Biological soil crusts in ecological restoration: emerging research and perspectives

Drylands encompass over 40% of terrestrial ecosystems and face significant anthropogenic degradation causing a loss of ecosystem integrity, services, and deterioration of social‐ecological systems. To combat this degradation, some dryland restoration efforts have focused on the use of biological soil crusts (biocrusts): complex communities of cyanobacteria, algae, lichens, bryophytes, and other organisms living in association with the top millimeters of soil. Biocrusts are common in many ecosystems and especially drylands. They perform a suite of ecosystem functions: stabilizing soil surfaces to prevent erosion, contributing carbon through photosynthesis, fixing nitrogen, and mediating the hydrological cycle in drylands. Biocrusts have emerged as a potential tool in restoration; developing methods to implement effective biocrust restoration has the potential to return many ecosystem functions and services. Although culture‐based approaches have allowed researchers to learn about the biology, physiology, and cultivation of biocrusts, transferring this knowledge to field implementation has been more challenging. A large amount of research has amassed to improve our understanding of biocrust restoration, leaving us at an opportune time to learn from one another and to join approaches for maximum efficacy. The articles in this special issue improve the state of our current knowledge in biocrust restoration, highlighting efforts to effectively restore biocrusts through a variety of different ecosystems, across scales and utilizing a variety of lab and field methods. This collective work provides a useful resource for the scientific community as well as land managers.     

Practices of biological soil crust rehabilitation in China: experiences and challenges

Biological soil crusts (biocrusts) are a central component of dryland ecosystems. However, they are highly vulnerable to disturbance and natural recovery may be slow. Therefore, finding ways to enhance the reestablishment of biocrusts after disturbance has been of great interest to researchers. This article provides a review of the laboratory cultivation and field inoculations of biocrust materials in China (mostly published in Chinese). Larger filamentous cyanobacteria (e.g. Microcoleus) are relatively easy, although slow, to grow in culture compared to other biocrust components. Thus, most researchers have focused their efforts on the cyanobacteria and a few species of mosses that are also easily grown but at smaller scale. For all the studies, a small amount of biocrust material was collected and its biomass enhanced under controlled conditions. However, the enhancement was done using various methods and techniques in different regions. These materials were then applied to disturbed field sites, again with various methods. Results show that keeping the inoculated soil surface wet for some time period after inoculation was crucial for restoration success. Cyanobacterial establishment was improved by installing automatic sprinkling using micro‐irrigation techniques and/or physical structures that reduced sediment moving onto the inoculated area. Experimental applications in China showed that cyanobacteria can be successfully inoculated at a large scale (hundreds of ha). Moss inoculation, on the other hand, was only accomplished at a small scale (several m²). To assess whether biocrust restoration can enhance the establishment of a self‐supporting ecosystem, further research is needed on how inoculation affects vegetation diversity and structure and ecological processes.     

Microsite enhancements for soil stabilization and rapid biocrust colonization in degraded drylands

In dryland ecosystems, natural recovery of biological soil crusts (biocrusts) following disturbance may be slow or inhibited, necessitating active restoration practices. While biocrusts can be readily propagated under environmentally controlled conditions, rehabilitation in the field is complicated by environmental stresses which may be particularly acute in degraded, destabilized soils with harsh climatic conditions at the soil surface. In this study, we first present the results of a field trial at a severely degraded rangeland site examining the stabilizing effects of various soil amendments (polysaccharide glues and polyacrylamides) in combination with biocrust inoculum. We found that a psyllium compound was the only amendment to maintain effectiveness after 19 months, and the only treatment that maintained biocrust inoculum throughout the trial. In a subsequent short‐term experiment where plots were shaded and watered, we examined how biocrust inoculation rate (0, 20, and 40% initial cover) and the psyllium‐based amendment affected biocrust growth. After 4 months, visible biocrust cover in inoculated plots was greater than in controls, but only chlorophyll a exhibited a dosage‐response to inoculum application rate, indicating preferential establishment of cyanobacteria. Psyllium did not affect biocrust development but did improve soil stability. Shade and watering buffered against temperature extremes (up to 15°C) and increased the duration of moist surface conditions necessary for biocrust growth by up to 30%, mimicking conditions more common in the fall and winter months. Our results suggest that inducing early successional biocrusts on a highly degraded site is possible with suitable microclimate conditions.     

Climate Change and Physical Disturbance Manipulations Result in Distinct Biological Soil Crust Communities

Biological soil crusts (biocrusts) colonize plant interspaces in many drylands and are critical to soil nutrient cycling. Multiple climate change and land use factors have been shown to detrimentally impact biocrusts on a macroscopic (i.e., visual) scale. However, the impact of these perturbations on the bacterial components of the biocrusts remains poorly understood. We employed multiple long-term field experiments to assess the impacts of chronic physical (foot trampling) and climatic changes (2°C soil warming, altered summer precipitation [wetting], and combined warming and wetting) on biocrust bacterial biomass, composition, and metabolic profile. The biocrust bacterial communities adopted distinct states based on the mechanism of disturbance. Chronic trampling decreased biomass and caused small community compositional changes. Soil warming had little effect on biocrust biomass or composition, while wetting resulted in an increase in the cyanobacterial biomass and altered bacterial composition. Warming combined with wetting dramatically altered bacterial composition and decreased Cyanobacteria abundance. Shotgun metagenomic sequencing identified four functional gene categories that differed in relative abundance among the manipulations, suggesting that climate and land use changes affected soil bacterial functional potential. This study illustrates that different types of biocrust disturbance damage biocrusts in macroscopically similar ways, but they differentially impact the resident soil bacterial communities, and the communities'' functional profiles can differ depending on the disturbance type. Therefore, the nature of the perturbation and the microbial response are important considerations for management and restoration of drylands.     

Do soil inoculants accelerate dryland restoration? A simultaneous assessment of biocrusts and mycorrhizal fungi

Biological soil crusts (biocrusts) and arbuscular mycorrhizal (AM) fungi are communities of soil organisms often targeted to assist in the achievement of multiple ecological restoration goals. In drylands, benefits conferred from biocrust and AM fungal inoculation, such as improved native plant establishment and soil stabilization, have primarily been studied separately. However, comparisons between these two types of soil inoculants and investigations into potential synergies between them, particularly at the plant community scale, are needed to inform on‐the‐ground management practices in drylands. We conducted two full‐factorial experiments—one in greenhouse mesocosms and one in field plots—to test the effects of AM fungal inoculation, biocrust inoculation, and their interaction on multiple measures of dryland restoration success. Biocrust inoculation promoted soil stabilization and plant drought tolerance, but had mixed effects on native plant diversity (positive in greenhouse, neutral in field) and productivity (negative in greenhouse, neutral in field). In greenhouse mesocosms, biocrust inoculation reduced plant biomass, which was antagonistic to % root length colonized by AM fungi. Inoculation with native or commercial AM fungi did not influence plant establishment, drought tolerance, or soil stabilization in either study, and few synergistic effects of simultaneous inoculation of AM fungi and biocrusts were observed. These results suggest that, depending on the condition of existing soil communities, inoculation with AM fungi may not be necessary to promote dryland restoration goals, while inoculation with salvaged biocrust inoculation may be beneficial in some contexts.     

Addressing barriers to improve biocrust colonization and establishment in dryland restoration

Methods to reduce soil loss and associated loss of ecosystem functions due to land degradation are of particular importance in dryland ecosystems. Biocrusts are communities of cyanobacteria, lichens, and bryophytes that are vulnerable to soil disturbance, but provide vital ecosystem functions when present. Biocrusts stabilize soil, improve hydrologic function, and increase nutrient and carbon inputs. Methods to reestablish biocrust rapidly, when lost from ecosystems, have the potential to restore important dryland ecosystem functions and thereby increase probability of successful rehabilitation. The aim of this study was to identify habitat ameliorations to enhance the success of biocrust inoculation by: (1) reducing physiological stress on biocrusts and increasing resource availability (using shade, soil surface roughening, and watering), and (2) stabilizing mobile soils (using straw borders, three soil tackifiers [soil stabilizers], and a combination of shade, water, roughening, and tackifier). In the Great Basin Desert on the Utah Test and Training Range near Salt Lake City, we applied field‐harvested biocrust material to experimental plots on coarse‐ and fine‐textured soils with the top 2 cm of soil and biocrust removed. Habitat ameliorations were applied with and without biocrust addition. Shade provision increased biocrust cover 50% over controls. Biocrust cover and soil stability were 65% lower in straw border plots relative to controls. Soil tackifiers, alone and in combination with resource augmentation and stress reduction, did not improve cover and stabilization over inoculated controls. We found variability in recovery by time and between soil types. These results suggest plausible strategies to improve success of biocrust inoculation.     

Cyanobacterial persistence and influence on microbial community dynamics over 15 years in induced biocrusts

Biocrusts provide numerous ecological functions in drylands. Recovering biocrusts via cyanobacterial inoculation recently gathered interest for ecological restoration, yet it still lacks long-term experiments to unravel biocrust community dynamics. To examine how cyanobacterial inoculants influenced local microbial community and biocrust development, we observed a 2 km² (Qubqi Desert, China) inoculation experiment after 10 and 15 years, following biocrust formation. Our results revealed that biocrust development was in line with ecological regime shift, providing evidence for biocrust community succession, from cyanobacteria- to moss-dominated types. Associated with biocrust development, microbial communities differed significantly with less specialists compared to shifting sands. Cyanobacterial community analysis showed that Microcoleus vaginatus and Scytonema javanicum are an ideal inoculating model, as they were still dominating the community after 15 years since inoculation, while other nitrogen-fixing cyanobacteria occurred profusely with biocrust development. Biocrust community composition combined with thickness, Chl-a and exopolysaccharide measurements revealed the large variation of cyanobacterial ecological functions along biocrust development, suggesting a main function shift: from carbon fixation associated with exopolysaccharide secretion in bare sandy soils to nitrogen fixation in developed biocrusts. This large-scale field study verifies that cyanobacterial inoculation accelerates biocrust development and forwards succession, shaping the biocrust community composition over a long time.     

Strategies for restoring the structure and function of lichen‐moss biocrust communities

Biological soil crusts (biocrusts) are crucial components of dryland ecosystems, but they are slow to recover following disturbance. Herein, we evaluated several methods for restoring lichen‐moss biocrusts that included factorial applications of moss fragments in a water‐slurry (1) with and without lichen fragments (to restore biocrust taxonomic structure), (2) with and without clay (to facilitate establishment), and (3) with and without jute ground cloth (to facilitate establishment). Three and four years after inoculation, moss and lichen cover was up to five and eight times higher on jute ground cloth than on bare ground, respectively. Lichen cover was six times higher in plots where lichen fragments were added. Clay amendments did not increase moss or lichen establishment. To understand the effects of biocrust recovery on soil properties, we measured soil inorganic nitrogen, microbial biomass carbon, and soil water availability in restoration and control plots. Restored biocrusts decreased inorganic NH₄‐N availability by 67% when compared to controls 3 years after inoculation, but did not influence the availability of inorganic NO₃‐N, soil water, or microbial biomass carbon. Our results demonstrate that adding a biocrust inoculant to jute ground cloth can expedite recovery of lichen‐moss biocrust and reestablish its influence on soil properties within a few years.     

黄土高原封禁林地藓结皮呼吸速率对放牧踩踏干扰强度的响应

放牧踩踏造成的土壤属性变化是引起土壤呼吸速率和碳固排波动的重要原因,但目前有关不同强度放牧踩踏对生物结皮呼吸的影响尚不明确。本研究针对黄土高原风沙土发育的藓结皮,以未干扰为对照,分别进行强度为10%、30%、50%和70%的模拟放牧踩踏干扰,连续测定了藓结皮呼吸速率的变化,分析了藓结皮呼吸速率对不同干扰强度的响应规律。结果表明: 1)适度干扰会激发藓结皮呼吸速率,而重度干扰则会抑制其呼吸速率。与未干扰相比,干扰10%和30%的藓结皮呼吸速率分别增加了41.1%和22.2%,而干扰50%和70%的藓结皮呼吸速率则分别降低了8.9%和15.3%。2)踩踏干扰显著改变了土壤温度,但对土壤含水量的影响不显著。与未干扰相比,干扰10%和30%的土壤温度分别显著降低了0.4和1.2 ℃,干扰50%和70%则分别显著增加了1.1和1.0 ℃。3)不同干扰强度下藓结皮呼吸速率与土壤温度和含水量分别呈显著指数和线性正相关关系,但与藓结皮基本特性无显著相关关系,土壤温度和水分可分别解释藓结皮呼吸速率动态变化的70.6%～96.3%和49.1%～70.0%。综上,放牧踩踏显著影响了藓结皮呼吸速率,短期适度放牧踩踏会激发藓结皮呼吸速率,而过度放牧踩踏则会抑制其呼吸速率。因此,在未来黄土高原地区土壤碳收支平衡研究中应考虑不同强度放牧踩踏对生物结皮呼吸的影响。     

Production of moss‐dominated biocrusts to enhance the stability and function of the margins of artificial water bodies

Margins of water reservoirs associated with dams can have high‐frequency tides, promoting soil erosion and nutrient leaching. We propose the use of biocrusts for restoration and ecological engineering purposes, due to their poikilohydric character, to stabilize reservoir margins. We promoted biocrust growth under controlled conditions, testing two types of substrate: native sand and organic substrate. After 2 months, biocrusts grew on organic substrate covering almost all the area, but not on native sand. This fast and easy nature‐based solution for soil stabilization can be used as an environmental engineering tool in highly degraded sites.     

Plant recovery techniques do not ensure biological soil‐crust recovery after gypsum quarrying: a call for active restoration

Biological soil crusts (biocrusts) are a key component of dryland ecosystems worldwide. However, large extensions of biocrusts are disturbed by human activities, gypsum quarry being an outstanding example. Restoration techniques applied have offered satisfactory results for vascular plants but they could greatly differ in promoting biocrust recovery. A basic question remains unaddressed: can measures for plant recovery accelerate or promote the recovery of biological crusts? We have examined eight different situations: undisturbed natural habitat, five treatments with no restoration measures (overgrazed area, abandoned quarry, topsoil removal from natural habitat, and two areas filled with gypsum mining spoil), and 2 areas receiving restoration measures (manual sowing and hydroseeding). We took 40 soil cores to determine cover of lichen, moss, and cyanobacteria. Biocrust richness and cover were higher in the undisturbed habitat, with remarkable differences for the different components among treatments. Cyanobacteria were well represented in all the cores (restored and non‐restored). Mosses were promoted the most by hydroseeding. Lichen cover was remarkably higher in undisturbed samples, very low in the quarry abandoned in 1992, and 0 in the rest. Complete spontaneous recovery of biocrusts was inefficient in the 25‐year period examined. Plant restoration measures could speed up its recovery comparing with non‐restored areas. Cyanobacteria and mosses can spontaneously recover fairly well. However, promoting them would accelerate the appearance of lichen. For lichen, inoculation or translocation of lichen thalli might be proposed. Therefore, our results call for the inclusion of active restoration measures of biocrust components in recovery plans, especially for lichens.     

Production of greenhouse‐grown biocrust mosses and associated cyanobacteria to rehabilitate dryland soil function

Mosses are an often‐overlooked component of dryland ecosystems, yet they are common members of biological soil crust communities (biocrusts) and provide key ecosystem services, including soil stabilization, water retention, carbon fixation, and housing of N₂ fixing cyanobacteria. Mosses are able to survive long dry periods, respond rapidly to precipitation, and reproduce vegetatively. With these qualities, dryland mosses have the potential to be an excellent dryland restoration material. Unfortunately, dryland mosses are often slow growing in nature, and ex situ cultivation methods are needed to enhance their utility. Our goal was to determine how to rapidly produce, vegetatively, Syntrichia caninervis and S. ruralis, common and abundant moss species in drylands of North America and elsewhere, in a greenhouse. We manipulated the length of hydration on a weekly schedule (5, 4, 3, or 2 days continuous hydration per week), crossed with fertilization (once at the beginning, monthly, biweekly, or not at all). Moss biomass increased sixfold for both species in 4 months, an increase that would require years under dryland field conditions. Both moss species preferred short hydration and monthly fertilizer. Remarkably, we also unintentionally cultured a variety of other important biocrust organisms, including cyanobacteria and lichens. In only 6 months, we produced functionally mature biocrusts, as evidenced by high productivity and ecosystem‐relevant levels of N₂ fixation. Our results suggest that biocrust mosses might be the ideal candidate for biocrust cultivation for restoration purposes. With optimization, these methods are the first step in developing a moss‐based biocrust rehabilitation technology.     

生物土壤结皮演替对土壤生态化学计量特征的影响

生物土壤结皮(生物结皮)是干旱半干旱地区重要的地表覆被物,能够固定碳氮,影响养分循环,从而可能引起土壤化学计量特征的变化。以黄土丘陵区不同演替阶段生物结皮为研究对象,研究该区生态恢复初期生物结皮演替对土壤碳(C)、氮(N)、磷(P)含量及其化学计量特征的影响。结果表明:(1)生物结皮显著增加了生物结皮层C、N、P含量,后期藓结皮较初期藻结皮C、N、P含量分别增加了161%、127%和9%,均显著高于其下0—10 cm土壤;(2)土壤C/N随着生物结皮演替变化较小,范围为10.0—11.8,C/P和N/P在演替后期分别是初期的2.4倍和2.1倍,均显著高于其下0—10 cm土壤;(3)生物结皮土壤C/N、C/P与N/P受坡向影响较大,并与藓生物量显著正相关,与土壤容重显著负相关;(4)生物结皮显著影响土壤C、N、P含量及其化学计量特征,使生物结皮层显著不同于其下层土壤及裸地;(5)生物结皮演替加速表层土壤养分恢复,影响程度可至其下2 cm土层。研究初次从土壤化学计量特征的角度揭示生物结皮对土壤养分循环的贡献,对干旱半干旱地区生态修复和管理具有指导意义。     

A practical guide to measuring functional indicators and traits in biocrusts

Biocrusts are multifunctional communities that are increasingly being used to restore degraded or damaged ecosystems. Concurrently, restoration science is shifting away from the use of purely structural metrics, such as relative abundance, to more functional approaches. Although biocrust restoration technology is advancing, there is a lack of readily available information on how to monitor biocrust functioning and set appropriate restoration goals. We therefore compiled a selection of 22 functional indicators that can be used to monitor biocrust functions, such as CO₂ exchange as an indicator of productivity or soil aggregate stability as a proxy for erosion resistance. We describe the functional importance of each indicator and the available protocols with which it may be measured. The majority of indicators can be measured as a functional trait of species by using patches of biocrust or cultures that contain only one species. Practitioners wishing to track the multifunctionality of an entire biocrust community would be advised to choose one indicator from each broad functional group (erosion resistance, nutrient accumulation, productivity, energy balance, hydrology), whereas a targeted approach would be more appropriate for projects with a key function of interest. Because predisturbance data are rarely available for biocrust functions, restoration goals can be based on a closely analogous site, literature values, or an expert elicitation process. Finally, we advocate for the establishment of a global trait database for biocrusts, which would reduce the damage resulting from repeated sampling, and provide a wealth of future research opportunities.     

Amendments fail to hasten biocrust recovery or soil stability at a disturbed dryland sandy site

In most drylands, biological soil crusts (biocrusts), an assemblage of lichens, bryophytes, fungi, green algae, and cyanobacteria, are critical to healthy ecosystem function. However, they are extremely sensitive to disturbance and attempts to facilitate their recovery have had variable success. In this study, we applied soil amendments designed to improve soil surface stability and accelerate biocrust recovery on an area disturbed by oil/gas exploration vehicles. Treatments included: (1) control (one time water only); (2) biocrust‐only: biocrust inoculum + nutrients in water; (3) polyacrylamide gels (which are known to stabilize soils) + biocrust inoculum + nutrients in water; (4) gypsum + biocrust inoculum + nutrients in water; and (5) saline (NaCl) solution + biocrust inoculum + nutrients in water. Only the NaCl treatment showed any effects on soil properties and these were only short term. These effects included an increase in soil strength and a reduction in soil aggregate stability, unsaturated hydraulic conductivity (K_h), and cyanobacterial biomass. The inoculated biocrust material failed to develop and even after 10 years, there was only a very low natural recolonization of the plots. These results show that inoculating soils or applying these levels of soil amendments does not guarantee recovery of soil stability or biocrust, and that some sites are unlikely to recover without assistance. Thus, there is a need for more research into ways to enhance soil stability and identify the factors limiting biocrust establishment.     

Establishing rates of lateral expansion of cyanobacterial biological soil crusts for optimal restoration

Aims

Biocrusts that form on topsoils contribute ecosystem services to drylands, and their loss under anthropogenic pressure has negative ecological consequences. Therefore, development of biocrust inoculation technology for restoration is of interest. This requires knowledge of biocrust growth and dispersal. To contribute to this, we determined the speed at which biocrusts expand laterally based on the self-propelled motility of cyanobacteria.

Methodology

We inoculated sterile soil with natural biocrusts and incubated them over a year in a greenhouse under conditions mimicking local precipitation, monitoring the crust’s lateral expansion using time-course photography, chlorophyll a content, and microscopic inspection. Concurrent uninoculated controls served to monitor, and discount, natural inoculation by aeolian propagules.

Results

While the expansion front was highly variable in space, biocrusts expanded in the order of 2 cm month^?1, but only in seasons with moderate temperatures (Spring and Fall). Microcoleus vaginatus, Microcoleus steenstrupii, and Scytonema spp. advanced at averages of 1 cm month^?1, the crust advance front being preferentially driven by specialized propagules (hormogonia). These rates are within expectations based on instantaneous gliding motility speeds of cyanobacteria.

Conclusions

Based on the expansion capability of biocrusts during growth seasons, greenhouse inoculum units can be optimally spaced to fill 4–8 cm gaps.

     

生物结皮光合作用对光温水的响应及其对结皮空间分布格局的解译——以黄土丘陵区为例

生物结皮是具有生命活性的地表覆被物,在黄土丘陵半干旱区广泛发育。迄今有关该区不同类型生物结皮的光合、呼吸作用潜力及其影响因素还不清楚。通过野外调查采样,利用室内半开放光合测定系统,测定了黄土丘陵区典型的藻类(3年生)、藓类(土生扭口藓)生物结皮(13年生)光合及呼吸作用过程对水分、温度及光照等环境因子的响应,在此基础上,解释了不同类型生物结皮空间分布格局的内在原因。结果表明:(1)在适合的光温水条件下,生物结皮的光合及呼吸作用强度取决于生物组成,研究区藓结皮的光合及呼吸作用速率显著高于藻结皮;(2)水分是生物结皮光合、呼吸作用的关键影响因子,只有当水分含量达到一定水平,才有光合及呼吸的可能。黄土丘陵区藓类及藻类生物结皮光合作用最适水分条件分别为80%-100%和40%-80%田间持水量。(3)藓类及藻类生物结皮光合作用光补偿点PAR均低于10μmol.m-2.s-1,光饱和点PAR分别为1000和800μmol.m-2.s-1,研究中2类生物结皮均未出现光抑制现象。(4)温度显著影响生物结皮的光合及呼吸作用,研究区藓类及藻类生物结皮的最适光合温度分别为20-25℃和25-30℃。(5)生物结皮光合作用对含水量及温度的响应可以解释结皮的空间分布特征。藻类植物光合作用的最适含水量较低而最适温度较高,因此在失水较快的阳坡及开阔地面上依然可以大面积发育;而藓类植物光合作用对水分条件要求较高,因此在水分条件较好的阴坡、低洼地才易演替至藓结皮阶段。     

A unique role for citizen science in ecological restoration: a case study in streams

Citizen science has the potential to generate valuable biologic data for use in restoration monitoring, while also providing a unique opportunity for public participation in local restoration projects. In this article, we describe and evaluate a citizen science program designed to monitor the effect of stream restoration construction disturbance on the macroinvertebrate community. We present the results of a 7‐year stream restoration study conducted by citizen scientists utilizing a Before‐After‐Control‐Impact (BACI) design. Trait‐based macroinvertebrate data showed a strong response to restoration construction disturbance and return to pre‐restoration conditions within 2 years. The findings of this study suggest that citizen science can generate meaningful BACI‐oriented data about ecological restoration; however, until more research is conducted, citizen data should only be used to augment professional data intended to demonstrate restoration success.     

Potential for synergy in soil inoculation for nature restoration by mixing inocula from different successional stages

Background and aims

Due to the well-known importance of biocrusts for several ecosystem properties linked to soil functionality, we aim to go deeper into the physiological performance of biocrusts components. Possible functional convergences in the physiology of biocrust constituents would facilitate the understanding of both species and genus distributional patterns and improve the possibility of modelling their response to climate change.

Methods

We measured gas exchange in the laboratory under controlled conditions of lichen- and moss-dominated biocrusts from four environmentally different locations in Europe. Field data were used to determine the natural hydration sources that drive metabolic activity of biocrusts.

Results

Our results show different activity drivers at the four sites. Within site analyses showed similar C fixation for the different crust types in the three sites without hydric stress whilst light use related parameters and respiration at 15 °C were similar in the between sites analyses. There were significant differences in water relations between the biocrusts types, with moss-dominated crusts showing higher maximum and optimum water contents.

Conclusions

The functional type approach for biocrusts can be justified from a physiological perspective when similar values are found in the within and between site analyses, the latter indicating habitat independent adaptation patterns. Our multi-site analyses for biocrusts functional performance provide comparisons of C fluxes and water relations in the plant-soil interface that will help to understand the adaptation ability of these communities to possible environmental changes.

     

黄土丘陵区生物结皮空间分布的代表性景观指数

景观指数可量化表征生物结皮的分布特征,但景观指数存在数量众多、冗余度较高的问题。本研究以位于典型黄土丘陵区的陕西省吴起县合沟小流域不同分布格局的58个样方的生物结皮为对象,计算其15个常用景观格局指数,基于相关分析、因子分析和敏感性分析,筛选出可描述生物结皮景观格局、具有特定生态学意义的代表性景观指数,并以陕西省定边县杨井镇鹰窝山涧小流域退耕地不同盖度的生物结皮为例,检验所选景观格局指数的可靠性和合理性。结果表明: 15个景观指数中,10个指数之间具有显著相关关系,而边缘长度(TE)和边缘密度(ED)分别与斑块数量(NP)、斑块密度(PD)、丛生度(CLUMPY)和散布与并列指数(IJI)无显著相关性。斑块占景观比例(PLAND)、ED、斑块连结度(COHESION)和分离度(SPLIT),分别从盖度、长度、连通度和破碎度方面描述了生物结皮空间分布特征,其代表的3个公因子在刻画生物结皮空间分布格局时累计贡献率达91.6%。本研究确定了可量化生物结皮空间分布复杂性的代表性景观指数,为研究生物结皮格局变化及其与生态过程的关系提供了理论依据。