Assessing recovery of biological soil crusts across a latitudinal gradient in Western Europe

Biological soil crusts are ecologically important communities in areas where vascular plant coverage is low, and their presence is often vital in prevention of soil erosion. Despite recurrent threats to biological soil crusts across different environments, their recovery after disturbance has been little studied. We therefore established experiments across a latitudinal gradient in Europe, from Öland, Sweden in the north, to Gössenheim, Germany and Hochtor, Austria, to Almeria, Spain in the south, spanning over 20° latitude and 2,300 m in altitude, and including natural and semi‐natural sites. At each site 10 (1 × 1 m) paired plots were constructed where the biological soil crusts were either completely removed, or left intact. Over a 2‐year period (2012–2014) the plots were regularly sampled to assess functional group recovery (cyanobacteria, algae, lichens, bryophytes, vascular plants), soil stability, and chlorophyll, carbon, and nutrient contents. Cyanobacterial assemblages were examined by denaturing gradient gel electrophoresis, a technique used to detect DNA in environmental samples. The results indicated that recovery was site dependent, suggesting that physical and climatic parameters play a major role in biological soil crust recovery. This conclusion is supported by the results of the soil properties, which were found to differ between sites, although they did not show meaningful recovery over the study period. Although 2 years was insufficient for pronounced biological soil crust recovery, this study documents changes over the initial recovery period, suggests management practices for future projects, and recommends proxies for measuring recovery over time.     

Warming reduces the growth and diversity of biological soil crusts in a semi-arid environment: implications for ecosystem structure and functioning

Biological soil crusts (BSCs) are key biotic components of dryland ecosystems worldwide that control many functional processes, including carbon and nitrogen cycling, soil stabilization and infiltration. Regardless of their ecological importance and prevalence in drylands, very few studies have explicitly evaluated how climate change will affect the structure and composition of BSCs, and the functioning of their constituents. Using a manipulative experiment conducted over 3 years in a semi-arid site from central Spain, we evaluated how the composition, structure and performance of lichen-dominated BSCs respond to a 2.4°C increase in temperature, and to an approximately 30 per cent reduction of total annual rainfall. In areas with well-developed BSCs, warming promoted a significant decrease in the richness and diversity of the whole BSC community. This was accompanied by important compositional changes, as the cover of lichens suffered a substantial decrease with warming (from 70 to 40% on average), while that of mosses increased slightly (from 0.3 to 7% on average). The physiological performance of the BSC community, evaluated using chlorophyll fluorescence, increased with warming during the first year of the experiment, but did not respond to rainfall reduction. Our results indicate that ongoing climate change will strongly affect the diversity and composition of BSC communities, as well as their recovery after disturbances. The expected changes in richness and composition under warming could reduce or even reverse the positive effects of BSCs on important soil processes. Thus, these changes are likely to promote an overall reduction in ecosystem processes that sustain and control nutrient cycling, soil stabilization and water dynamics.     

高寒草甸植被退化过程中生物土壤结皮演变特征

对生物土壤结皮(BSCs)在青藏高原的分布和作用还知之甚少,为了解其在高寒草甸退化过程中的演变特征,以高寒草甸为对象,研究其退化和人工重建对BSCs生长和分布影响。结果表明:随草甸退化,植被盖度不断降低,苔藓类结皮在禾草嵩草草甸和6年人工草地出现频度最高,随植被退化出现频度降低,"黑土滩"阶段最低(43.6%);黑斑、菌斑和地衣都是在矮嵩草阶段形成,在小嵩草草甸裂缝期或者剥蚀期出现频度最高,在禾草嵩草群落、6年人工草地及黑土滩均未出现;相关分析表明,苔藓类结皮出现频率与植被盖度呈极显著相关(P0.01),而与黑斑和地衣出现频率呈负相关,菌斑与地衣出现频度呈显著相关(P0.05)。BSCs在高寒草甸广泛分布,不同演替阶段,其分布和物种组成存在较大差别。植被生长和表土稳定性状况导致了BSCs的异质性分布格局。     

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.     

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.     

Biodiversity of Klebsormidium (Streptophyta) from alpine biological soil crusts (Alps,Tyrol, Austria,and Italy)

Forty Klebsormidium strains isolated from soil crusts of mountain regions (Alps, 600–3,000 m elevation) were analyzed. The molecular phylogeny (internal transcribed spacer rDNA sequences) showed that these strains belong to clades B/C, D, E, and F. Seven main (K. flaccidum, K. elegans, K. crenulatum, K. dissectum, K. nitens, K. subtile, and K. fluitans) and four transitional morphotypes (K. cf. flaccidum, K. cf. nitens, K. cf. subtile, and K. cf. fluitans) were identified. Most strains belong to clade E, which includes isolates that prefer humid conditions. One representative of the xerophytic lineage (clade F) as well as few isolates characteristic of temperate conditions (clades B/C, D) were found. Most strains of clade E were isolated from low/middle elevations (<1,800 m above sea level; a.s.l.) in the pine‐forest zone. Strains of clades B/C, D, and F occurred sporadically at higher elevations (1,548–2,843 m a.s.l.), mostly under xerophytic conditions of alpine meadows. Comparison of the alpine Klebsormidium assemblage with data from other biogeographic regions indicated similarity with soil crusts/biofilms from terrestrial habitats in mixed forest in Western Europe, North America, and Asia, as well as walls of buildings in Western European cities. The alpine assemblage differed substantially from crusts from granite outcrops and sand dunes in Eastern Europe (Ukraine), and fundamentally from soil crusts in South African drylands. Epitypification of the known species K. flaccidum, K. crenulatum, K. subtile, K. nitens, K. dissectum, K. fluitans, K. mucosum, and K. elegans is proposed to establish taxonomic names and type material as an aid for practical studies on these algae, as well as for unambiguous identification of alpine strains. New combination Klebsormidium subtile (Kützing) Mikhailyuk, Glaser, Holzinger et Karsten comb. nov. is made.     

黄土丘陵区不同类型生物结皮下的土壤生态化学计量特征

生物结皮在土壤养分累积和循环中起着重要作用.本研究以黄土丘陵区浅色藻结皮、深色藻结皮、藻藓混合结皮、藓结皮、地衣结皮和普通念珠藻结皮6类典型生物结皮为对象,分析不同类型生物结皮土壤碳(C)、氮(N)、磷(P)含量及其化学计量学特征,研究不同类型生物结皮对土壤养分的影响.结果表明:不同类型生物结皮土壤C、N、P、C/N、C/P、N/P差异显著;生物结皮层C、N、P、C/N、C/P、N/P均显著高于0~10 cm土层土壤.6类生物结皮土壤C、N含量均随土层加深而下降,P含量受土层深度影响较小.对于生物结皮层,藓结皮C、N、P含量分别为27.07、2.42、0.67 g·kg^-1,显著高于其他类型生物结皮.念珠藻结皮的0~2 cm土层土壤C、N、P、C/N、C/P、N/P显著高于其他类型生物结皮.     

Warming and increased precipitation frequency on the Colorado Plateau: implications for biological soil crusts and soil processes

Aims

Changes in temperature and precipitation are expected to influence ecosystem processes worldwide. Despite their globally large extent, few studies to date have examined the effects of climate change in desert ecosystems, where biological soil crusts are key nutrient cycling components. The goal of this work was to assess how increased temperature and frequency of summertime precipitation affect the contributions of crust organisms to soil processes.

Methods

With a combination of experimental 2°C warming and altered summer precipitation frequency applied over 2?years, we measured soil nutrient cycling and the structure and function of crust communities.

Results

We saw no change in crust cover, composition, or other measures of crust function in response to 2°C warming and no effects on any measure of soil chemistry. In contrast, crust cover and function responded to increased frequency of summer precipitation, shifting from moss to cyanobacteria-dominated crusts; however, in the short timeframe we measured, there was no accompanying change in soil chemistry. Total bacterial and fungal biomass was also reduced in watered plots, while the activity of two enzymes increased, indicating a functional change in the microbial community.

Conclusions

Taken together, our results highlight the limited effects of warming alone on biological soil crust communities and soil chemistry, but demonstrate the substantially larger effects of altered summertime precipitation.     

生物土壤结皮对全球气候变化的响应

生物土壤结皮在干旱、半干旱区和极地、亚极地区等脆弱生态区广泛存在,生物土壤结皮对脆弱生态系统的稳定、碳氮循环和生态平衡等具有重要的生态意义.概述了脆弱生态区生物结皮的碳氮循环与气候变化关系,综合分析了脆弱生态区生物结皮对气候变暖、降水变化和UV-B增加的响应.分析认为应该加强区域尺度下干旱与半旱区生物结皮对气候变化响应研究.     

古尔班通古特沙漠生物土壤结皮对土壤氮库垂直分布特征的影响

生物土壤结皮在干旱区氮素地球化学循环中具有重要作用,研究不同生物土壤结皮下不同形态氮素含量的变化,解析生物土壤结皮对土壤养分影响过程和范围,有助于进一步理解生物土壤结皮的生态功能。本研究以古尔班通古特沙漠藻-地衣混生结皮和藓类结皮两种生物土壤结皮为研究对象,以裸沙为对照,测定生物土壤结皮层和0—100 cm内8个土层全氮、无机氮、可溶性有机氮、游离态氨基酸氮、微生物生物量氮等氮库含量,和土壤脲酶、硝酸盐还原态酶、亮氨酸氨基肽酶等土壤胞外酶活性。结果表明：1)结皮层各形态氮素含量和各土壤酶活性显著高于其下层土壤,结皮层和结皮下各层土壤氮库整体上表现为藓类结皮>藻-地衣混生结皮>裸沙;土壤氮库各形态氮素含量和土壤酶活性在垂直分布上均呈现先显著下降(0—20 cm)后稳定(20—100 cm)的趋势;在20—30 cm土层,除裸沙的无机氮、铵态氮以及藻-地衣混生结皮的硝态氮外,其余速效氮(无机氮、硝态氮、铵态氮)含量具有增加的特点。2)土壤各氮库含量与全磷、有机碳、电导率、土壤脲酶和亮氨酸氨基肽酶活性呈正相关,与pH、土壤含水率呈负相关。3)利用氮循环相关指标建立土壤氮循环多功能...     

Development of artificially induced biological soil crusts in fields and their effects on top soil

Aims

Biological soil crusts (BSCs) could improve severe environment ecological conditions by increasing soil moisture, soil nitrogen concentration, and so on. In order to control desertification and recover the destroyed soil fertility utilizing a new means using BSCs, the soil surface was artificially inoculated with Microcoleus vaginatus and Scytonema javanicum. Relationships between the development of the artificially induced biological soil crusts and the distribution and dynamic changes of nitrogen and phosphorus in the soil crusts have been analyzed.

Methods

Crusts of different ages were investigated by measuring soil physical and chemical factors, such as moisture, pH, total and available N content, and total and available P, which were correlated with the depths of the crusts.

Results

This study found that the types of color, shape, and species components of the algal crusts increased with crust development. Soil moisture, total N, available N, and available P increased gradually with crust growth. Soil with crusts was wetter than the controlled naked sandy soil, and a significant correlation was observed between biomass and total nitrogen (r?=?0.946, P?=?0.015). Soil pH was lower than that of control. The scytonemin on the soil surface was exceptionally higher than the other pigments, and all the pigments were mainly distributed at the soil surface level. Though the crusts were mainly distributed on soil surface, the available P was mainly stored below the crust layer.

Conclusions

Pearson correlation tests indicated that artificially inoculated biological crusts could improve soil fertility and micro-environment of the top soil: The development of artificially induced BSCs was very well, and this was favorable to inducing the following crust succession.     

The prokaryotic diversity of biological soil crusts in the Sonoran Desert (Organ Pipe Cactus National Monument, AZ)

We studied prokaryotic community structure and composition in biological soil crusts (BSCs) from the Sonoran Desert, and their variability over space and time, using statistically analyzed, PCR-based molecular surveys of environmental 16S rRNA genes. Four sites, tens of km apart, were sampled, 3 times over a 1 year period, collecting 10 duplicate samples every 50 m in each site. Denaturing gradient gel electrophoresis (DGGE) revealed communities much less diverse than those of typical soil assemblages, displaying dominance of some bacterial types. No differences in crust microbial diversity or composition were detected between crusts under plant canopies and those in plant interspaces, indicating a likely crust independence from higher plant resources. However, statistically significant variability with space and time could be detected, and samples within a site were more similar than samples between sites. Both temporal and spatial variability in community composition involved non-dominant members of the community. Extensive sequencing and phylogenetic analysis revealed a large array of bacterial types, many novel. The most common included members of Cyanobacteria, Proteobacteria, Actinobacteria and Acidobacteria. Bacteriodetes, Chloroflexi and Gemmatimonadetes were not seen in high numbers, but were present in all sites, and Deinococci were also detected. Archaea were present, but as minor components. Sonoran BSC communities were distinct in rough compositional character from those in bulk arid soils or agricultural soils, and contained reoccurring, uncultured microbes.     

Activation of methanogenesis in arid biological soil crusts despite the presence of oxygen

Methanogenesis is traditionally thought to occur only in highly reduced, anoxic environments. Wetland and rice field soils are well known sources for atmospheric methane, while aerated soils are considered sinks. Although methanogens have been detected in low numbers in some aerated, and even in desert soils, it remains unclear whether they are active under natural oxic conditions, such as in biological soil crusts (BSCs) of arid regions. To answer this question we carried out a factorial experiment using microcosms under simulated natural conditions. The BSC on top of an arid soil was incubated under moist conditions in all possible combinations of flooding and drainage, light and dark, air and nitrogen headspace. In the light, oxygen was produced by photosynthesis. Methane production was detected in all microcosms, but rates were much lower when oxygen was present. In addition, the δ(13)C of the methane differed between the oxic/oxygenic and anoxic microcosms. While under anoxic conditions methane was mainly produced from acetate, it was almost entirely produced from H(2)/CO(2) under oxic/oxygenic conditions. Only two genera of methanogens were identified in the BSC-Methanosarcina and Methanocella; their abundance and activity in transcribing the mcrA gene (coding for methyl-CoM reductase) was higher under anoxic than oxic/oxygenic conditions, respectively. Both methanogens also actively transcribed the oxygen detoxifying gene catalase. Since methanotrophs were not detectable in the BSC, all the methane produced was released into the atmosphere. Our findings point to a formerly unknown participation of desert soils in the global methane cycle.     

Structure and Reactivity of a Biological Soil Crust from a Xeric Sandy Soil in Central Europe

The investigation was designed to explore the structure, composition, and activity of a biological soil crust on an acidic, sandy soil from a temperate climate. The crust covers several hundreds of square meters on the hilltop of a large terminal moraine. The conjugate alga Zygogonium ericetorum forms the essential matrix for the crust, a dense web of algal filaments with interspersed lichens and mosses. The crust is composed of three layers, with an uppermost layer consisting nearly entirely of a dense algal mat. In lower layers, a parasitic fungus, penetrating the algal cells, is another important component of the crust community. In this soil crust, photosynthetic and respiratory activity is stabilized at low water activities.     

不同类型生物土壤结皮覆盖下风沙土的入渗特征及模拟

干旱荒漠区广泛分布的生物土壤结皮(BSCs)对土壤水分入渗过程有重要影响。以古尔班通古特沙漠南缘的BSCs为研究对象,基于野外采样与室内模拟实验等方法,探究藓类、地衣和藻等3种类型BSCs覆盖下沙土的入渗特征。结果表明:与无结皮覆盖的风沙土对照,3种类型BSCs均显著降低了沙土初渗速率,藓类结皮、地衣结皮、藻结皮覆盖下初渗速率降低幅度依次为36.10%、46.42%、50.39%;藓类结皮、地衣结皮(P0.05)和藻结皮(P0.05)均明显降低了沙土稳渗速率,降低幅度依次为16.50%、33.98%和35.92%;3种类型BSCs均限制了湿润锋在沙土的推进过程,表现为:藓类结皮、地衣结皮、藻结皮的渗漏时间分别为裸沙对照的2.13、3.04和2.98倍;各类型BSCs均减小了沙土累积入渗量,阻碍了沙土水分入渗,与裸沙对照相比,藓类结皮、地衣结皮、藻结皮的1 h累积入渗量分别降低16.10%、28.56%和26.56%。在实验条件下,Kostiakov模型最适用于模拟不同类型BSCs覆盖下土壤水分入渗过程,Horton模型模拟效果次之。     

Relationships between biological soil crusts,bacterial diversity and abundance,and ecosystem functioning: Insights from a semi‐arid Mediterranean environment

Questions: To what degree do biological soil crusts (BSCs), which are regulators of the soil surface boundary, influence associated microbial communities? Are these associations important to ecosystem functioning in a Mediterranean semi‐arid environment? Location: Gypsum outcrops near Belmonte del Tajo, Central Spain. Methods: We sampled a total of 45 (50 cm × 50 cm) plots, where we estimated the cover of every lichen and BSC‐forming lichen species. We also collected soil samples to estimate bacterial species richness and abundance, and to assess different surrogates of ecosystem functioning. We used path analysis to evaluate the relationships between the richness/abundance of above‐ and below‐ground species and ecosystem functioning. Results: We found that the greatest direct effect upon the ecosystem function matrix was that of the biological soil crust (BSC) richness matrix. A few bacterial species were sensitive to the lichen community, with a disproportionate effect of Collema crispum and Toninia sedifolia compared to their low abundance and frequency. The lichens Fulgensia subbracteata and Toninia spp. also had negative effects on bacteria, while Diploschistes diacapsis consistently affected sensitive bacteria, sometimes positively. Despite these results, very few of the BSC effects on ecosystem function could be ascribed to changes within the bacterial community. Conclusion: Our results suggest the primary importance of the richness of BSC‐forming lichens as drivers of small‐scale changes in ecosystem functioning. This study provides valuable insights on semi‐arid ecosystems where plant cover is spatially discontinuous and ecosystem function in plant interspaces is regulated largely by BSCs.     

外源输入氮素在藓类生物土壤结皮中各氮组分的分配特征与归趋途径

针对黄土高原典型藓类生物结皮,以无结皮裸地为对照,通过野外原位同位素标记试验,在标记后1~30 d连续取样测定,示踪外源添加¹⁵N在生物结皮中各氮组分的分配特征,并分析¹⁵N在土壤-微生物-藓植株中的归趋途径,对比揭示生物结皮对土壤氮循环的影响。结果表明: 1)生物结皮的全氮、微生物生物量氮、可溶性有机氮中¹⁵N含量分别平均比裸地高2.9、17.5、9.0倍,且藓植株固定的¹⁵N含量高达4.73 mg·kg^-1。2)生物结皮的¹⁵N残留率平均为13.0%,其氮固持能力是裸地(3.3%)的4.0倍;生物结皮中各组分¹⁵N占全氮的比率依次为微生物生物量氮(54.3%)>藓植株氮(22.5%)>可溶性有机氮(6.2%),而裸地则为微生物生物量氮(11.5%)>可溶性有机氮(2.6%),显示生物结皮中微生物和藓植株的氮固持能力合计比裸地高65.3%。3)生物结皮中微生物生物量¹⁵N的转移量和库容量分别比裸地高17.2和20.5倍,但其周转率为每月5.8次,低于裸地的每月7.2次,其周转期是裸地的1.2倍。综上,与无结皮裸地相比,生物结皮具有更高的氮固持能力,同时能够改变土壤各氮组分的分配率,因此,在干旱生态系统土壤氮循环过程中具有重要作用。     

荒漠地表生物土壤结皮形成与演替特征概述

土壤表面结皮是世界范围内干旱沙漠地区土壤表面广泛存在的自然现象,包括物理结皮和生物土壤结皮两大类型。其中,生物土壤结皮作为干旱沙漠地区特殊环境的产物,是由细菌、真菌、蓝绿藻、地衣和苔藓植物与土壤形成的有机复合体。它的形成使土壤表面在物理、化学和生物学特性上均明显不同于松散沙土,具有较强的抗风蚀功能和重要的生态效应,成为干旱沙漠地区植被演替的重要基础。随着形成生物土壤结皮的物种更替,维持结皮结构的主要胶结方式亦随之发生变化,即由胞外多糖的粘结作用逐渐转变为蓝藻和荒漠藻的藻丝体、地衣菌丝体以及苔藓植物假根的缠绕和捆绑作用,物种更替是结皮微结构和胶结方式转化的生物基础。生物土壤结皮的形成通常可以分为以下几个阶段:生物土壤结皮的早期阶段(土壤酶和土壤微生物),藻结皮阶段、地衣结皮阶段和苔藓结皮阶段。即随着土壤微生物在沙土表面的生长,随后出现丝状蓝藻和荒漠藻类植物,形成以藻类植物为主体的荒漠藻结皮;当土壤表面得到一定固定后,便开始出现地衣和苔藓植物,形成以地衣和苔藓植物为优势的生物结皮类型。其中,前一阶段的完成又为后一阶段的开始提供良好的环境条件。当环境条件适宜时,生物土壤结皮也可以不经历其中某个阶段而直接发育到更高级的阶段。     

古尔班通古特沙漠生物土壤结皮下土壤有机碳垂直分布特征及影响因素

为了探究沙漠不同生物土壤结皮类型覆盖下土壤有机碳(SOC)垂直分布特征及与土壤理化因素的关系,解析影响因素,以古尔班通古特沙漠腹地藻类、藓类生物土壤结皮和裸沙三种不同地被覆盖类型为研究对象,通过野外采集不同类型结皮样品及其下层0-2 cm、2-5 cm、5-10 cm、10-20 cm、20 -30 cm、30-50 cm、50-70 cm以及70-100 cm土层土壤(裸沙对照),测定不同土层的SOC含量及土壤理化指标,开展相关研究。结果表明:(1)不同地被类型下0-100 cm SOC含量随着土壤深度增加呈减小的趋势,在10-30 cm 土层存在SOC含量升高的现象,藓类、藻类和裸沙三种地被类型0-100 cm土层SOC含量范围分别在:1.61-2.70、1.41-2.56、1.21-1.92 g/kg;(2)不同地被类型下同一土层SOC含量在0-5 cm土层存在显著差异,5-100 cm土层SOC整体无显著差异,同层SOC含量均表现为:藓类>藻类>裸沙对照;(3)Pearson法分析结果表明不同地被覆盖下SOC含量与养分(全氮、全磷)呈现正相关关系,与pH和电导率(EC)呈现负相关关系。通过结构方程模型结果表明,土壤养分和粒径(砂粒占比)是影响垂直分布的主要因子,其中粒径是裸沙和藻类的最主要影响因子,藓类最重要的影响因子是养分(全磷)。生物土壤结皮的发育会逐步提高土壤碳的积累,改变SOC的垂直分布特征,对SOC的影响主要集中在5 cm以上土层,土壤理化特征对垂直分布特征具有调控作用。