Biological soil crust development affects physicochemical characteristics of soil surface in semiarid ecosystems

Water and nutrients are scarce resources in arid and semiarid ecosystems. In these regions, biological soil crusts (BSCs) occupy a large part of the soil surface in the open spaces surrounding patches of vegetation. BSCs affect physicochemical soil properties, such as aggregate stability, water retention, organic carbon (OC) and nitrogen (N) content, associated with primary ecosystem processes like water availability and soil fertility. However, the way BSCs modify soil surface and subsurface properties greatly depends on the type of BSC. We hypothesised that physicochemical properties of soil crusts and of their underlying soils would improve with crust development stage. Physicochemical properties of various types of soil crusts (physical crusts and several BSC development stages) and of the underlying soil (soil layers 0–1 cm and 1–5 cm underneath the crusts) in two semiarid areas in SE Spain were analysed. The properties that differed significantly depending on crust development stage were aggregate stability, water content (WC) (at −33 kPa and −1500 kPa), OC and N content. Aggregate stability was higher under well-developed BSCs (cyanobacterial, lichen and moss crusts) than under physical crusts or incipient BSCs. WC, OC and N content significantly increased in the crust and its underlying soil with crust development, especially in the first centimetre of soil underneath the crust. Our results highlight the significant role of BSCs in water availability, soil stability and soil fertility in semiarid areas.     

Spatial distribution and physiology of biological soil crusts from semi-arid central Spain are related to soil chemistry and shrub cover

Despite the critical role of biological soil crusts (BSCs) in arid and semi-arid ecosystem function, few studies are found concerning the most important environmental variables affecting their distribution and physiology. This study seeks to determine soil and microenvironmental factors affecting the spatial distribution and pigment production of BSC-forming lichens and mosses in open patches of a semi-arid Mediterranean kermes oak thicket. We measured late-successional BSC cover, shrub cover, distance to nearest kermes oak (to test for effects of kermes oak thicket microenvironment on BSC), and pigment concentration of one lichen (Cladonia foliacea) and one moss (Pleurochaete squarrosa) species in the Nature Reserve El Regajal-Mar de Ontígola (Central Spain). At the macroscale (>0.5 m), results showed that BSC distribution and pigments were tightly coupled to a suite of soil properties, in particular soil pH, Fe, and Ca. Specifically, soil pH had a positive relationship with the cover of five individual BSC-forming lichen species and was negatively related to pigment production in C. foliacea. When pH was excluded from the analysis, Ca appeared as the main soil variable and was correlated with total BSC cover and total lichen cover. The micronutrient Fe had a significant positive relationship with the concentration of eight pigments in P. squarrosa and was also coupled with the cover of two BSC-forming lichens. Manganese, previously proposed as a key limiting micronutrient for BSCs, affected lichen diversity in a negative way. At the microscale (∼0.5 m), kermes oak microenvironment, shrub cover, and moss cover were determinants of BSC distribution, and total lichen and total BSC cover were overrepresented on N and E-facing shrub microsites. Our findings suggest that soil chemical variability and microsite diversity created by neighbouring vegetation affect BSC distribution in complex and essential ways and that studies aiming to explore BSC-environment relationships should be conducted at various spatial scales. Studies based on species- or group-specific responses are, thus, inadequate to unveil the main factors determining the distribution of the diverse organisms that constitute BSCs and/or to propose potential tools aiming to restore BSC in arid and semiarid ecosystems.     

Hydrolase enzyme activities in a successional gradient of biological soil crusts in arid and semi-arid zones

In arid and semi-arid regions, pioneer organisms form complex communities that penetrate the upper millimetres of the bare substrate, creating biological soil crusts (BSC). These thin crusts play a vital role in whole ecosystem functioning because they enrich bare surfaces with organic matter, initiate biogeochemical cycling of elements, modify hydrological cycles, etc., thus enabling the ground to be colonized by vascular plants. Various hydrolase enzymes involved in the carbon (cellulase, β-glucosidase and invertase activities), nitrogen (casein-protease and BAA-protease activities) and phosphorus (alkaline phosphomonoesterase activity) cycles were studied at three levels (crust, middle and deep layers) of three types of BSCs from the Tabernas Desert (SE Spain), representing an ecological gradient ranging from crusts predominated by cyanobacteria to crusts predominated by lichens (Diploschistes diacapsis, Lepraria crassissima). All enzyme activities were higher in all layers of all BSCs than in the bare substrate. The enzymes that hydrolyze low molecular weight substrates were more active than those that hydrolyze high molecular weight substrates (cellulase, casein-protease), highlighting the pioneering characteristics of the BSCs. The hydrolytic capacity developed in parallel to that of ecological succession, and the BSCs in which enzyme activity was highest were those under L. crassissima. The enzyme activity per unit of total organic C was extremely high; the highest values occurred in the BSCs formed by cyanobacteria and the lowest in those formed by lichens, which indicates the fundamental role that the primary colonizers (cyanobacteria) play in enriching the geological substrate with enzymes that enable degradation of organic remains and the establishment of more developed BSCs. The results of the study combine information on different enzyme activities and provide a clear vision of how biogeochemical cycles are established in BSCs, thus confirming the usefulness of enzyme assays as key tools for examining the relationship between biodiversity and ecosystem function in biological soil crusts.     

From peds to paradoxes: Linkages between soil biota and their influences on ecological processes

Soils and their biota have been studied by a variety of observational and experimental methods that have allowed biologists to infer their structural and functional interactions. Viewing progress made over the last 10 years, it is apparent that an increasing diversity of analytical and chemical methods are providing much more detailed information about feeding preferences and niche overlaps of speciose groups such as oribatid mites.Other topics in which considerable progress has been made include new advances in knowledge of root and mycorrhiza production and turnover and their impacts on soil ecosystems; soil biota, exudations and secretions and soil aggregation phenomena; the biology of invasive species in soils; soil biodiversity, legacies and linkages to soil processes; and soil biodiversity and ecosystem functional responses. Throughout the review I suggest ways for new progress to be made in future research.     

Terrestrial ecosystem processes of Victoria Land, Antarctica

Terrestrial environments of Victoria Land, Antarctica are ideal systems to test hypotheses about the sensitivity of ecosystem processes to climate variability, and the relationships between soil biodiversity and ecosystem functioning because of their high sensitivity to climate change and their limited diversity. This region is also considered among the most pristine of ecosystems, and therefore may serve as an indicator for detecting the response of other ecosystems to global environmental change. Rates and controls over key ecosystem processes remain poorly documented over much of Victoria Land, but it is generally held that the distribution and functioning of soil communities are most limited by the availability of liquid water and organic carbon. Here we review examples of ecosystem processes from several sites in North and South Victoria Land and develop a regional synthesis accounting for variation in the availability of soil resources (i.e. liquid water, organic matter, inorganic nutrients). Variation in soil microclimate, organic matter, moisture and salinity encountered over gradients of coastal to interior sites, latitude, and soil chronosequences are the primary controls over the structure of soil communities and their functioning. Imbalanced stoichiometric nutrient ratios frequently encountered in Victoria Land ecosystems also contribute to limited distribution of soil biota, and where they occur these elemental imbalances indicate lower biological activity and little biotic control over bulk element ratios in soils. Priorities and future directions of Victoria Land soil and ecosystem research are also discussed.     

Effect of invasive Acacia dealbata Link on soil microorganisms as determined by PCR-DGGE

Acacia dealbata Link is an Australian woody legume that has become a serious environmental problem in Northwest Spain where it forms dense monospecific patches modifying the structure of different native ecosystems and threatening native aboveground biodiversity. In spite of the dramatic changes observed in the vegetation of invaded sites little is known about the consequences of invasion for soil microorganisms. To investigate the effect of A. dealbata invasion on the structure of soil fungi and bacteria communities, samples were taken from invaded and non-invaded areas from three different ecosystems in Northwest Spain: pine forest, shrubland and grassland. In each ecosystem type, soil samples were taken in areas of native vegetation, areas invaded by A. dealbata and in the transition zone between native and invaded vegetation. Soil microorganisms were analyzed in the different samples by PCR-DGGE using general primers for eubacteria and fungi. Soil analyses were also performed to evaluate the effect of A. dealbata invasion on soil fertility.The invasion by A. dealbata consistently increased soil N, C, organic matter and exchangeable P content in the three studied ecosystems. A clear effect of the invasion on the overall structure of microorganism communities was only observed in the shrubland where soil fungal communities in the invaded and transition areas clustered together and apart from the native soil. Significant differences in soil microorganisms richness and diversity between invaded and not invaded soils were only found in the grassland. Grassland invasion by A. dealbata lead to a significant increase of bacterial richness and to a significant reduction in fungal richness and diversity. Our results show that although the changes on soil chemistry due to A. dealbata invasion are consistent among the studied ecosystems, the effect on soil microorganisms depends on the ecosystem type affected by the invasion.     

Positive relationship between herbaceous layer diversity and the performance of soil biota in a temperate forest

The current decline in biodiversity is particularly pronounced in the herbaceous layer of forest ecosystems. We explored the relationship between a naturally occurring plant diversity gradient in the understory vegetation of a deciduous forest and several above-and belowground ecosystem processes. We show that particularly soil microbial parameters and microarthropod densities are positively correlated with plant species richness. These results confirm recent findings in grassland ecosystems and highlight the intimate interconnectance between the diversity and functioning of above-and belowground compartments. We conclude that irrespective of a potential causal relationship between plant species richness and belowground processes, it is essential to consider the performance of soil biota in order to understand the relationship between herbaceous layer composition and ecosystem function.     

黄土高原生物土壤结皮研究进展与展望

黄土高原是典型的生态脆弱敏感区和世界上水土流失最为严重的地区,也是当今我国生态恢复和生态文明建设的重点区域.生物土壤结皮是细菌、藻类、真菌和孢子植物与土壤颗粒胶结而成的有机复合体,是干旱半干旱地区地表系统的重要组成部分,它们对黄土高原的水土保护、养分积累和生态恢复具有重要的生态功能.本文论述了生物土壤结皮类型与演替过程...     

生物结皮土壤-水文-侵蚀效应研究进展

生物土壤结皮(简称生物结皮)是由隐花植物、微小生物和土壤表层颗粒胶结形成的一种特殊复合体,广泛分布于各类气候和生境条件。作为生态系统的重要组分,生物结皮在不同生物气候区土壤的生态过程、水文过程、生物过程、地球化学循环过程以及生态修复过程中发挥着重要作用。从生物结皮影响土壤物理、化学、生物学性质以及土壤水文与侵蚀过程等方面对其生态功能进行总结和概述,在此基础上,从研究区域和时空尺度、多过程耦合机制、生物结皮影响氮循环的过程及其对氮沉降的响应机制、生物结皮与维管植物空间分布及互动关系等方面,展望了该领域有待深化的问题及今后的发展方向,以期促进我国生物结皮相关研究工作,加深对生物结皮生态功能及地表过程的认识。     

Discriminating soil crust type,development stage and degree of disturbance in semiarid environments from their spectral characteristics

Biological soil crusts (BSCs) are increasingly recognized as common features in arid and semiarid ecosystems and play an important role in the hydrological and ecological functioning of these ecosystems. However, BSCs are very vulnerable to, in particular, human disturbance. This results in a complex spatial pattern of BSCs in various stages of development. Such patterns, to a large extent, determine runoff and erosion processes in arid and semiarid ecosystems. In recent years, visible and near infrared (Vis‐NIR) diffuse reflectance spectroscopy has been used for large‐scale mapping of the distribution of BSCs. Our goals were (i) to demonstrate the efficiency of Vis‐NIR spectroscopy in discriminating vegetation, physical soil crusts, various developmental stages of BSCs, and various types of disturbance on BSCs and (ii) to develop a classification system for these types of ground cover based on Vis‐NIR spectroscopy. Spectral measurements were taken of vegetation, physical crusts and various types of BSCs prior to, and following, trampling or removal with a scraper in two semiarid areas in SE Spain. The main spectral differences were: (i) absorption by water at about 1450 nm, more intense in the spectra of vegetation than in those of physical crusts or BSCs, (ii) absorption features at about 500 and 680 nm for the BSCs, which were absent or very weak for physical crusts, (iii) a shallower slope between about 750 and 980 nm for physical crusts and early‐successional BSCs than for later‐successional BSCs and (iv) a steeper slope between about 680 and 750 nm for the most developed BSCs. A partial least squares regression‐linear discriminant analysis of the spectral data resulted in a reliable classification (Kappa coefficients over 0.90) of the various types of ground cover and types of BSC disturbance. The distinctive spectral features of vegetation, physical crusts and the various developmental stages of BSCs were used to develop a classification system. This will be a promising tool for mapping BSCs with hyperspectral remote sensing.     

Interaction between humus form and herbicide toxicity to Collembola (Hexapoda)

Laboratory experiments were conducted using intact collembolan communities, exposed to Madit D^®, a phenylurea herbicide (active ingredient isoproturon). Effects were investigated using two distinct humus types, an acid Dysmoder and a neutral Eumull. Within two weeks, no effect of the herbicide was displayed by the Eumull population, while the Dysmoder population was stimulated. When animals were able to escape from the herbicide through a perforated wall separating two compartments filled with natural soil, the behavior of collembolan communities exhibited interactive (non-additive) effects of humus type and herbicide application. The combination of an acid soil (supposedly providing greater tolerance to organic pollutants) with a neutral soil, increased biodiversity of Collembola, but caused the disappearance of some acido-sensitive species, pointing to complex relationships between pesticides, soils and soil organisms. Parallel experiments with single species demonstrated that at the recommended dose Madit D^® may cause avoidance effects, but no toxicity.     

Spatial congruence between biodiversity and ecosystem services in South Africa

Ecosystems services sustain humans all over the world. The unsustainable use of ecosystem services around the world has led to widespread degradation which now threatens human health and livelihoods. Although the maintenance of ecosystem services is often used to justify biodiversity conservation actions, it is still unclear how ecosystem services relate to different aspects of biodiversity and to what extent the conservation of biodiversity will ensure the provision of services. The aim of this study was to find out whether biodiversity priorities, biomes, species richness and vegetation diversity hotspots co-occur in space with ecosystem services. The distribution of the ranges and hotspots of five ecosystem services (surface water supply, water flow regulation, carbon storage, soil accumulation, and soil retention) was assessed in South African biomes. Coincidence, overlap, and correlation analyses were used to assess spatial congruence between ecosystem services and species richness (plants and animals) and vegetation diversity hotspots. The grassland and savanna biomes contained significant amounts of all five ecosystem services. There was moderate overlap and a generally positive but low correlation between ecosystem services hotspots and species richness and vegetation diversity hotspots. Species richness was mostly higher in the hotspots of water flow regulation and soil accumulation than would be expected by chance. The water services showed varying levels of congruence with species richness hotspots and vegetation diversity hotspot. These results indicate that actions taken to conserve biodiversity in South Africa will also protect certain ecosystem services and ecosystem services can be used to strengthen biodiversity conservation in some instances.     

Functional aspects of soil animal diversity in agricultural grasslands

There has been recent interest in the characterization of soil biodiversity and its function in agricultural grasslands. Much of the interest has come from the need to develop grassland management strategies directed at manipulating the soil biota to encourage a greater reliance on ecosystem self-regulation. This review summarises information on selected groups of soil animals in grasslands, the factors influencing their abundance, diversity and community structure and their relationships to the functioning and stability of grassland ecosystems. Observations on the impacts of agricultural managements on populations and communities of soil fauna and their interactions confirm that high input, intensively managed systems tend to promote low diversity while lower input systems conserve diversity. It is also evident that high input systems favour bacterial-pathways of decomposition, dominated by labile substrates and opportunistic, bacterial-feeding fauna. In contrast, low-input systems favour fungal-pathways with a more heterogeneous habitat and resource leading to domination by more persistent fungal-feeding fauna. In view of this, we suggest that low input grassland farming systems are optimal for increasing soil biotic diversity and hence self-regulation of ecosystem function. Research is needed to test the hypothesis that soil biodiversity is positively associated with stability, and to elucidate relationships between productivity, community integrity and functioning of soil biotic communities.     

Plant–soil interactions in metal contaminated soils

The legacy of industrialization has left many soils contaminated. However, soil organisms and plant communities can thrive in spite of metal contamination and, in some cases, metabolize and help in remediation. The responses of plants and soil organisms to contamination are mutually dependent and dynamic. Plant–soil feedbacks are central to the development of any terrestrial community; they are ongoing in both contaminated and healthy soils. However, the theory that governs plant–soil feedbacks in healthy soils needs to be studied in contaminated soils. In healthy soils, negative feedbacks (i.e. pathogens) play a central role in shaping plant community structure. However to our knowledge, the nature of feedback relationships has never been addressed in contaminated soils. Here we review literature that supports a plant–soil feedback approach to understanding the ecology of metal-contaminated soil. Further, we discuss the idea that within these soils, the role of positive as opposed to negative plant–soil feedbacks may be more important. Testing this idea in a rigorous way in any ecosystem is challenging, and metal contamination imposes an additional abiotic constraint. We discuss research goals and experimental approaches to study plant–soil interactions applicable to metal-contaminated soils; these insights can be extended to other contaminated environments and restoration efforts.     

A future for soil ecology ? Connecting the system levels: moving from genomes to ecosystems: Opening Lecture to the XIII ICSZ “Biodiversity of soil organisms and ecosystem functioning”

As an introduction to the XIII ICSZ “Biodiversity of soil organisms and ecosystem functioning” the question is raised what contribution soil ecology has made to general ecology. Although the appearance of soil ecological papers in general ecological journals is limited, soil ecologists have had a major contribution to especially functional aspects of ecology like nutrient release and energy turnover mechanisms, perhaps partly due to the applied character of soil ecology. As a way forward it is suggested to unify further work and scientific discussions according to five research themes: 1. Combined spatial and temporal heterogeneity; 2. Scaling up from individual mobility via distribution patterns to bio-geography; 3. Structural and functional biodiversity: from gene- to ecosystem level; 4. Nutrient cycling/energy transfer at the micro- to macro-level; 5. Adaptability from bacteria to ecosystems: is there a mutual mechanism connecting genetic variation with ecosystemś adaptive mechanisms?     

Linking species richness, biodiversity and ecosystem function in soil systems

Soils are the central organizing entities in terrestrial ecosystems and possess extremely diverse prokaryotic and eukaryotic biota. They are physically and chemically complex, with micro- and macro-aggregates embedded within a solid, liquid and gaseous matrix that is continually changing in response to natural and human-induced perturbations. Recent advances in molecular techniques in systematics have provided opportunities for the study of biodiversity and biocomplexity of soil biota. A symposium and workshop on soil biogeochemistry and biodiversity International Symposium on Impacts of Soil Biodiversity on Biogeochemical Processes in Ecosystems, Taiwan Forestry Research Institute, Taipei, Taiwan April 18-24, 2004. Convened an international array of participants working in biomes on virtually every continent on the planet (ranging from polar to tropical regions). This special issue reports on the theoretical bases and applications of molecular methods for the measurement of soil biodiversity.

Themes addressed include a melding of classical taxonomic investigations with biochemical fingerprinting and molecular probing of organism identities. Several papers highlight new advances in identifications of prokaryotic and eukaryotic organisms. Examples include new developments in “fingerprinting” of microbes active in “mycorrhizospheres” using immunocapture and other innovative techniques. Developments in the study of impacts of invasive plant and animal species on ecosystem function and subsequent microbial community composition and function have been very great in the last 2-3 years. Soils are major repositories of legacies, including fine and coarse woody debris and other organic products, which have feedbacks on soil diversity. The ways in which species diversity and function of microbial and faunal communities interact and their importance to ecosystem function are examined in biological and biochemical detail. This paper provides an overview of soil biodiversity and its feedbacks on soil biogeochemical processes in ecosystems.     

Increased frequency of drought reduces species richness of enchytraeid communities in both wet and dry heathland soils

Studies of biological responses in the terrestrial environment to rapid changes in climate have mostly been concerned with above-ground biota, whereas less is known of belowground organisms. The present study focuses on enchytraeids (Oligochaeta) of heathland ecosystems and how the enchytraeid community has responded to simulated climate change in a long-term field experiment. Either increased temperature or repeated drought was applied for 13 years to field plots located in Wales, The Netherlands and Denmark representing a gradient in precipitation and annual temperature fluctuations thereby providing an opportunity to study biological responses on a local (within sites) and regional scale. Warming treatments increasing night-time temperature (0.5–1 °C higher than ambient at 5 cm soil depth) had no detectable effects on the enchytraeid communities. Increased intensity and frequency of drought had rather weak persistent effects on total enchytraeid abundance suggesting that ecosystem functions of enchytraeids may only be transiently impacted by repeated spring or summer drought. However, drought treatment had persistent negative effects on species richness and community structure across sites. Drought treated plots harboured only 35–65% of the species present in control plots, and the reduction of species richness was most pronounced at the driest sites. It is discussed that soil invertebrates, due to their weak migratory potential, may be more liable to extinction under changing climatic conditions than above-ground species, and therefore consequences of climate change to soil organisms need particular attention in future research.     

Ethanol outperforms multiple solvents in the extraction of chlorophyll-a from biological soil crusts

In order to estimate the biomass of photoautotrophic organisms in biological soil crusts (BSCs), based on the extraction and determination of phytoplanktonic chlorophyll-a (Chl-a) in aquatic ecosystems, this paper comparatively studied the Chl-a extraction efficiencies in ethanol, acetone, N, N -dimethyl -formamide (DMF) and dimethyl sulphoxide (DMSO) from algae, lichen, and moss crusts, analyzed the effects of dominant organisms or development degree of BSCs, mass of sample, and soil characteristics on the extraction efficiency. The results showed that the extraction efficiencies of different organic solvents were significantly different, and such efficiencies declined with the increase of mass of sample. DMSO resulted in the greatest extraction efficiency, but was not suitable for lichen crusts; DMF strongly underestimated the Chl-a content of man-made crusts relative to the other extractants, and was particularly affected by interfering pigments from the well-developed crusts. Generally, the extraction efficiency of ethanol was greater than that of acetone, and ethanol method showed the greatest stability compared with other 3 solvents. In addition, Chl-a content showed an increasing trend with the succession of BSCs when it was expressed on an areal basis. On the basis of this and related experiments we suggest Chl-a content of BSCs be extracted by ethanol and expressed on an areal basis.     

Microbiota accompanying different arbuscular mycorrhizal fungal isolates influence soil aggregation

Arbuscular mycorrhizal fungi (AMF) are key components of ecosystems through their influence on plant communities and ecosystem processes. A major source of information regarding the importance of AMF species richness on process rates are mesocosm experiments using different levels of diversity of AMF as provided by single-species cultures of AMF. Since AMF inocula are generally made available in the form of non-sterile pot culture material, it is possible that AMF symbiosis-associated microbiota are at least partially responsible for some effects hitherto directly attributed to the AMF mycelium. Here, we provide evidence that microbiota associated with single-species cultures of AMF (after long-term pot culture enrichment of 7–8 years) can strongly affect the ecosystem process of soil aggregation. This effect occurred in an AMF isolate specific manner, but in the absence of live and active AMF mycelium. We additionally documented large differences in microbiota communities associated with the different AMF inocula (using PLFA analyses), suggesting that these differences were at least partly responsible for the observed changes in soil aggregation. This result points to AMF–microbiota interactions as a largely unexplored mechanism underlying soil aggregation (and potentially other ecosystem processes). We suggest that a reinterpretation of previous experiments using greenhouse-derived AMF cultures may be necessary, and the need to consider AMF symbiosis-associated microbiota in mechanistic studies of AMF and mycorrhizae in general is emphasized.     

植物多样性对土壤动物影响的研究进展

生物多样性与生态系统功能一直是生态学研究的一个热点。近些年来的研究表明,植物多样性除了影响陆地生态系统地上部分的初级生产力等生态系统功能,还会间接影响地下生物多样性及土壤生态系统过程。本文概述了植物多样性对土壤动物的影响及其主要机制,归纳了植物多样性通过改变输入土壤中的资源数量与资源多样性、微生境结构、土壤环境因子等影响土壤动物的途径。目前关于土壤动物群落对植物多样性的响应仍存在很多问题和争议,本文总结了需要进一步深入研究的方向,特别指出了要加强研究影响植物多样性与土壤动物关系的生物与非生物因子、后续的生态效应和反馈、不同机制和途径的贡献定量化等。