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.     

Biological soil crust salvage for dryland restoration: an opportunity for natural resource restoration

Biocrusts' functional importance and vulnerability to disturbance have motivated consistent interest in biocrust restoration, as well as a recent increase in research to cultivate biocrusts in laboratory and greenhouse settings for use in ecological restoration. As part of a sustainable approach to developing biocrust restoration, we argue that a complementary step is to improve and accelerate methods for salvaging biocrusts that would otherwise be destroyed in a forthcoming disturbance. The increasing rate and scale of disturbance pressures in drylands where biocrusts flourish means that the supply of salvageable biocrust and demand for that material in restoration greatly exceed the present cultivable supply. In this article we describe the state of knowledge for biocrust salvage, present a simple set of steps for conducting a salvage harvest, discuss risks and benefits when considering using salvage, and suggest future research directions to facilitate scaling up biocrust restoration using salvaged material. A focus on the use of salvaged biocrust as a restoration source may prove an important step to improve ecological restoration in notoriously difficult to restore dryland ecosystems.     

Insolation and disturbance history drive biocrust biodiversity in Western Montana rangelands

Background and Aims

Biological soil crust (biocrust) communities, though common and important in the intermountain west, have received little research attention. There are gaps in understanding what influences biocrust species’ abundance and distributions in this ecoregion. Climatic, edaphic, topographic, and biotic forces, in addition to anthropogenic disturbance can all influence the biocrust.

Methods

We determined the relative influence of several possible environmental filters in biocrust communities of western Montana (USA) grasslands at two spatial scales. The larger scale exploited strong topographically-dictated climatic variation across >60km², while the smaller scale focused on differences among distinct microsites within ~700m² plots.

Results

We detected a total of 96 biocrust taxa, mostly lichens. Biocrust richness at each site ranged from 0 to 39 species, averaging 14 species. Insolation, aspect, and disturbance history were the strongest predictors of biocrust richness, abundance, and species turnover across the landscape; soil texture was influential for some biocrust community properties. Steep, north-facing slopes that receive longer periods of shade harbored higher diversity and cover of biocrust than south-facing sites. At a small scale, interspaces among native herbaceous communities supported the greatest diversity of biocrust species, but microsites under shrub canopies supported the greatest cover.

Conclusions

We found that, among the variables investigated, tillage, insolation, soil texture and the associated vegetation community were the most important drivers of biocrust abundance and species richness. This study can inform the practice of restoration and conservation, and also guide future work to improve predictions of biocrust properties.

     

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.     

Inoculation and habitat amelioration efforts in biological soil crust recovery vary by desert and soil texture

As dryland degradation continues, it is increasingly important to understand how to effectively restore biocrust communities. Potential techniques include the addition of biocrust inoculum to accelerate biocrust recovery. Enhanced erosion typical of degraded environments creates a challenge for these approaches, due to loss by wind or water and burial by saltating particles. To retain and protect added inoculum, the inclusion of habitat‐amelioration techniques can improve recovery rates. This study tested three different types of inoculum (field‐collected, greenhouse‐cultivated, and laboratory‐cultivated biocrust) coupled with two treatments to augment soil stability and ameliorate habitat limitations: soil surface polyacrylamide additions and installation of straw barriers. This was done across two deserts (Great Basin and Chihuahuan) and separated into generally coarse‐ or finer‐textured soils in each desert, with results monitored for 3 years (2015, 2016, 2017). While the inoculum type, coupled with habitat ameliorations, occasionally enhanced biocrust growth across years and treatments, in other cases, it made no difference compared to natural recovery rates. Rather, the desert location and soil texture groupings were the most prominent factors in determining recovery trajectories. Recovery proportions were similar in the finer‐textured sites in both the Great Basin and the Chihuahuan deserts, while the coarser‐textured site in the Great Basin did show some recovery over time and the Chihuahuan coarser‐textured site did not. This study demonstrates the importance of understanding site potential and identifying key limitations to biocrust recovery for successful restoration projects.     

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.     

Seven years of carbon dioxide enrichment, nitrogen fertilization and plant diversity influence arbuscular mycorrhizal fungi in a grassland ecosystem

? We tested the prediction that the abundance and diversity of arbuscular mycorrhizal (AM) fungi are influenced by resource availability and plant community composition by examining the joint effects of carbon dioxide (CO(2) ) enrichment, nitrogen (N) fertilization and plant diversity on AM fungi. ? We quantified AM fungal spores and extramatrical hyphae in 176 plots after 7 yr of treatment with all combinations of ambient or elevated CO(2) (368 or 560 ppm), with or without N fertilization (0 or 4 g Nm(-2) ), and one (monoculture) or 16 host plant species (polyculture) in the BioCON field experiment at Cedar Creek Ecosystem Science Reserve, Minnesota, USA. ? Extramatrical hyphal lengths were increased by CO(2) enrichment, whereas AM spore abundance decreased with N fertilization. Spore abundance, morphotype richness and extramatrical hyphal lengths were all greater in monoculture plots. A structural equation model showed AM fungal biovolume was most influenced by CO(2) enrichment, plant community composition and plant richness, whereas spore richness was most influenced by fungal biovolume, plant community composition and plant richness. ? Arbuscular mycorrhizal fungi responded to differences in host community and resource availability, suggesting that mycorrhizal functions, such as carbon sequestration and soil stability, will be affected by global change.     

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.     

Adapting mechanized vascular plant seed dispersal technologies to biocrust moss restoration

Biocrust restoration is an emerging field relevant to management of rangelands. Manual dispersal of biocrust is an effective approach, though there are few examples of biocrust restoration greater than a square meter, in part because specialized machinery has yet to be developed or adapted for dispersal across larger areas. Restoration with vascular plants is now conducted using a variety of equipment for work at small to large scales. We investigated the potential of two tractor draft implements, a rangeland seed drill and imprinter, to establish moss. We assessed three treatments: moss passed through a seed drill, manually broadcasted over imprinted soil, and manually broadcasted onto unaltered soil. We conducted these treatments at two sites that differed in management histories and vegetative cover. To use the seed drill to disperse moss we needed to amend moss materials to prevent jamming, which reduced application rate. Imprinted treatments established the most cover, a result we attribute to creation of favorable microsites for establishment. Broadcast methods established a comparable number of moss colonies to broadcast‐over‐imprinted soil, but less cover. Drill treatments were not effective by any metric, likely due to burial of moss. Final differences between our highest cover treatment and controls were positive, but small: 2.38% in imprinted versus 0.44% in controls. However, moss cover in imprinted areas increased during our two‐year experiment at this site, while controls did not. Future work should focus on integrating biocrust and vascular plant treatments and identifying equipment to efficiently achieve these goals.     

Improving field success of biocrust rehabilitation materials: hardening the organisms or softening the environment?

Drylands are a widely degraded biome characterized by low productivity and high abiotic stress. Biological soil crust (biocrust) inoculants hold promise as a rehabilitation material in drylands, useful for boosting ecosystem functions including stabilization of eroding soil surfaces. However, biocrust materials cultivated ex situ by humans inconsistently establish under field conditions. We tested two approaches aimed at improving field establishment of biocrust inoculum: exposing the organisms within the inoculum to abiotic stress in an attempt to harden them, and applying habitat ameliorations intended to reduce the stressfulness of the environment. We hypothesized that both approaches in concert would lead to the most consistent field establishment of biocrusts. Overall, addition of biocrust inoculum did enhance biocrust establishment over the 1.5‐year duration of the study but did not result in full recovery. Generally, hardened biocrust inoculum performed no better than inoculum that was not hardened, although one indicator (chlorophyll a) was enhanced by addition of hardened inoculum in some circumstances. Temporary irrigation was initially an effective habitat amelioration but had no effect on biocrust establishment by 1.5 years. In contrast, application of jute net to the soil surface promoted biocrust establishment both in synergy with and in the absence of inoculum addition. We hypothesize that jute net stabilizes the soil surface, reduces abiotic stress, and enhances resource availability, overcoming barriers to establishment of biocrusts. Currently, there is broad support for the efficacy of habitat amelioration approaches in biocrust rehabilitation, but effective hardening techniques remain elusive.