively; they also increased in moss-lichen crust with horizontal and sloping treatments, but there was no significant difference. Available phosphorus in cultivated BSCs was reduced, soil pH was lower and cationic exchange capacity was higher in cultivated BSCs than in the control. Alkaline phosphatase, urease and invertase activities were increased in artificially cultivated BSCs, and alkaline phosphatase activity in all cultivated BSCs was obviously higher than that in the control. Numbers of soil bacteria, fungi and actinomycetes were increased in the formation process of cultivated BSCs. These results indicate that BSCs could be formed rapidly in short-term cultivation and improve the mi- cro-environment of soil surface, which provides a scientific reference for vegetation restoration and ecological reconstruction in the Loess Plateau. China.

The effects of extreme rainfall events on carbon release from biological soil crusts covered soil in fixed sand dunes in the Tengger Desert, northern China

In May to August of 2011, we assessed the effects of extreme rainfall （quantity and intensity） events on the carbon release from soils covered by different types of biological soil crusts （BSCs） in fixed sand dunes in the Tengger Desert, northern China. A Li-6400-09 Soil Chamber was used to measure the respiration rates of the BSCs immediately after the rainfall stopped, and continued until the respiration rates of the BSCs returned to the pre-rainfall basal rate. Our results showed that almost immediately after extreme rainfall events the respiration rates of algae crust and mixed crust were significantly inhibited, but moss crust was not significantly affected. The respiration rates of algae crust, mixed crust, and moss crust in extreme rainfall quantity and intensity events were, respectively, 0.12 and 0.41 μmolCO2/（m2.s）, 0.10 and 0.45 gmolCO2/（m2·s）, 0.83 and 1.69 gmolCO2/（m2.s）. Our study indicated that moss crust in the advanced succession stage can well adaot to extreme rainfall events in the short term.

Precipitation and soil particle size co-determine spatial distribution of biological soil crusts in the Gurbantunggut Desert, China

Biological soil crusts （BSCs） are bio-sedimentary associations that play crucial ecological roles in arid and semi-arid regions. In the Gurbantunggut Desert of China, more than 27% of the land surface is characterized by a predominant cover of lichen-dominated BSCs that contribute to the stability of the desert. However, little is known about the major factors that limit the spatial distribution of BSCs at a macro scale. In this study, the cover of BSCs was investigated along a precipitation gradient from the margins to the center of the Gurbantunggut Desert. Environmental variables including precipitation, soil particle size, soil pH, electrical conductivity, soil organic carbon, total salt, total nitrogen, total phosphorus and total potassium were analyzed at a macro scale to determine their association with differing assemblages of BSCs （cyanobacteria crusts, lichen crusts and moss crusts） using constrained linear ordination redundancy analysis （RDA）. A model of BSCs distribution correlated with environmental variables that dominated the first two axes of the RDA was constructed to clearly demonstrate the succession stages of BSCs. The study determined that soil particle size （represented by coarse sand content） and precipitation are the most significant drivers influencing the spatial distribution of BSCs at a macro scale in the Gurbantunggut Desert. The cover of lichen and moss crusts increased with increasing precipitation, while the cover of cyanobacteria crusts decreased with increasing precipitation. The cover of lichen and moss crusts was negatively associated with coarse sand content, whereas the cover of cyanobacteria crusts was positively correlated with coarse sand content. These findings highlight the need for both the availability of soil moisture and a relatively stable of soil matrix, not only for the growth of BSCs but more importantly, for the regeneration and rehabilitation of disturbed BSC communities in arid and semi-arid lands. Thereby, this study will provide a theory basis to effectively increase soil stability in desert regions.

Distribution of algae and cyanobacteria of biological soil crusts along the elevation gradient in mountain plant communities at the Northern Urals(Russian European Northeast)

This paper describes the biodiversity of cyanobacteria and microalgae of biological soil crusts(BSC)on bare substrates in different mountain vegetation types at the Northern Urals.In total,we identified 99 algal species from six divisions in all sampled sites.The species diversity and structure of BSC algal communities show a relationship with environmental factors(altitude,soil p H and humidity,and illumination).Taxonomic diversity of algae decreases along the altitude gradient from mountain meadow to mountain tundra.Algae and cyanobacteria species from six divisions were identified in meadow communities,five in mountain forests and four in mountain tundra.We observed a positive correlation between species diversity of phototrophic microorganisms and altitude in the forest communities,but a negative correlation in the tundra.The dominant complex of cyanobacterial and algal species in BSC was specific for each type of plant community and was reflective of the habitat conditions.The species diversity and morphological organization of the BSC algae thalli can be used as a criterion for the ongoing assessment of climatic changes in high latitudes and mountain regions.

Snowpack shifts cyanobacterial community in biological soil crusts

Winter snowpack is an important source of moisture that influences the development ofbiological soil crusts(BSCs)in desert ecosystems.Cyanobacteria are important photosynthetic organismsin BSCs.However,the responses of the cyanobacterial community in BSCs to snowpack,snow depth andmelting snow are still unknown.In this study,we investigated the cyanobacterial community compositionand diversity in BSCs under different snow treatments(doubled snow,ambient snow and removed snow)and three snow stages(stage 1,snowpack;stage 2,melting snow;and stage 3,melted snow)in theGurbantunggut Desert in China.In stages 1 and 2,Cyanobacteria were the dominant phylum in the bacterialcommunity in the removed snow treatment,whereas Proteobacteria and Bacteroidetes were abundant inthe bacterial communities in the ambient snow and doubled snow treatments.The relative abundances ofProteobacteria and Bacteroidetes increased with increasing snow depth.The relative abundances ofCyanobacteria and other bacterial taxa were affected mainly by soil temperature and irradiance.In stages 2and 3,the relative abundance of Cyanobacteria increased quickly due to the suitable soil moisture andirradiance conditions.Oscillatoriales,Chroococcales,Nostocales,Synechococcales and unclassifiedCyanobacteria were detected in all the snow treatments,and the most dominant taxa were Oscillatorialesand Chroococcales.Various cyanobacterial taxa showed different responses to snowpack.Soil moisture andirradiance were the two critical factors shaping the cyanobacterial community structure.The snowpackdepth and duration altered the soil surface irradiance,soil moisture and other soil properties,whichconsequently were selected for different cyanobacterial communities.Thus,local microenvironmentalfiltering(niche selection)caused by snow conditions may be a dominant process driving shifts in thecyanobacterial community in BSCs.

Discussion on wind factor influencing the distribution of biological soil crusts on surface of sand dunes

Biological soil crusts are widely distributed in arid and semi-arid regions, whose formation and development have an important impact on the restoration process of the desert ecosystem. In order to explore the relationship between surface airflow and development characteristics of biological soil crusts, we studied surface airflow pattern and development characteristics of biological soil crusts on the fixed dune profile through field observation. Results indicate that the speed of near-surface airflow is the lowest at the foot of windward slope and the highest at the crest, showing an increasing trend from the foot to the crest. At the leeward side, although near-surface airflow increases slightly at the lower part of the slope after an initial sudden decrease at upper part of the slope, its overall trend decreases from the crest. Wind velocity variation coefficient varied at different heights over each observation site. The thickness, shear strength of biological soil crusts and percentage of fine particles at crusts layer decreased from the slope foot to the upper part, showing that biological soil crusts are less developed in high wind speed areas and well developed in low wind speed areas. It can be seen that there is a close relationship between the distribution of biological soil crusts in different parts of the dunes and changes in airflow due to geomorphologic variation.

Weathering of Carbonate Rocks by Biological Soil Crusts in Karst Areas

The weathering of carbonate rocks by biological soil crusts （BSC） in karst areas is very common. It is helpful to understand the weathering mechanisms and processes for avoiding karst rock-desertification. The weathering of carbonate rocks by BSC in karst areas, namely the expansion, contraction and curl resulting from environmental wetting-drying cycles, was investigated and ana- lyzed in this paper. The bulk density, area and thickness of BSC were determined and the weathering amount of limestone and dolomite per unit area of BSC was calculated as 3 700 and 3 400 g·m-2; the amount of biomass on the surface of limestone and dolomite was calculated as 1 146 and 1 301 g·m-2, respectively. Such an increased weathering amount was not only the result of chemical and physical weathering of BSC on carbonate rocks, but also the attachment and cementation of BSC to clay particles, dust-fall, sand particles, solid particles brought by strong air currents, wind and other factors in the surrounding environment, which may also be related to the special environment and the special time period. Based on the results obtained, a weathering mode of BSC is studied, and the mechanisms of weathering by BSC are discussed. In conclusion, we suggest that the mechanical force exerted by the expansion and constriction of gelatinous and mucilaginous substances through wetting and drying of BSC play a significant role in the physical weathering process of the carbonate substrates.

The influence of biological soil crusts on ^(15)N translocation in soil and vascular plant in a temperate desert of northwestern China

Aims Desert ecosystems are often characterized by patchy distribution of vascular plants,with biological soil crusts(BSC)covering interplant spaces.However,few studies have comprehensively examined the linkage between BSC and vascular plants through nitrogen(N)or element translocation.the objective of this study was to evaluate the ecological roles of BSC on N translocation from soil to the domi-nant herb Erodium oxyrrhynchum bieb.(geraniaceae)in a temper-ate desert in China.Methods Isotopes(including 15N-glu,15N-NH4Cl and 15N-NaNo3)were used as a tracer to detect translocation of N in two types of desert soil(BSC covered;bare)to the dominant herb E.oxyrrhynchum.three different forms of 15N-enriched N compounds were applied as a point source to small patches of BSC and to bare soil.and we measured isotopes(14N and 15N)and obtained the concentration of labeled-15N in both vascular plants and soils at different distances from substrate application Important Findings Plants of E.oxyrrhynchum growing in BSC-covered plots accumulated moreδ15N than those growing in the bare soil.similarly,soil from b Ccovered plots showed a higher concentration of labeled-N irrespective of form of isotope,than did the bare soil.the concentration of dissolved organic N(15N-glu)in E.oxyrrhynchum was higher than that of dis-solved inorganic N(15N-NH4Cl and 15N-NaNo3).soil covered by BSC also accumulated considerably more dissolved organic N than bare soil,whereas the dominant form of 15N concentrated in bare soil was dissolved inorganic N.Correlation analysis showed that the concentra-tion of labeled-N in plants was positively related to the concentration of labeled-N in soils and the N%recorded in E.oxyrrhynchum.our study supports the hypothesis that BSC facilitates ^(15)N translocation in soils and vascular plants in a temperate desert of northwestern China.

Ammonia-oxidizing archaea and bacteria are structured by geography in biological soil crusts across North American arid lands

Introduction:Biological soil crusts(BSCs)can dominate surface cover in dry lands worldwide,playing an integral role in arid land biogeochemistry,particularly in N fertilization through fixation and cycling.Nitrification is a characteristic and universal N transformation in BSCs that becomes important for the export of N beyond the microscopic bounds of the crust itself.The contribution of ammonia-oxidizing bacteria(AOB)in BSCs has been shown,but the role and extent of the recently discovered ammonia-oxidizing archaea(AOA)have not.Methods:We sampled various types of crusts in four desert regions across the western United States and characterized the composition and size of ammonia-oxidizing communities using clone libraries and quantitative PCR targeting the amoA gene,which codes for the ammonia monooxygenase enzyme,universally present in ammonia-oxidizing microbes.Results:All archaeal amoA sequences retrieved from BSCs belonged to the Thaumarchaeota(Nitrososphaera associated Group I.1b).Sequences from the Sonoran Desert,Colorado Plateau,and Great Basin were indistinguishable from each other but distinct from those of the Chihuahuan Desert.Based on amoA gene abundances,archaeal and bacterial ammonia oxidizers were ubiquitous in our survey,but the ratios of archaeal to bacterial ammonia oxidizers shifted from bacterially dominated in northern,cooler deserts to archaeally dominated in southern,warmer deserts.Conclusions:Archaea are shown to be potentially important biogeochemical agents of biological soil crust N cycling.Conditions associated with different types of BSCs and biogeographical factors reveal a niche differentiation between AOA and AOB,possibly driven by temperature.

Long-term light grazing does not change soil organic carbon stability and stock in biocrust layer in the hilly regions of drylands

Livestock grazing is the most extensive land use in global drylands and one of the most extensive stressors of biological soil crusts(biocrusts).Despite widespread concern about the importance of biocrusts for global carbon(C)cycling,little is known about whether and how long-term grazing alters soil organic carbon(SOC)stability and stock in the biocrust layer.To assess the responses of SOC stability and stock in the biocrust layer to grazing,from June to September 2020,we carried out a large scale field survey in the restored grasslands under long-term grazing with different grazing intensities(represented by the number of goat dung per square meter)and in the grasslands strictly excluded from grazing in four regions(Dingbian County,Shenmu City,Guyuan City and Ansai District)along precipitation gradient in the hilly Loess Plateau,China.In total,51 representative grassland sites were identified as the study sampling sites in this study,including 11 sites in Guyuan City,16 sites in Dingbian County,15 sites in Shenmu City and 9 sites in Ansai District.Combined with extensive laboratory analysis and statistical analysis,at each sampling site,we obtained data on biocrust attributes(cover,community structure,biomass and thickness),soil physical-chemical properties(soil porosity and soil carbon-to-nitrogen ratio(C/N ratio)),and environmental factors(mean annual precipitation,mean annual temperature,altitude,plant cover,litter cover,soil particle-size distribution(the ratio of soil clay and silt content to sand content)),SOC stability index(SI)and SOC stock(SOCS)in the biocrust layer,to conduct this study.Our results revealed that grazing did not change total biocrust cover but markedly altered biocrust community structure by reducing plant cover,with a considerable increase in the relative cover of cyanobacteria(23.1%)while a decrease in the relative cover of mosses(42.2%).Soil porosity and soil C/N ratio in the biocrust layer under grazing decreased significantly by 4.1%–7.2%and 7.2%–13.3%,respectively,compared with those under grazing exclusion.The shifted biocrust community structure ultimately resulted in an average reduction of 15.5%in SOCS in the biocrust layer under grazing.However,compared with higher grazing(intensity of more than 10.00 goat dung/m2),light grazing(intensity of 0.00–10.00 goat dung/m2 or approximately 1.20–2.60 goat/(hm2•a))had no adverse effect on SOCS.SOC stability in the biocrust layer remained unchanged under long-term grazing due to the offset between the positive effect of the decreased soil porosity and the negative effect of the decreased soil C/N ratio on the SOC resistance to decomposition.Mean annual precipitation and soil particle-size distribution also regulated SOC stability indirectly by influencing soil porosity through plant cover and biocrust community structure.These findings suggest that proper grazing might not increase the CO_(2) release potential or adversely affect SOCS in the biocrust layer.This research provides some guidance for proper grazing management in the sustainable utilization of grassland resources and C sequestration in biocrusts in the hilly regions of drylands.

Biocrust-induced partitioning of soil water between grass and shrub in a desert steppe of Northwest China

Maintaining the stability of exotic sand-binding shrub has become a large challenge in arid and semi-arid grassland ecosystems in northern China.We investigated two kinds of shrublands with different BSCs(biological soil crusts)cover in desert steppe in Northwest China to characterize the water sources of shrub(Caragana intermedia Kuang et H.C.Fu)and grass(Artemisia scoparia Waldst.et Kit.)by stable 18O isotopic.Our results showed that both shrublands were subject to persistent soil water deficiency from 2012 to 2017,the minimum soil depth with CV(coefficient of variation)<15% and SWC(soil water content)<6% was 1.4 m in shrubland with open areas lacking obvious BSC cover,and 0.8 m in shrubland covered by mature BSCs.For C.intermedia,a considerable proportion of water sources pointed to the surface soil.Water from BSCs contributed to averages 22.9%and 17.6%of the total for C.intermedia and A.scoparia,respectively.C.intermedia might use more water from BSCs in rainy season than dry season,in contrast to A.scoparia.The relationship between shrub(or grass)and soil water by δ^(18)O shown significant differences in months,which partly verified the potential trends and relations covered by the high variability of the water source at seasonal scale.More fine roots at 0-5 cm soil layer could be found in the surface soil layer covered by BSCs(8000 cm/m^(3))than without BSCs(3200 cm/m^(3)),which ensured the possibility of using the surface soil water by C.intermedia.The result implies that even under serious soil water deficiency,C.intermedia can use the surface soil water,leading to the coexistence between C.intermedia and A.scoparia.Different with the result from BSCs in desert areas,the natural withdrawal of artificial C.intermedia from desert steppe will be a long-term process,and the highly competitive relationship between shrubs and grasses also determines that its habitat will be maintained in serious drought state for a long time.

Soil total organic carbon/total nitrogen ratio as a key driver deterministically shapes diazotrophic community assemblages during the succession of biological soil crusts

The diazotrophic community in biological soil crusts(biocrusts)is the key supplier of nitrogen in dryland.To date,there is still limited information on how biocrust development influences the succession of diazotrophic community,and what are the most important factors mediating diazotrophic communities during biocrust succession.Using the high throughput nifH amplicon sequencing,the diazotrophs in soils at different developmental stages of biocrust were comparatively studied.The results evidenced the decreases of TOC/TN ratio and pH value with biocrust development.Nostoc and Scytonema were the most dominant diazotrophic genera at all biocrust stages,while Azospirillum and Bradyrhizobium were abundant only in bare soil.Diazotrophic co-occurrence networks tended to be less complex and less connected with biocrust succession.The soil TOC/TN ratio was the most dominant factor mediating diazotrophic diversity,community composition and assembly processes,while diazotrophic-diversity and NO3–-N/NH4+-N ratio were positively correlated with the nitrogenase activity during biocrust succession.This study provided novel understandings of nitrogen fixation and succession patterns of diazotrophic community,by showing the effects of biocrust succession on diazotrophic diversity,community composition,community assembly and co-occurrence networks,and recognizing TOC/TN ratio as the most dominant factor mediating diazotrophs during biocrust succession.

Effects of biological soil crusts on profile distribution of soil water,organic carbon and total nitrogen in Mu Us Sandland,China

Aims Biological soil crusts(BSCs)can affect soil properties including water dynamics and cycling of soil carbon and nitrogen in dryland ecosystems.Previous research has mostly focused on effects of BSCs on soil water distribution or carbon and nitrogen fixation in the surface soil layer.Thus,little is known about effects of BSCs on properties throughout the soil profile.In the current study,we assessed the effects of BSCs on the distribution of soil water content(SW),soil organic carbon content(SOC)and soil total nitrogen content(STN)throughout the soil profile as well as the influence of water conditions on the effects of BSCs.Methods In a field investigation in Mu Us Sandland,North China,soil samples were taken from plots with and without BSCs on 13 and 28 September 2006,respectively.On the two sampling dates,average soil gravimetric water content was 3.83%(61.29%)and 5.08%(60.89%),respectively,which were regarded as low and high water conditions.Soil samples were collected every 5 cm to a depth of 60 cm,and SW,SOC and STN were measured in the laboratory.Important Findings(i)BSCs affected profile distribution of SW,SOC and STN.In addition,water conditions within the plots significantly modified BSCs’effects on the profile distribution of SW,but marginally affected the effects on SOC and STN.(ii)Under high water conditions,SW in the surface soil layer(0–10 cm)was higher in soils with BSCs compared to those without BSCs,while the opposite was true in the deep soil layer(30–55 cm).(iii)Under low water conditions,SW was lower with BSCs compared with no BSCs in near-surface(5–20 cm)and deep(25–40 cm)soil layers.(iv)BSCs affected SOC and STN only in the surface soil layer(0–5 cm)and were modified by plot water conditions.

Long-term changes in biological soil crust cover and composition

Introduction:Communities change over time due to disturbances,variations in climate,and species invasions.Biological soil crust communities are important because they contribute to erosion control and nutrient cycling.Crust types may respond differently to changes in environmental conditions:single-celled organisms and bryophytes quickly recover after a disturbance,while lichens are slow growing and dominate favorable sites.Community change in crusts has seldom been assessed using repeated measures.For this study,we hypothesized that changes in crust composition were related to disturbance,topographic position,and invasive vegetation.Methods:We monitored permanent plots in the Columbia Basin in 1999 and 2010 and compared changes in crust composition,cover,richness,and turnover with predictor variables of herbivore exclosure,elevation,heat load index,time since fire,presence of an invasive grass,and change in cover of the invasive grass.Results:Bryophytes were cosmopolitan with high cover.Dominant lichens did not change dramatically.Indicator taxa differed by monitoring year.Bryophyte and total crust cover declined,and there was lower turnover outside of herbivore exclosures.Lichen cover did not change significantly.Plots that burned recently had high turnover.Increase in taxon richness was correlated with presence of an invasive grass in 1999.Change in cover of the invasive grass was positively related to proportional loss and negatively related to gain.Conclusions:Composition and turnover metrics differed significantly over 11 years,though cover was more stable between years.This study can be a baseline for assessing change in crust composition due to anthropogenic influences.

Chlorophytes of biological soil crusts in Gurbantunggut Desert,Xinjiang Autonomous Region,China

In this paper,chlorophytes collected from 253 biological soil crust samples in Gurbantunggut Desert in Xinjiang Autonomous Region,China were studied by field investigation and microscopical observation in lab.The flora composition,ecological distribution of chlorophytes in the desert and dynamic changes of species composition of chlorophytes in different developing stages of biological soil crusts are preliminarily analyzed.Results showed that there were 26 species belonging to 14 genera and 10 families,in which unicellular chlorophytes were dominant.There existed some differences in distribution of varied sand dune positions.The taxa of chlorophytes in leeward of sand dunes are most abundant,but the taxa in windward,interdune and the top of sand dunes reduced gradually.Chlorophytes were mainly distributed within the crust and the taxa of chlorophytes decrease obviously under the crust.In the devel-oping stages of the biological soil crust,species diversity of chlorophytes changed a little,but species composition pre-sented some differences.Chlorococcum humicola,Chlorella vulgaris,Chlamydomonas ovalis and Chlamydomonas sp.nearly existed in all developing stages of biological crusts.In several former stages of the biological soil crust there were spherical chlorophytes and filamentous ones.When moss crust formed,filamentous chlorophytes disappeared,such as Microspora and Ulothrix.

Progress in the study of algae and mosses in biological soil crusts

Algae and mosses are not only two of the familiar communities in the process of desert vegetational succession,but also have the highest biomass in biological soil crusts.Meanwhile,being the pioneer plants,algae and mosses are involved in the establishment of biological soil crusts,which have great importance in arid environments and play a major role in desert ecosystems,such as being the indicator of the vegetation type,soil-holding,preventing erosion by water and wind,and sand fixation.This paper reviews the advances in the study of algae and mosses in arid and semi-arid areas.It mainly describes the ecological functions of algae and mosses including their influences on water cycle,circulation of substances,and community succession.In addition,the relationships between algae and mosses are discussed.Finally,some suggestions are proposed for the research orientations of algae and mosses in biological soil crusts.Ecologically,algae and mosses have significant ecological importance in arid areas,especially in those areas where environmental problems are becoming increasingly serious.