Microbiotic Crusts and Ecosystem Processes

Microbiotic crusts are biological soil crusts composed of lichens, cyanobacteria, algae, mosses, and fungi. The biodiversity of these crusts is poorly understood; several cosmopolitan species dominate in most areas, but many species are confined to one or a few sites. Nitrogen fixation by organisms within the crust can be the dominant source of nitrogen input into many ecosystems, although rates of nitrogen input are limited by water availability, temperature, and nitrogen loss from the crust. Photosynthetic rates of the microbiotic crust can be 50% of those observed for higher plants, but the contribution of crusts to carbon cycling is not known. The microbiotic crust binds soil particles together, and this significantly increases soil surface stability and resistance to erosion. Greenhouse studies have found that crusts can enhance seed germination, seedling survivorship, and plant nutrient status, but further experiments are needed under field conditions. Crusts are extremely susceptible to surface disturbance and fire, and disruption of crusts can decrease soil fertility and stability resulting in lower nutrient availability for vascular plants and significant soil loss from the ecosystem.     

Effects of Biological Soil Crusts on Seed Bank, Germination and Establishment of Two Annual Plant Species in the Tengger Desert (N China)

The presence of biological soil crusts can affect the germination and survival of vascular plants, but the reasons are not well investigated. We have conducted a field investigation and greenhouse experiments to test the effect of crusts on two desert annual plants, which occur on the stabilized dunes of the Tengger Desert in N China. The results showed that biological soil crusts negatively influenced the seed bank of Eragrostis poaeoides and Bassia dasyphylla. The important effect of biological soil crusts on seed germination and establishment were performed indirectly through reducing the amount of germinating seeds. Field investigation and experimental results with regard to the seed bank indicated that higher seedling density was found in disturbed crust soil and bare soil surface than in intact crust soils. Greenhouse experiments showed that the effects of biological soil crusts on germination and establishment of the two plants are not obvious in moist condition, while disturbed crusts are more favorable to seed germination in dry treatment. Significant differences in biomass were found between disturbed crust soil and bare soil. Survival and growth of the two annual plants were enhanced in both algal and moss crusts during the season of rainfall or in moist environment, but crusts did not affect seedling survival in the dry period. The small seeded E. poaeoides has higher germination than larger-seeded B. dasyphylla in crust soils, but B. dasyphylla has a relatively higher survival rate than E. poaeoides in crust soils.     

Germination and seed water status of four grasses on moss-dominated biological soil crusts from arid lands

Biological soil crusts dominated by drought-tolerant mosses are commonly found through arid and semiarid steppe communities of the northern Great Basin of North America. We conducted growth chamber experiments to investigate the effects of these crusts on the germination of four grasses: Festuca idahoensis, Festuca ovina, Elymus wawawaiensis and Bromus tectorum. For each of these species, we recorded germination time courses on bare soil and two types of biological soil crusts; one composed predominantly of the tall moss Tortula ruralis and the other dominated by the short moss Bryum argenteum. On the short-moss crust, the final germination percentage was about half of that on bare soil. Also, the mean germination time was 4 days longer on short-mosses than on bare soil. In contrast to the short-moss crust, the tall-moss crust did not reduce the final germination percentage but increased the mean germination time. Similar results were observed in the four grasses studied. To investigate the mechanism by which moss crusts affected germination, we analyzed the water status of seeds on bare soil and moss crusts. Six days after seeding, the water content of seeds on bare soil was approximately twice that of seeds on tall- or short-moss crust. Analysis of the time course of changes in seed weight and water potential in Bromus tectorum revealed that overtime seeds on tall mosses reached higher water content than those on short mosses. The increase in the water content of seeds on tall mosses occurred as the seeds gradually fell through the moss canopy. Taken together, our results indicate that biological soil crusts with distinct structural characteristics can have different effects on seed germination. Furthermore, this study revealed that a biological soil crust dominated by short mosses had a negative effect on seed water status and significantly reduced seed germination.     

生物土壤结皮对荒漠土壤种子库和种子萌发的影响

研究了腾格里沙漠东南缘在不同自然条件（风、温度、水分）下,人工固沙植被区（24龄、41龄、50龄）和相邻天然植被区的两种生物土壤结皮对荒漠土壤种子库和种子萌发的影响。结果表明,荒漠土壤种子库在苔藓结皮上的储量显著高于藻类结皮。随着生物土壤结皮的发育,种子萌发量在苔藓结皮上增加,在藻类结皮上减少。生物土壤结皮层的含水量对种子萌发有显著的影响（p〈0.05）,植物种子在湿润处理的生物土壤结皮上的萌发量高于干燥处理的生物土壤结皮上的种子萌发量。生物土壤结皮表层温度和亚表层温度对荒漠植物种子萌发无显著影响（p〉0.05）,但总体而言,对于苔藓结皮,植物种子在较高温度下的萌发量略高于在较低温度下的萌发量,而对于藻类结皮,植物种子在较低温度下的萌发量略高。     

实验室条件下两种生物土壤结皮对荒漠植物种子萌发的影响

生物土壤结皮广泛分布在干旱、半干旱区,深刻影响着土壤表层特性,进而对植物种子散布、萌发和定居产生极大的影响。到目前为止,生物土壤结皮与植物关系的研究很少见到,并且这些有限的研究所得出的结论存在着争议。研究了不同年龄的两种生物土壤结皮（苔藓结皮和藻类结皮）对油蒿（Artemisia ordosica）和雾冰藜（Bassia dasyphylla）种子萌发的影响,同时也研究了这两种结皮在失去活性前后对油蒿、雾冰藜和小画眉草（Eragrostis poaeoides）种子萌发的影响。苔藓和藻类结皮的出现对油蒿和雾冰藜种子的萌发均有显著的促进作用,而结皮年龄对植物种子的萌发没有显著的差异。对于不同的植物种,结皮类型和活性对种子的萌发具有不同的作用。雾冰藜在两种结皮上的萌发有显著的差异而油蒿和小画眉草在两种结皮上的萌发没有显著差异。活藻类显著地增加了三种植物的种子萌发,活苔藓仅增加了油蒿和雾冰藜种子萌发量而对小画眉草种子的萌发没有作用,研究表明,生物土壤结皮对一些植物种子萌发具有明显的促进作用。     

Seed characteristics and soil surface patch type interact to affect germination of semi-arid woodland species

Biological soil crusts are common in many arid and semi-arid regions and they can alter microenvironments which are likely to directly and indirectly influence vascular plant establishment. The effect of biological soil crusts on germination is also influenced by the biological characteristics of the seeds themselves but rarely have the effects of both crust type and seed morphology on germination been examined in the same study. In this study, seed of five semi-arid woodland species with contrasting seed morphology were sown on top of patch types that commonly occur in natural woodlands (foliose lichen, short-turf moss, tree leaf litter, disturbed crust) and their emergence was followed. Percent germination varied between patch types and, for the largest-seeded species (Maireana excavata), final germination was significantly lower on the biological soil crust and litter patch types because they strongly acted as a physical barrier to seed penetration into the soil. The remaining four species showed no significant differences in final percent germination with patch type because most seeds either completely or partially penetrated the surface layer. Germination time courses, however, showed that biological soil crusts delayed the timing of germination of these species. Hence, soil crusts might differentially affect the spatial patterning of species in semi-arid woodlands by their subtle influence on seedling emergence that is determined by differences in seed morphology and subsequent positioning within crusts.     

Effects of biological soil crusts on emergence of desert vascular plants in North China

Biological soil crusts are a universal and common feature in arid and semi-arid regions and their appearance profoundly affects soil surface properties which may greatly change the seed dispersal, germination and establishment. To date, only a handful of experiments have exerted to investigate the effects of crusts on vascular plants and the conclusions from these studies are variable. In this study, we investigate the influences of two different crusts universally spreading in southeastern part of the Tengger Desert with four chronosequences (24, 41, 50 years old in artificial vegetation area and natural vegetation crusts) on vascular plants. Crusts were placed at three different sites to simulate different environmental factors (wind velocity and soil crust moisture), we set two soil moisture regimes for crusts to investigate how vascular plants respond under different moisture regimes in crusts. Emergence densities of vascular plants were significantly higher in moss crust than in algae crust. With the development of crusts, seed emergence increased in moss crust while decreased in algae crust. As for effects of moisture, our results showed that soil moisture had a significant effect on seed emergence in both types of crusts at all developing phases. Crusts with higher moisture had more seedlings than those with lower moisture. The above results indicated that the appearance of crusts changed the surface soil properties, which had greatly influenced the entrapment and lodgement of seeds in the study area, thus subsequently influence seed emergence through affecting natural factors.     

Initial effects of experimental warming on carbon exchange rates, plant growth and microbial dynamics of a lichen-rich dwarf shrub tundra in Siberia

Biological soil crusts can affect seed germination and seedling establishment. We have investigated the effect of biological soil crusts on seed water status as a potential mechanism affecting seed germination. The seed water potential of two annual grasses, one exotic Bromus tectorum L. and another native Vulpia microstachys Nutt., were analyzed after placing the seeds on bare soil, on a crust that contains various lichens and mosses (mixed crust), or on a crust dominated by the crustose lichen Diploschistes muscorum (Scop.) R. Sant. (Diploschistes crust). Seed water potential and germination were similar on the bare soil and the mixed crust, except for the initial germination of V. microstachys, which was higher on the mixed crust than on the bare soil. For the two grasses studied, seed water potential was significantly higher on the bare soil and mixed crust than on the Diploschistes crust. These differences in water potential correlated with differences in germination, which was much lower on the lichen crust. Experiments were conducted under two watering regimens. Increasing the frequency of watering amplified the differences in seed water potential and germination between the Diploschistes crust and the other two surfaces. For a particular watering regimen, the bare soil, mixed crust, and Diploschistes crust received the same amount of water, but they reached significantly different water potentials. Throughout the experiments, the water potential of the soil and mixed crust remained above −0.6 MPa, while there was a marked decline in the water potential of the Diploschistes surface to about −4 MPa. To ascertain that water was the major factor limiting germination on the Diploschistes crust, we conducted germination tests in an environment with 100% relative humidity. Under these conditions, germination on the Diploschistes crust was similar to that on the bare soil. However, the seeds that germinated on the Diploschistes crust did not penetrate this surface and approximately 60% of their root tips became necrotic. Our results indicate that the presence of D. muscorum can inhibit seedling establishment by two mechanisms: a reduction in seed water absorption and an increase in root tip mortality.     

Germination and seedling establishment of two annual grasses on lichen-dominated biological soil crusts

Biological soil crusts dominated by lichens are common components of shrub-steppe ecosystems in northwestern US. We conducted growth chamber experiments to investigate the effects of these crusts on seed germination and initial seedling establishment of two annual grasses; the highly invasive exotic Bromus tectorum L. and the native Vulpia microstachys Nutt. We recorded germination time courses on bare soil and two types of biological soil crusts; one composed predominantly of the lichen Diploschistes muscorum (Scop.) R. Sant. (lichen crust) and the other comprised of an assortment of lichens and mosses (mixed crust). Final germination on the lichen crust for both grass species was about a third of that on the bare soil surface. Mean germination time (MGT) was 3–4 days longer on the lichen crust compared with the bare soil. In contrast, there was no difference in germination percentage or MGT between the mixed crust and bare soil, and results were similar for both grass species. For both species, root penetration of germinating seeds on the lichen crust was lower than on the bare soil or mixed crust surfaces. The combined effects of the lichen crust on germination and root penetration resulted in an overall reduction in seedling establishment of 78% for V. microstachys and 85% for B. tectorum relative to the bare soil treatment. Our results clearly demonstrate that lichen-dominated biological soil crust can inhibit germination and root penetration, but the extent of these effects depends on the composition of the crust. Responsible Editor: Tibor Kalapos     

Can atmospheric input of nitrogen affect seed bank dynamics in habitats of conservation interest? The case of dune slacks

Questions: Does the increased atmospheric deposition of nitrogen, which can have major effects on the established vegetation of nutrient‐poor habitats, also impact germination from the soil seed bank? Location: Coastal dune slacks at Newborough Warren, Wales, UK. Methods: The effects of nitrogen addition (15 kg.ha^‐1.a^‐1) on seed germination from the soil seed bank were investigated using the seedling emergence method between September 2004 and February 2005. Results: More seedlings emerged from fertilised samples than unfertilised controls. Most species showed enhanced germination after fertilisation with nitrogen, with seedling numbers statistically significantly greater in nitrogen addition samples in a quarter of species abundant enough for analysis. Species that responded positively to fertilisation were species with low Ellenberg indicator values indicative of infertile sites. Conclusions: Most species showed increased germination after fertilisation with nitrogen, including early successional species normally growing in nutrient‐poor conditions. This suggests that the increased atmospheric deposition of nitrogen probably not only impacts on established vegetation, but also has the potential to alter seed bank dynamics.     

生物结皮对5种不同形态的荒漠植物种子萌发的影响

生物结皮广泛分布于干旱、半干旱区, 强烈影响着土壤表层理化特性, 进而对种子散布、萌发和定居产生影响。目前关于生物结皮与植物种子萌发关系的研究结论存在争议。该文通过室内人工控制实验, 研究了生物结皮对古尔班通古特沙漠5种具不同种子形态特征的荒漠植物白梭梭(Haloxylon persicum)、蛇麻黄(Ephedra distachya)、角果藜(Ceratocarpus arenaarius)、涩芥(Malcolmia africana)和狭果鹤虱(Lappula semiglabra)的种子萌发的影响。结果表明, 在干燥和湿润两种条件下, 生物结皮对不同形态植物种子萌发均具有不同的作用。在干燥条件下, 生物结皮显著抑制了角果藜和涩芥种子的萌发(p<0.05), 对其它3种植物无显著影响; 而湿润条件下, 生物结皮显著抑制了白梭梭、角果藜和狭果鹤虱种子的萌发(p<0.05), 对蛇麻黄、涩芥则无显著影响。     

Effects of biological soil crusts on seed germination of four endangered herbs in a xeric Florida shrubland during drought

Soil crusts of rosemary scrubs in south-central Florida were examined for effects on seed germination of four herbs that are killed by fire and must recruit from seed: Eryngium cuneifolium (Apiaceae), Hypericum cumulicola (Hypericaceae), Polygonella basiramia (Polygonaceae), and Paronychia chartacea ssp. chartacea (Caryophyllaceae). Biological soil crusts in these sites are dominated by algae, cyanobacteria, fungi, and bacteria. Because crusts can change soil stability, water, and nutrients, they can affect seed germination. A series of greenhouse and field experiments were designed to first examine the effects of crusts in isolation and then to determine their role in the context of other environmental factors – time since fire and distance to the dominant shrub in this system, Ceratiola ericoides. In the greenhouse experiment, germination in autoclaved crusts was dramatically reduced relative to germination in living crusts for all but P. basiramia. In four field experiments where crusts were left intact, disturbed (mechanically or by flaming), or completely removed, the effects of crusts were variable and species-specific, but were significant enough to impact aboveground population sizes. More germination was consistently observed in recently burned sites away from C. ericoides shrubs. Overall rates of germination were generally very low during this study, possibly as a result of seasonal droughts that could have reduced germination, increased seed dormancy, and/or decreased seed viability. The importance of water for germination was confirmed in an experiment with two watering regimes and three crust treatments designed to create a gradient of soil water availability. Germination was significantly greater in the high water treatment and unaffected by soil crust moisture. Dry years are not uncommon in scrub and the results of this study help us to understand how scrub herbs fare during drought and what role biological soil crusts play in germination.     

Crust Composition and Disturbance Drive Infiltration Through Biological Soil Crusts in Semiarid Ecosystems

Soil crusts influence many soil parameters that affect how water moves into and through the soil, and therefore, critically influence water availability, erosion processes, nutrient fluxes, and vegetation distribution patterns in semiarid ecosystems. Soil crusts are quite sensitive to disturbance, and their alteration can lead to modification of the local hydrological regime, thus affecting general functioning of the ecosystem. The aim of this study was to analyze the influence of different types of soil crusts, physical, and biological in different developmental stages, as well as the impact of their disturbance, on infiltration. This was assessed by means of rainfall simulations conducted in two semiarid ecosystems in southeast Spain characterized by different lithologies, topographies, and soil crust distributions. Two consecutive rainfall simulation experiments (50 mm h⁻¹ rainfall intensity), the first on dry soil and the second on wet soil, were carried out in microplots (0.25 m²) containing the most representative soil crust types at each site, each crust type subjected to three disturbance treatments: (a) undisturbed, (b) trampling, and (c) removal. Infiltration in the crusts was higher on coarse- than on fine-textured soils and almost two times greater on dry than on wet soil. Biological soil crusts (BSC) showed higher infiltration rates than physical soil crusts (PSC). Within BSC, infiltration increased as cyanobacterial biomass increased and was the highest in moss crusts. However, late-successional crustose and squamulose lichen crusts showed very low infiltration rates. Trampling reduced infiltration rates, especially when soil was wet, whereas crust removal enhanced infiltration. But this increase in infiltration after removing the crust decreased over time as the soil sealed again due to raindrop impact, making runoff rates in the scraped microplots approach those registered in the respective undisturbed crust types. Our results demonstrate that water redistribution in semiarid ecosystems strongly depends on the type of crusts that occupy the interplant spaces and the characteristics of the soils which they overly, as well as the antecedent moisture conditions of the soil. Disturbance of these crust patches results in increased runoff and erosion, which has important consequences on general ecosystem functioning.     

生物土壤结皮——荒漠化地区研究的热点问题

生物土壤结皮在荒漠化地区广为分布。从结皮的概念、结皮对生态系统和景观变化的影响（包括水文循环、土壤侵蚀、土壤养分循环、维管植物的萌发和生长、动物多样性、景观异质性以及生态系统和景观变化监测和评价等）以及生物土壤结皮对干扰的反应及其恢复机理等方面详细地论述了生物结皮在荒漠化发生发展及其防治中所起的重要作用,并提出了今后生物土壤结皮研究的方向和着眼点。     

石漠化地区苔藓结皮对土壤养分及生态化学计量特征的影响

苔藓结皮是石漠化生态系统的重要地表覆被物,但其在土壤养分累积和元素循环过程中的作用尚不明确。以我国贵州典型喀斯特高原峡谷石漠化区-花江大峡谷两岸不同等级石漠化生境下的苔藓结皮及其覆被土壤为研究对象,研究了苔藓结皮覆被对土壤养分及生态化学计量特征的影响。结果表明：（1）苔藓结皮层养分含量显著高于下层土壤,结皮覆被土壤有机碳（SOC）、全氮（TN）、全磷（TP）、全钾（TK）、碱解氮（AN）、速效磷（AP）、速效钾（AK）平均含量分别为25.95、3.05、1.00、5.10 g/kg和189.61、1.59、275.10 mg/kg,较无结皮覆被的裸土分别增加46.08%、26.50%、53.62%、20.25%、25.24%、110.47%和83.76%。（2）苔藓结皮覆被土壤C/N、C/K、P/K显著高于裸土,且随土层加深而递减;N/P显著低于裸土,且随土层加深而升高。（3）苔藓结皮覆被土壤养分恢复指数为33.16%-72.48%,呈现随石漠化等级升高而增加的趋势,中度和强度石漠化阶段较无石漠化阶段分别增加83.26%和118.58。本研究表明苔藓结皮能有效促进土壤养分累积,加速石漠化土壤养分恢复进程,可作为补充手段联合其它生态恢复措施共同推动石漠化地区的生态恢复与重建。     

陇中黄土高原典型草原生物土壤结皮发育对土壤养分的影响

为了解生物土壤结皮发育与土壤养分特征的关系,该研究选取处于不同发育阶段和不同盖度的藓结皮(M-crust)和地衣结皮(L-crust),对其结皮层生物体及结皮下层3 cm土壤进行养分分析。结果显示：(1)藓结皮生物体对土壤全氮(TN)、全磷(TP)、铜(Cu)和锌(Zn)的生物富集作用较强,其富集系数分别高达807.4%、175%、122.4%和244.5%,而地衣结皮生物体对TN、TP和钙(Ca)的生物富集作用较强,其富集系数分别高达950.8%、126.2%和208.6%;除Ca外,藓结皮生物体中TP、钠(Na)、钾(K)、Cu、铁(Fe)、镁(Mg)、锰(Mn)和Zn均显著高于地衣结皮。(2)土壤生物结皮盖度与其对应的土壤养分含量之间无显著线性相关关系,但发育有不同盖度藓结皮的土壤其平均TN、TP、Cu、Fe、K、Na和Ca含量均高于地衣结皮。研究表明,不同类型生物土壤结皮对养分吸收、富集效率不同,处于发育后期的藓结皮较发育中期的地衣结皮具有更强的养分累积效应,提高了土壤的养分贮存量,对黄土高原典型草原生态系统的恢复与重建具有潜在的重要影响。     

黄土高原水蚀风蚀交错区藓结皮对土壤酶活性的影响

以黄土高原水蚀风蚀交错区黄绵土和风沙土上发育30年的典型藓结皮为对象,分6层采集0～12 cm土层土样,研究了藓结皮对不同土层4种水解酶活性的影响及其与土壤养分的关系.结果表明: 与无结皮相比,黄绵土藓结皮的脲酶、碱性磷酸酶、蔗糖酶和蛋白酶活性分别提升了2.4、7.6、20.7、2.4倍,而风沙土藓结皮的这4种水解酶活性分别提升了3.5、22.2、22.3、2.0倍;黄绵土和风沙土藓结皮的体积含水量分别降低了6.5%和0.8%,温度分别降低了0.8和2.5 ℃;黄绵土藓结皮的有机质、碱解氮和速效磷含量分别提升了2.5、2.9和3.6倍,风沙土分别提升了3.6、3.0和6.6倍.4种水解酶的活性与土壤养分含量呈显著正相关,与土壤含水量呈显著负相关;而土壤温度与脲酶活性呈显著正相关,与蛋白酶活性呈显著负相关.黄土高原水蚀风蚀交错区2种土壤藓结皮均能显著提升水解酶活性,这是藓结皮加速土壤养分周转的重要因素;通过改变土壤含水量和温度等酶促反应条件间接影响酶活性,是藓结皮促进土壤酶活性的重要途径.     

The germination of mucilaginous seeds of t Plantago coronopus,Reboudia pinnata,and t Carrichtera annua on cyanobacterial soil crust from the Negev Desert

We studied the effect of intact, crushed or sterile cyanobacterial soil crust from the Negev Desert highlands of Israel as substrates for the germination of seeds of three annual plant species from local populations that produce mucilaginous seeds t Plantago coronopus, Reboudia pinnata and t Carrichtera annua. Mucilaginous seeds of these species were wetted on local intact cyanobacterial soil crust which inhibited their germination in comparison with their germination on filter paper. However, the percentage of germination of each species differed on sterile and crushed soil crust after 72 h of wetting. Germination of t Plantago coronopus seeds was inhibited mechanically and biologically on intact soil crust, since it was significantly higher on both crushed and sterile soil crust. Germination of t Reboudia pinnata seeds was slightly mechanically and biologically inhibited on intact soil crust, but germination on live crushed crust was higher than on sterile crushed crust. By contrast, germination of t Carrichtera annua seeds was not mechanically inhibited by live soil crust but there was significant biological acceleration of germination on intact soil crust relative to crushed soil crust. Each of the three species exhibited different requirements for germination. Germination of t P. coronopus is promoted by disturbances and removing the crust components (including its biological and structural effects). t R. pinnata and t C. annua germination is less affected by crust disturbance but is generally inhibited after removing the live components of the cyanobacterial crust.These species differences suggest different functions of the mucilage of the seed coat for the three species.     

温带荒漠灌丛和藓类结皮对土壤养分的贡献

灌丛与生物土壤结皮镶嵌分布是温带荒漠常见的地表景观之一,二者的发育均显著影响了地表土壤养分的空间分布特征及循环过程。然而,灌丛和生物土壤结皮对荒漠土壤表层养分的影响存在怎样的差异,二者对养分变化的贡献度如何尚不清楚。因此,选择中国北方典型温带荒漠古尔班通古特沙漠为研究区,以荒漠中的优势灌丛膜果麻黄(Ephedra przewalskii)灌丛和生物土壤结皮发育高级阶段的藓类结皮的结皮层及结皮下层土壤为研究对象,采集不同微生境(裸露地、灌丛下)的裸沙与藓类结皮的土壤样品。为探究不同微生境下不同土层碳、氮、磷和钾养分变化特征,测定了不同土层土壤有机碳(SOC)、全氮(TN)、全磷(TP)、全钾(TK)、速效氮(AN)、速效磷(AP)和速效钾(AK)含量。结果表明:(1)相较于裸露地,灌丛显著提高了藓类结皮不同土层有机碳、全氮和全钾的含量和裸沙全磷的含量,降低了灌丛下藓类结皮土壤全磷的含量。(2)对于速效养分而言,与裸沙相比,裸露地藓类结皮覆盖降低了土壤速效氮含量,增加了速效磷和速效钾含量;而灌丛下藓类结皮覆盖提高了土壤速效氮和速效钾的含量,但降低了速效磷的含量。 (3) 相关性分析显示,在0-2 cm土壤中速效磷与速效氮呈现显著负相关,而在2-6 cm速效磷与速效氮呈现显著正相关(P<0.01)。(4)贡献度分析表明,土壤中灌丛效应对养分的贡献(42.54%)要远大于藓类结皮的贡献(2.43%),但二者交互作用却降低了除速效氮以外的其他土壤养分含量。综上,灌丛、藓类结皮覆盖和土层深度变化对土壤碳、氮、磷和钾养分均具有显著影响(P<0.05),但三者间的交互效应对养分的影响不显著(P>0.05)。相对于裸露地,灌丛与藓类结皮的覆盖均对表层土壤碳、氮、磷和钾养分具有促进作用,且随着土层深度的下降,土壤养分含量呈现显著的下降趋势(P<0.05)。因此,在荒漠生态系统中耐旱灌丛与生物土壤结皮这两个最重要的斑块单元联合调控了微尺度土壤养分的空间异质性变化。     

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.