Biological Soil Crusts of the Great Plains: A Review

Biological soil crusts (BSCs), or biocrusts, are composed of fungi, bacteria, algae, and bryophytes (mosses, etc.) that occupy bare soil, entwining soil particles with filaments or rootlike structures and/or gluing them together with polysaccharide exudates to form a consolidated surface crust that stabilizes the soil against erosion. BSCs are common in arid and semiarid regions where vascular plant cover is naturally sparse, maximizing the exposure of surface-dwelling organisms to direct sunlight. Although less prominent and less studied there, BSC organisms are also present in more mesic areas such as the Great Plains where they can be found in shortgrass and mixed-grass prairie, in the badlands of several states, where burrowing animals have created patches of bare soil, on damaged road-cuts, strip-mines, gas and oil drill pads, military training areas, heavily grazed areas, and burn scars. Even where BSCs are not readily visible to the naked eye, many of the organisms are still present. BSC organisms are passively dispersed to the Great Plains as airborne organismal fragments, asexual diaspores, or sexual spores that accompany wind-blown dust from as far away as northern China and Mongolia. BSCs can best be studied and managed by 1) acknowledging their presence; 2) documenting their diversity, abundance, and functional roles; and 3) minimizing unnecessary disturbance, particularly when the soils are dry. This paper describes the current knowledge of Great Plains BSCs in an effort to heighten awareness of these cryptic but crucial ecosystem components and to encourage new research initiatives to better understand and manage them in this biome. Some specific actions may include refined taxonomic and ecologic studies of BSC organisms in underexplored areas, particularly those previously less or not recognized as BSC habitat, and incorporation of techniques to sample airborne organisms

Using an Ultraviolet Light Test to Improve Sagebrush Identification and Predict Forage Quality for Wildlife

Sagebrush identification can be improved by using a relatively easy ultraviolet (UV) light test on specimens. Sagebrush produces a variety of water-soluble polyphenols called coumarins, which fluoresce a blue color under UV light and can help differentiate species, subspecies, and hybrids. We tested 16 different sagebrush taxa (including species and subspecies) from herbarium specimens and found 3 taxa (low sagebrush, Artemisia arbuscula; Wyoming sagebrush, A. tridentata wyomingensis; and basin sagebrush, A. t. tridentata) that were often misidentified. We show that the UV light test can greatly improve identification of these species. Moreover, given that the UV+ chemicals that discriminate taxa are also considered an indirect biomarker of sagebrush palatability for some herbivores, the UV light test can be used to predict forage quality for threatened species like sage-grouse (Centrocercus spp.) and pygmy rabbits (Brachylagus idahoensis). Collecting voucher specimens of sagebrush at wildlife study sites and comparing their UV intensity to historical herbarium specimens could help identify both current and changing availability of palatable sagebrush for wildlife. We found that even herbarium specimens \u3e80 years old still fluoresce under UV light

Aspen Indicator Species in Lichen Communities in the Bear River Range of Idaho and Utah

Aspen are thought to be declining in this region due to a combination of fire suppression, grazing and wildlife management practices, and potentially cool/wet climates of the past century which favor advancing conifer succession. Many scientists are concerned that aspen’s related species may also be losing habitat, thereby threatening the long-term local and regional viability of this important community. To date, few studies have specifically examined the role of aspen’s epiphytic lichen community. This paper presents basic community research describing the application of Indicator Species Analysis for lichens growing on aspen stems in the central Rocky Mountains of North American. Results show unique lichen assemblages between conifers and aspen – the dominant hardwood of mid-elevations in this region

\u3cem\u3eBromus tectorum\u3c/em\u3e Litter Alters Photosynthetic Characteristics of Biological Soil Crusts from a Semiarid Shrubland

Invasion by the exotic annual grass Bromus tectorum has increased the cover and connectivity of fine litter in the sagebrush steppes of western North America. This litter tends to cover biological soil crusts, which could affect their metabolism and growth. To investigate this possible phenomenon, biological soil crusts dominated by either the moss Bryum argenteum or the lichen Diploschistes muscorum were covered with B.tectorum litter (litter treatment) or left uncovered (control treatment) and exposed to natural field conditions. After periods of five and ten months, we removed the litter and compared the photosynthetic performance of biological soil crusts from the two treatments. Litter induced photosynthetic changes in our samples. In bothB. argenteum and D. muscorum, biological soil crusts that had been covered with litter for ten months had lower rates of gross photosynthesis and lower chlorophyll content than control samples. Similarly in both biological soil crust types, litter reduced the rate of dark respiration. For D. muscorum, the reduction in dark respiration fully compensated for the decrease in gross photosynthesis, resulting in similar values of net photosynthesis in the two treatments. In contrast, for B. argenteum, net photosynthesis was four-times greater in the control than the litter treatment. Also under litter cover, D. muscorum showed three common adaptations to shade conditions: a decrease in the light compensation point, in the light intensity needed to achieve 95% of maximal net photosynthesis, and in the chlorophyll a/b ratio. None of these changes was apparent in B. argenteum. Overall, our results indicate that photosynthetic responses to the presence of litter varied among species of the crust biota and that the litter can reduce the photosynthetic capacity of biological soil crusts. These results help to explain field observations of decreases in biological soil crust cover and changes in biological soil crust composition with increases in litter cover, and suggest that the landscape-wide invasion by B. tectorum may have substantial effects on biological soil crust performance and therefore their capacity to function in semiarid shrublands

Could Biological Soil Crusts Act as Natural Fire Fuel Breaks in the Sagebrush Steppe?

For decades, large portions of the semi-arid sagebrush ecosystem have been experiencing increased frequency and extent of wildfire, even though small, infrequent fire is a natural disturbance in this ecosystem (Baker, 2006). Increased wildfire is threatening the existence of sagebrush ecosystems and the wildlife species that depend upon them (Baker, 2006; Coates et al., 2016). Increased wildfire in sagebrush ecosystems is often driven by invasive annual grasses, especially cheatgrass, Bromus tectorum (L.). Invasion can initiate a trajectory toward a “grass-fire cycle”, in which cheatgrass increases fine fuel loadings that promote fire, and native plant species do not recover quickly after fire, leading frequently burned sites to transition to monocultures of cheatgrass (Brooks et al., 2004). Although cheatgrass has been extensively studied in the sagebrush steppe, less attention has been given to the organisms that would have filled the interspaces that cheatgrass replaces, namely, biological soil crusts (“biocrusts”). Semi-arid environments are characterized by sparse cover of vascular plants and substantial cover of biocrusts (Belnap & Lange, 2001). Biocrusts contain organisms that live on the soil surface and include lichens, mosses, and light algal crusts (including cyanobacteria). Although biocrusts were included in some of the first descriptions of the vegetation in the region (Flowers, 1934), biocrusts are rarely included in contemporary studies of sagebrush ecosystems. Comprehensive community studies have concluded consistent negative relationships between abundance of biocrusts and annual invasive grasses, specifically cheatgrass (Condon & Pyke, 2018a,b; Daubenmire, 1970). We postulate that biocrusts, and particularly lichens, facilitate a pattern of small, infrequent, low intensity fire given their association with reduced fine fuels (cheatgrass)

Shotgun Sequencing Decades-Old Lichen Specimens to Resolve Phylogenomic Placement of Type Material

Natural history collections, including name-bearing type specimens, are an important source of genetic information. These data can be critical for appropriate taxonomic revisions in cases where the phylogenetic position of name-bearing type specimens needs to be identified, including morphologically cryptic lichen-forming fungal species. Here, we use high-throughput metagenomic shotgun sequencing to generate genome-scale data from decades-old (i.e., more than 30 years old) isotype specimens representing three vagrant taxa in the lichen-forming fungal genus Rhizoplaca, including one species and two subspecies. We also use data from high-throughput metagenomic shotgun sequencing to infer the phylogenetic position of an enigmatic collection, originally identified as R. haydenii, that failed to yield genetic data via Sanger sequencing. We were able to construct a 1.64 Mb alignment from over 1200 single-copy nuclear gene regions for the Rhizoplaca specimens. Phylogenomic reconstructions recovered an isotype representing Rhizoplaca haydenii subsp. arbuscula within a clade comprising other specimens identified as Rhizoplaca haydenii subsp. arbuscula, while an isotype of R. idahoensis was recovered within a clade with substantial phylogenetic substructure comprising Rhizoplaca haydenii subsp. haydenii and other specimens. Based on these data and morphological differences, Rhizoplaca haydenii subsp. arbuscula is elevated to specific rank as Rhizoplaca arbuscula. For the enigmatic collection, we were able to assemble the nearly complete nrDNA cistron and over 50 Mb of the mitochondrial genome. Using these data, we identified this specimen as a morphologically deviant form representing Xanthoparmelia aff. subcumberlandia. This study highlights the power of high-throughput metagenomic shotgun sequencing in generating larger and more comprehensive genetic data from taxonomically important herbarium specimens

A First Phylogenetic Assessment of \u3ci\u3eDictyonemo\u3c/i\u3e s.lat in Southwestern North America Reveals Three New Basidiolichens, Described in Honor James D. Lawrey

Three species of lichenized basidiomycetes in the Dictyonema clade from southeastern North America are described as new to science: Cyphellostereum georgianum, C. jamesianum and Dictyonema lawreyi, all with a crustose-filamentous growth form. Based on ITS sequences, the species form well-supported monophyletic clades in a phylogeny and are represented by at least two specimens each. They are also distinguishable by morphological and anatomical characters. These new findings emphasize the importance of lichenological studies in North America, especially in historically understudied taxonomic groups, such as basidiolichens. This study is dedicated to James D. Lawrey on the occasion of his 70th birthda

Biocrusts Indicators of Livestock Grazing Effects on Soil Stability in Sagebrush Steppe: A Case Study from a Long-Term Experiment in the Northern Great Basin

Biocrusts are sensitive to changes in livestock grazing intensity in arid rangelands and may be useful indicators of ecosystem functions, particularly soil properties like soil stability, which may suggest the potential for soil erosion. We compared biocrust community composition and surface soil stability in a big sagebrush (Artemisia tridentata) steppe rangeland in the northwestern Great Basin in several paired sites, with or without long-term cattle grazing exclusion, and similar soils (mostly sandy loams), climate, and vegetation composition. We found that livestock grazing was associated with both lower surface soil stability and cover of several biocrust morphogroups, especially lichens, compared with sites with long-term livestock exclusion. Surface soil stability did not modify the effects of grazing on most biocrust components via interactive effects. Livestock grazing effects on total biocrust cover were partially mediated by changes in surface soil stability. Though lichens were more sensitive to grazing disturbance, our results suggest that moss (mostly Tortula ruralis in this site) might be a more readily observable indicator of grazing-related soil stability change in this area due to their relatively higher abundance compared with lichens (moss: mean, 8.5% cover, maximum, 96.1%, lichens: mean, 1.0% cover, maximum, 14.1%). These results highlight the potential for biocrust components as sensitive indicators of change in soil-related ecosystem functions in sagebrush steppe rangelands. However, further research is needed to identify relevant indicator groups across the wide range of biocrust community composition associated with site environmental characteristics, variable grazing systems, other rangeland health metrics, and other disturbance types such as wildfire

Bromus tectorum litter alters photosynthetic characteristics of biological soil crusts from a semiarid shrubland

a b s t r a c t Invasion by the exotic annual grass Bromus tectorum has increased the cover and connectivity of fine litter in the sagebrush steppes of western North America. This litter tends to cover biological soil crusts, which could affect their metabolism and growth. To investigate this possible phenomenon, biological soil crusts dominated by either the moss Bryum argenteum or the lichen Diploschistes muscorum were covered with B. tectorum litter (litter treatment) or left uncovered (control treatment) and exposed to natural field conditions. After periods of five and ten months, we removed the litter and compared the photosynthetic performance of biological soil crusts from the two treatments. Litter induced photosynthetic changes in our samples. In both B. argenteum and D. muscorum, biological soil crusts that had been covered with litter for ten months had lower rates of gross photosynthesis and lower chlorophyll content than control samples. Similarly in both biological soil crust types, litter reduced the rate of dark respiration. For D. muscorum, the reduction in dark respiration fully compensated for the decrease in gross photosynthesis, resulting in similar values of net photosynthesis in the two treatments. In contrast, for B. argenteum, net photosynthesis was four-times greater in the control than the litter treatment. Also under litter cover, D. muscorum showed three common adaptations to shade conditions: a decrease in the light compensation point, in the light intensity needed to achieve 95% of maximal net photosynthesis, and in the chlorophyll a/b ratio. None of these changes was apparent in B. argenteum. Overall, our results indicate that photosynthetic responses to the presence of litter varied among species of the crust biota and that the litter can reduce the photosynthetic capacity of biological soil crusts. These results help to explain field observations of decreases in biological soil crust cover and changes in biological soil crust composition with increases in litter cover, and suggest that the landscape-wide invasion by B. tectorum may have substantial effects on biological soil crust performance and therefore their capacity to function in semiarid shrublands