Restoration,soil organisms,and soil processes: emerging approaches

Better understanding of the connection between aboveground plant communities and belowground soil organisms and processes has led to an explosion in recent research on the applications of this link to the field of ecological restoration. Research is only beginning to have the capacity to link soil organisms and specific ecosystem functions. Establishing general ecological principles of the role microbial communities have during ecological restoration is also still in its infancy. As such, the literature is at a critical point to generate a Special Feature that brings together novel approaches of linking soil and restoration to promote more regular inclusion and consideration of soil organisms and soil‐based processes in ecological restoration. In this special feature, we bring together nine research articles from different ecosystems that study the relationship between restoration activities, soil microbial communities, and soil properties. From these research articles, we describe two primary themes: (1) research on the impacts of ecosystem‐specific restoration activities on soil organisms and processes and (2) research testing methods of soil manipulation to improve restoration outcomes. We hope to inspire readers and restoration practitioners to consider soil microbes and soil processes in their research, restoration projects, and world views.     

Site preparation impacts on soil biotic and abiotic properties,weed control,and native grass establishment

In severely degraded systems active restoration is required to overcome legacies of past land use and to create conditions that promote the establishment of target plant communities. While our understanding of the importance of soil microbial communities in ecological restoration is growing, few studies have looked at the impacts different site preparation techniques have on these communities. We trialed four methods of site preparation: fire, top‐soil removal (TSR; removal of top 50 mm of soil), slashing (vegetation cut to 30 mm, biomass removed), and carbon (C; as sugar and saw‐dust) addition, and quantified resulting soil bacterial communities using DNA metabarcoding. We compared the effectiveness of these techniques to reduce weed biomass, improve native grass establishment, and induce changes in soil nutrient availability. TSR was the most effective technique, leading to a reduction in both available nutrients and competition from weeds. In comparison, the remaining methods had little or no effect on weed biomass, native grass establishment, or soil nutrient availability. Both TSR and C addition resulted in changes in the soil bacterial community. These changes have the potential to alter plant community assembly in many ways, such as via nutrient acquisition, pathogenic effects, nutrient cycling, and decomposition. We recommend TSR for ecological restoration of old‐fields and suggest it is a much more effective technique than burning, slashing, or C addition. Restoration practitioners should consider how their management techniques may influence the soil biota and, in turn, affect restoration outcomes.     

Soil Organism and Plant Introductions in Restoration of Species-Rich Grassland Communities

Soil organisms can strongly affect competitive interactions and successional replacements of grassland plant species. However, introduction of whole soil communities as management strategy in grassland restoration has received little experimental testing. In a 5-year field experiment at a topsoil-removed ex-arable site ( receptor site ), we tested effects of (1) spreading hay and soil, independently or combined, and (2) transplanting intact turfs on plant and soil nematode community development. Material for the treatments was obtained from later successional, species-rich grassland ( donor site ). Spreading hay affected plant community composition, whereas spreading soil did not have additional effects. Plant species composition of transplanted turfs became less similar to that in the donor site. Moreover, most plants did not expand into the receiving plots. Soil spreading and turf transplantation did not affect soil nematode community composition. Unfavorable soil conditions (e.g., low organic matter content and seasonal fluctuations in water level) at the receptor site may have limited plant and nematode survival in the turfs and may have precluded successful establishment outside the turfs. We conclude that introduction of later successional soil organisms into a topsoil-removed soil did not facilitate the establishment of later successional plants, probably because of the "mismatch" in abiotic soil conditions between the donor and the receptor site. Further research should focus on the required conditions for establishment of soil organisms at restoration sites in order to make use of their contribution to grassland restoration. We propose that introduction of organisms from "intermediate" stages will be more effective as management strategy than introduction of organisms from "target" stages.     

Earthworms in Soil Restoration: Lessons Learned from United Kingdom Case Studies of Land Reclamation

Restoration ecology requires theoretical consideration of a habitat’s former structure and function before the practice of ecological restoration is applied. However, experience has shown that this does not always occur and aspects such as soil ecology have often been an afterthought. Here, case study material relates the use of earthworms at selected sites in the United Kingdom. Due to their soil‐forming capabilities, these organisms may be essential to reconstruction of soils when drastic activities have despoiled an area. While describing in brief the type of work undertaken, these case studies seek to illustrate some of the misunderstandings/problems/deliberately negative acts that have too often accompanied use of earthworms in soil restoration. From such experiences, implications for practice are suggested that should lead to a greater understanding and appropriate utilization of earthworms in future projects.     

Tallgrass prairie ants: their species composition,ecological roles,and response to management

Ants are highly influential organisms in terrestrial ecosystems, including the tallgrass prairie, one of the most endangered ecosystems in North America. Through their tunneling, ants affect soil properties and resource availability for animals and plants. Ants also have important ecological roles as consumers of plant tissue and seeds. In the last several decades, various organizations, agencies, and agricultural producers have attempted to create wildlife habitat or reduce soil erosion by seeding thousands of hectares of bare cropland in the central United States with tallgrass prairie seed mixes. Although initially, monitoring of these restorations and of unplowed prairie remnants focused on plants and birds, in recent years the response of invertebrates such as ants has increasingly been the subject of research. An understanding of tallgrass prairie ant communities can help land managers and scientists better monitor the ecological condition of tallgrass prairie and guide management and restoration efforts. Here I review our current knowledge of ant species found within tallgrass prairie, their ecological roles, and their response to management.     

Barriers to ecosystem restoration presented by soil legacy effects of invasive alien N2‐fixing woody species: implications for ecological restoration

Impacts of invasive alien N₂‐fixing woody species and how they can persist as soil legacy effects after invasive species control are well appreciated, but how soil legacy effects can present barriers to restoration is poorly understood. Finding better ways to deal with these barriers to restoration is essential to improving restoration outcomes. In this study, we review 440 studies to identify barriers to restoration and potential management actions for the barriers to restoration, and provide practical application examples of the management actions. Our findings suggest that altered soil microbial communities, depleted native soil seed banks, elevated N status, secondary invasion and weedy native species dominance, and reinvasion are potential barriers to restoration. Furthermore, carbon addition, litter removal, soil microbial treatments, establishing species adapted to low N levels, prescribed burning, classical biological control, grazing, mowing, herbicide or graminicide application, manual weeding, soil N management, soil solarization, weed mats, native species reintroduction, and nurse plants are potential management actions for these barriers to restoration. However, there is little evidence suggesting that several of these barriers to restoration hinder improved restoration outcomes and this could be due to little research on them. More research is needed to assess their relative importance in hindering improved restoration outcomes. Management actions are rarely applied in combination, despite that they often address distinct barriers to restoration. Management actions should be combined into an integrated management effort to improve restoration outcomes.     

Using Stable Isotopes to Differentiate Trophic Feeding Channels within Soil Food Webs

The soil is probably the most diverse habitat there is, with organisms ranging in sizes from less than 1 μm to several metres in length. However, it is increasingly evident that we know little about the interactions occurring between these organisms, the functions that they perform as individual species, or together within their different feeding guilds. These interactions between groups of organisms and physical and chemical processes shape the soil as a habitat and influence the nature of the soil food web with consequences for the above‐ground vegetation and food web. Protists are known as one of the most abundant groups of bacterivores within the soil; however, they are also consumers of a number of other food sources. Even though they are responsible for a large proportion of the mineralisation of bacterial biomass and have a large impact on the C and N cycles within the soil they are regularly overlooked when investigating the complete soil food web. Recently, stable isotopes have been used to determine trophic interactions and here we describe how this technique has been used to highlight linkages between protists and the soil food web.     

稀土尾矿土壤细菌群落结构对植被修复的响应

选用赣州-安远稀土弃废尾矿及其不同植被修复的堆浸田为研究对象,调查废弃尾矿及6种不同植被修复方案下土壤理化性质的变化,并利用变性凝胶梯度电泳(DGGE)技术,分析土壤微生物群落结构对植被修复的响应。结果表明:与未修复尾矿土壤相比,经不同植被修复后的土壤理化性质均得到明显改良,其中土壤含水量、有机质含量均比未修复尾矿土壤增加2—3倍。微生物群落结构分析表明,植被修复后土壤微生物群落与废弃尾矿土壤微生物群落亲缘度仅为0.21,表明植被修复后,土壤微生物群落结构发生了明显变化,且微生物多样性、均匀度、丰富度与未修复尾矿土壤相比均有了明显的提高。而在不同植被修复方案中,以湿地松和山胡椒为优势群落的两种植被修复方案对土壤改良效果最为明显,这两种修复方案不仅能显著改善土壤的固水性、有机质含量,并且对微生物群落的改善作用也最为显著。典范对应分析表明,废弃尾矿土壤微生物群落结构受土壤p H影响最为显著,而植被修复后土壤微生物群落的环境影响因子则转变为含水量、有机质、有机碳及总磷含量。进一步揭示了微生物在植被修复过程中所起到的重要作用,并为矿山生态重建过程中的土壤改良工作提供了丰富的理论依据。     

Invasive species cover,soil type,and grazing interact to predict long‐term grassland restoration success

Grasslands are undergoing tremendous degradation as a result of climate change, land use, and invasion by non‐native plants. However, understanding of the factors responsible for driving reestablishment of grassland plant communities is largely derived from short‐term studies. In order to develop an understanding of the factors responsible for longer term restoration outcomes in California annual grasslands, we surveyed 12 fields in Davis, CA, U.S.A., in 2015 that were seeded with native species mixtures starting in 2004. Using field surveys, we investigated how invasive plant richness and cover, native plant richness and cover, aboveground biomass, grazing, soil type, and restoration species identity might provide utility for explaining patterns of restoration success. We found a negative relationship between invasive cover and restoration cover, which was attributed to the slow establishment of seeded species and subsequent dominance by weeds. The relationship between invasive cover and restoration cover was modified by grazing, likely due to a change in the dominance of exotic forbs, which have a more similar growing season to restoration species, and therefore compete more strongly for late season moisture. Finally, we found that soil type was responsible for differences in the identity and abundance of invasive plants, subsequently affecting restoration cover. This work highlights the value of focusing resources on reducing invasive species cover, limiting grazing to periods of adequate moisture, and considering soil type for successful long‐term restoration in California annual grasslands. Moreover, observations of long‐term restoration outcomes can provide insight into the way mechanisms driving restoration outcomes might differ through time.     

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.     

Native soil microbial amendments generate trade‐offs in plant productivity,diversity, and soil stability in coastal dune restorations

Given the important role that soil microbes play in structuring plant communities and mediating ecosystem functions, there is growing interest in harnessing microbial communities to restore degraded ecosystems. Dune restorations, in particular, may benefit from native soil amendments because microbial diversity and abundance are very low in unvegetated areas. In an outdoor mesocosm experiment simulating Texas Gulf Coast dune restorations, we tested how native soil microbial amendments and restored diversity of foundational grasses influenced three key restoration responses: plant performance, plant diversity (including the colonization of native forbs), and soil stability. We found that native microbial amendments increased plant diversity and have the potential to increase soil stability, but this came at the cost of decreased plant biomass. Our results suggest that soil enemies in the native microbial amendments increased plant diversity by decreasing the performance of the dominant grass species and that arbuscular mycorrhizal fungi in the native microbial amendments increased the density of fungal hyphae in the soil, which can increase soil stability. Depending on the goals of the restoration, native soil microbial amendments may be a simple and inexpensive method to provide restoration benefits.     

Soils,time, and primate paleoenvironments

Soils are the skin of the earth. From both poles to the equator, wherever rocks or sediment are exposed at the surface, soils are forming through the physical and chemical action of climate and living organisms. The physical attributes (color, texture, thickness) and chemical makeup of soils vary considerably, depending on the composition of the parent material and other variables: temperature, rainfall and soil moisture, vegetation, soil fauna, and the length of time that soil–forming processes have been at work. United States soil scientists¹ have classified modern soils into ten major groups and numerous subgroups, each reflecting the composition and architecture of the soils and, to some extent, the processes that led to their formation. The physical and chemical processes of soil formation have been active throughout geologic time; the organic processes have been active at least since the Ordovician.² Consequently, nearly all sedimentary rocks that were deposited in nonmarine settings and exposed to the elements contain a record of ancient, buried soils or paleosols. A sequence of these rocks, such as most ancient fluvial (stream) deposits, provides a record of soil paleoenvironments through time. Paleosols are also repositories of the fossils of organisms (body fossils) and the traces of those organisms burrowing, food–seeking, and dwelling activities (ichnofossils). Indeed, most fossil primates are found in paleosols. Careful study of ancient soils gives new, valuable insights into the correct temporal reconstruction of the primate fossil record and the nature of primate paleoenvironments.     

Soil bioengineering and the ecological restoration of riverbanks at the Airport Town,Shanghai, China

Ecological, soil bioengineering, and traditional techniques were integrated to obtain a structurally sound, ecologically sustainable and socio-economically beneficial method for restoring the riverbanks at the Airport Town, Shanghai, which was the first project applying soil bioengineering to riverbank restoration in China. Soil bioengineering is the use of living plant materials to construct structures that perform some engineering and ecological functions and can provide an effective means for slope stabilization and site restoration of riverbanks. The restoration and management strategy was based on a plan to integrate the natural landscape using live staking, live fascines, brush layer, vegetated geo-grids and geo-gabions, along with native vegetation for riverbank preservation. Ecological parameters including root characteristics and their biomass, species and habitat diversity, and soil moisture and shear stress were measured for site characterization and evaluation of a demonstration project The riverbank erosion was reduced significantly, along with an increase in species and habitat diversity, and improvement in aesthetics and water quality after a ten-month project implementation period, when compared to the control site. Our project of ecological restoration of riverbanks can be viewed through the perspective of the 19 principles presented by Mitsch and Jorgensen, which shows also how the principles and methods of soil bioengineering, and the concepts of ecological engineering that have recently been much developed in the West have been absorbed into Chinese practices of ecological engineering and can be applied to ecological restoration of riverbanks in China.     

New Zealand island restoration: seabirds, predators, and the importance of history

New Zealand?s offshore and outlying islands have long been a focus of conservation biology as sites of local endemism and as last refuges for many species. During the c. 730 years since New Zealand has been settled by people, mammalian predators have invaded many islands and caused local and global extinctions. New Zealand has led international efforts in island restoration. By the late 1980s, translocations of threatened birds to predator-free islands were well under way to safeguard against extinction. Non-native herbivores and predators, such as goats and cats, had been eradicated from some islands. A significant development in island restoration in the mid-1980s was the eradication of rats from small forested islands. This eradication technology has been refined and currently at least 65 islands, including large and remote Campbell (11 216 ha) and Raoul (2938 ha) Islands, have been successfully cleared of rats. Many of New Zealand?s offshore islands, especially those without predatory mammals, are home to large numbers of breeding seabirds. Seabirds influence ecosystem processes on islands by enhancing soil fertility and through soil disturbance by burrowing. Predators, especially rats, alter ecosystem processes and cause population reductions or extinctions of native animals and plants. Islands have been promoted as touchstones of a primaeval New Zealand, but we are now increasingly aware that most islands have been substantially modified since human settlement of New Zealand. Archaeological and palaeoecological investigations, together with the acknowledgement that many islands have been important mahinga kai (sources of food) for Maori, have all led to a better understanding of how people have modified these islands. Restoration technology may have vaulted ahead of our ability to predict the ecosystem consequences of its application on islands. However, research is now being directed to help make better decisions about restoration and management of islands, decisions that take account of island history and key drivers of island ecosystem functioning.     

Soil microbial communities in restored and unrestored coastal dune ecosystems in California

Most restoration projects involving invasive plant eradication tend to focus on plant removal with little consideration given to how these invasives change soil microbial communities. However, soil microorganisms can determine invasibility of habitats and, in turn, be altered by invasives once established, potentially inhibiting native plant establishment. We studied soil microbial communities in coastal dunes with varying invasion intensity and different restoration approaches (herbicide, mechanical excavation) at Point Reyes National Seashore. Overall, we found evidence of a strong link between bacterial and fungal soil communities and the presence of invasives and restoration approach. Heavily invaded sites were characterized by a lower abundance of putatively identified nitrifiers, fermentative bacteria, fungal parasites, and fungal dung saprotrophs and a higher abundance of cellulolytic bacteria and a class of arbuscular mycorrhizal fungi (Archaeosporomycetes). Changes in soil microbiota did not fully dissipate following removal of invasives using herbicide, with exception of reductions in cellulolytic bacteria and Archaeosporomycetes abundance. Mechanical restoration effectively removed both invasives and soil legacy effects by inverting or “flipping” rhizome‐contaminated surface soils with soils from below and may have inadvertently induced other adverse effects on soils that impeded reestablishment of native dune plants. Land managers should consider additional measures to counteract lingering legacy effects and/or focus restoration efforts in areas where legacy effects are less pronounced.     

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.     

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.     

Contribution of nematodes to the structure and function of the soil food web

As carbon and energy flow through the soil food web they are depleted by the metabolic and production functions of organisms. To be sustained, a "long" food web, with a large biomass at higher trophic levels, must receive a high rate of rhizodeposition or detrital subsidy, or be top-populated by organisms of slow growth and long life cycle. Disturbed soil food webs tend to be bottom heavy and recalcitrant to restoration due to the slow growth of upper predator populations, physical and chemical constraints of the soil matrix, biological imbalances, and the relatively low mobility and invasion potential of soil organisms. The functional roles of nematodes, determined by their metabolic and behavioral activities, may be categorized as ecosystem services, disservices or effect-neutral. Among the disservices attributable to nematodes are overgrazing, which diminishes services of prey organisms, and plant-damaging herbivory, which reduces carbon fixation and availability to other organisms in the food web. Unfortunately, management to ameliorate potential disservices of certain nematodes results in unintended but long-lasting diminution of the services of others. Beneficial roles of nematodes may be enhanced by environmental stewardship that fosters greater biodiversity and, consequently, complementarity and continuity of their services.     

Rates of root and organism growth, soil conditions, and temporal and spatial development of the rhizosphere

BACKGROUND: Roots growing in soil encounter physical, chemical and biological environments that influence their rhizospheres and affect plant growth. Exudates from roots can stimulate or inhibit soil organisms that may release nutrients, infect the root, or modify plant growth via signals. These rhizosphere processes are poorly understood in field conditions. SCOPE AND AIMS: We characterize roots and their rhizospheres and rates of growth in units of distance and time so that interactions with soil organisms can be better understood in field conditions. We review: (1) distances between components of the soil, including dead roots remnant from previous plants, and the distances between new roots, their rhizospheres and soil components; (2) characteristic times (distance(2)/diffusivity) for solutes to travel distances between roots and responsive soil organisms; (3) rates of movement and growth of soil organisms; (4) rates of extension of roots, and how these relate to the rates of anatomical and biochemical ageing of root tissues and the development of the rhizosphere within the soil profile; and (5) numbers of micro-organisms in the rhizosphere and the dependence on the site of attachment to the growing tip. We consider temporal and spatial variation within the rhizosphere to understand the distribution of bacteria and fungi on roots in hard, unploughed soil, and the activities of organisms in the overlapping rhizospheres of living and dead roots clustered in gaps in most field soils. CONCLUSIONS: Rhizosphere distances, characteristic times for solute diffusion, and rates of root and organism growth must be considered to understand rhizosphere development. Many values used in our analysis were estimates. The paucity of reliable data underlines the rudimentary state of our knowledge of root-organism interactions in the field.