Lichens of soil crust communities in the Intermountain Area of the western United States

Lichens are common components of microbiotic soil crusts. A total of 34 species from 17 genera are reported from soil crust communities throughout the Intermountain Area. Distribution of terricolous lichens is determined by various physical and biological factors: physical and chemical characteristics of the soil, moisture regimes, temperature, insolation, and development and composition of the vascular plant community. Some species demonstrate a broad ecological amplitude while others have a more restricted distribution. All growth forms are represented; however, the vast majority of soil crust lichens are squamulose (minutely foliose). Fruticose species are least abundant. In exposed, middle-elevation sites vagrant (detached) species are common. This paper describes and discusses terricolous lichen communities of desert habitats of the intermountain western United States. Effects of various human-related activities including grazing, wildfire, air pollution, and recreation vehicles on soil crust lichens are discussed. Gypsoplaca macrophylla (Zahlbr.) Timdal, a rare squamulose lichen which occurs on gypsifersous soils, was recently collected in Emery County, Utah, and is reported as new to the state.     

Cryptogamic soil crusts: recovery from grazing near Camp Floyd State Park, Utah, USA

The effects of grazing on the cryptogamic and vascular plant communities at two sites near Camp Floyd State Park, Utah County, Utah, were studied. The grazed site was subject to heavy grazing up until seven years prior to the study. The ungrazed site within the park boundaries had been protected from grazing for 20 years prior to the study and had a well-developed algal-lichen-moss crust. We found that the algae of the grazed site had recovered in terms of degree of crusting. There were no significant differences in the algal communities of the two sites when prevalent species were used as blocks in the ANOVAR analysis. However, when major algal groups were used as blocks, the analysis was significant, with the more recently grazed site having lower algal frequency. This difference, together with a few compositional differences, indicates that, although the algal community seven years following grazing is very similar to the algal community free of grazing for 20 years, the seven-year site is still in the process of recovery and community development. The diatom collections had a higher density in the grazed site, though the difference was not significant. Recovery of the lichen and moss community was not complete. The lichen Collema tenax and the mosses Bryum pallescens and Tortula ruralis were all significantly more abundant in the ungrazed area. Total cover of the lichen and moss components of the soil crusts was significantly lower in the more recently grazed area. Vascular cover was also lower.     

Effect of cryptobiotic crust type on microarthropod assemblages in pi?on-juniper woodland in central New Mexico

Cryptobiotic crusts make up an important part of the ground cover in arid systems. Along with their roles of retarding soil erosion and enhancing soil fertility, crusts may also be supporting local and regional arthropod biodiversity. We inventoried arthropod species in mossy, lichen, and mixed (lichen and mossy) cryptobiotic crusts at 2 sites in central New Mexico pi?on-juniper habitat. We collected 240 crust samples and used a heptane flotation technique to extract the microarthropods. We found 39 species of microarthropods and small macroarthropods, with significantly fewer species on lichen. Species richness was higher in March than in August, but diversity was lower because of dominance by the mite Neonanorchestes sp. Mean area differed significantly for different crust types, but arthropod species richness did not follow the pattern of more species on crusts with larger area. Arthropod species may be influencing such processes as nutrient cycling; therefore, the crust/arthropod interaction may be critical to aridland health.     

Natural history of a saline mound ecosystem

Along the margins of playas in northwestern Nevada, a salt-tolerant plant community occupies mounds that dot a largely unvegetated landscape. In this environment we studied soil development and plant-soil relationships. The mounds, averaging 0.3 m in height, are occupied by the shrubs Allenrolfea occidentalis (iodine bush), Sacrobatus vermiculatus (black greasewood), and Atriplex lentiformis spp. torreyi (Torrey saltbush). Distichlis spicata (desert saltgrass) is the only herbaceous plant occupying this community. Soil salinity decreases with depth in this environment, and content of aqueous-extractable solutes is significantly influenced by site-specific vegetation. Content of silt, clay, and salt in mound surface horizons suggests a chronosequence of mound formation, with the youngest at the barren playa interface and the oldest at the upland vegetation border. Plant demography and mound soil stratigraphy suggest that a pulse of plant recruitment and mound building occurred during a time of neoglacial cooling. As a substrate for plant recruitment, mounds have a limited lifespan because deposition of eolian-transported salts and geochemical cycling by plants quickly render them too saline for seed germination. The apparent productivity of mound formation precludes definitive conclusions regarding those mound characteristics favorable for plant recruitment and survivorship.     

Small-stone content of Mima mounds of the Columbia Plateau and Rocky Mountain regions: implications for mound origin

Mima moundfields were investigated at the Lawrence Memorial Grassland Preserve, located on the Columbia Plateau in southern Wasco County, Oregon, and at three locations in the San Luis Valley and Sangre de Cristo Mountains, southern Colorado, to test the alternative hypotheses of mound origins by erosion, frost action, and soil translocation by geomyid pocket gophers. The concentrations of two size classes of small stones, gravel (8-15 mm diameter) and pebbles (15-50 mm diameter), were sampled along mound-to-intermound transects and at different depths within the mounds. Numbers and masses of small stones per unit soil volume increased from intermounds to mounds tops at the Colorado sites and from mound edge to mound top at the Oregon site, where thin intermound soils lay directly on the weathering surface of basalt bedrock. Numbers and masses of small stones in the surface soil of mound tops were greater than or similar to concentrations in deeper layers. Mean masses of individual pebbles were greater in the intermound zone than in mound soils at the Oregon site, but did not differ along mound-intermound gradient at the Oregon site and at one Colorado site, being highest at mound edges or in intermounds. These observations support the hypothesis that mounds are formed by centripetal translocation of soil by geomyid pocket gophers, and are contrary to predictions based on theories assuming erosion or frost action to be the mechanism of mound formation.     

Recovery rates of cryptobiotic crusts: inoculant use and assessment methods

Recovery rates of cyanobacterial-lichen soil crusts from disturbance were examined. Plots were either undisturbed or scalped, and scalped plots were either inoculated with surrounding biological crust material or left to recover naturally. Natural recovery rates were found to be very slow. Inoculation significantly hastened recovery for the cyanobacterial/green algal component, lichen cover, lichen species richness, and moss cover. Even with inoculation, however, lichen and moss recovery was minimal. Traditional techniques of assessing recovery visually were found to underestimate time for total recovery. Other techniques, such as extraction of chlorophyll a from surface soil and measurement of sheath material accumulation, were used and are discussed.     

Vegetation response to fire and postburn seeding treatments in juniper woodlands of the Grand Staircase-Escalante National Monument, Utah

We compared 3 naturally ignited burns with unburned sites in the Grand Staircase-Escalante National Monument. Each burn site was restored with native and nonnative seed mixes, restored with native seeds only, or regenerated naturally. In general, burned sites had significantly lower native species richness (1.8 vs. 2.9 species), native species cover (11% vs. 22.5%), and soil crust cover (4.1% vs. 15%) than unburned sites. Most burned plots, seeded or not, had significantly higher average nonnative species richness and cover and lower average native species richness and cover than unburned sites. Regression tree analyses suggest site variation was equally important to rehabilitation results as seeding treatments. Low native species richness and cover, high soil C, and low cover of biological soil crusts may facilitate increased nonnative species richness and cover. Our study also found that unburned sites in the region had equally high cover of nonnative species compared with the rest of the Monument. Cheatgrass ( Bromus tectorum ) dominated both burned and unburned sites. Despite the invasion of cheatgrass, unburned sites still maintain higher native species richness; however, the high cover of cheatgrass may increase fire frequency, further reduce native species richness and cover, and ultimately change vegetation composition in juniper woodlands.     

The biological soil crusts of the San Nicolas Island: enigmatic algae from a geographically isolated ecosystem

Composite soil samples from 7 sites on San Nicolas Island were evaluated quantitatively and qualitatively for the presence of cyanobacteria and eukaryotic microalgae. Combined data demonstrated a rich algal flora with 19 cyanobacterial and 19 eukaryotic microalgal genera being identified, for a total of 56 species. Nine new species were identified and described among the cyanobacteria and the eukaryotic microalgae that were isolated: Leibleinia edaphica, Aphanothece maritima, Chroococcidiopsis edaphica, Cyanosarcina atroveneta, Hassallia californica, Hassallia pseudoramosissima, Microchaete terrestre, Palmellopsis californicus, and Pseudotetracystis compactis. Distinct distributional patterns of algal taxa existed among sites on the island and among soil algal floras of western North America. Some algal taxa appeared to be widely distributed across many desert regions, including Microcoleus vaginatus, Nostoc punctiforme, Nostoc paludosum, and Tolypothrix distorta, Chlorella vulgaris, Diplosphaera cf. chodatii, Myrmecia astigmatica, Myrmecia biatorellae, Hantzschia amphioxys, and Luticola mutica. Some taxa share a distinctly southern distribution with soil algae from southern Arizona, southern California, and Baja California (e.g., Scenedesmus deserticola and Eustigmatos magnus ). The data presented herein support the view that the cyanobacterial and microalgal floras of soil crusts possess significant biodiversity, much of it previously undescribed.     

Habitat relationships of the blackbrush community ( Coleogyne ramosissima ) of southwestern Utah

Eight general study sites in the blackbrush ( Coleogyne ramosissima ) zone of southwestern Utah were examined. Soils data were gathered and plant cover was estimated. Shrubs were found to he positively correlated with shallow, sandy soil. Nonwoody plants were found to be positively correlated with deeper, silty soils. Cryptogamic soil crusts were positively correlated with silt and nitrogen in the soil and therefore may play a role in elevating soil fertility. Shrubs and grasses were negatively correlated. Management implications are discussed.         

Bryophytes on the calcareous soils of Mungo National Park, an arid area of southern central Australia

Bryophytes were found to be abundant as components of microbiotic soil crusts on the calcareous soils of Mungo National Park, an arid area of southern central Australia. Six sites that reflected differences in soils, topography, and vascular plant vegetation were studied. At each site bryophytes were abundant, both in terms of number of species present and percent ground cover. Number of species present did not differ significantly between sites, but percent bryophyte cover was lower at a site on sand dunes in mallee woodland and a site on a silcrete ridge than at the four other sites. Environmental factors (soil texture, pH, conductivity, nutrient status, vascular plant vegetation, light level, leaf litter, and fire frequency) appear to play a significant part in determining bryophyte distribution. Mosses that occur at Mungo are also widespread on calcareous substrates throughout arid southern Australia. Many of the bryophyte species present at Mungo also occur on limestones in high rainfall areas of eastern Australia. Environmental factors favoring bryophytic soil crusts in arid Australia are also present on limestones in high rainfall areas of eastern Australia and may account for tile presence of many arid zone bryophyte species on limestones. In Australia there appears to be a relationship between rainfall and the ratios of acrocarpous to pleurocarpous mosses, and thallose to leafy liverworts. Recognition of calcareous soils, widespread in arid areas of southern Australia, may be possible by assessing a combination of characteristics of bryophyte assemblages.     

Trampling disturbance and recovery of cryptogamic soil crusts in Grand Canyon National Park

Cryptogamic soil crusts in Grand Canyon National Park were trampled by hikers, under controlled conditions, to determine how rapidly they were pulverized and how rapidly they recovered. Only 15 trampling passes were required to destroy the structure of the crusts; visual evidence of bacteria and cryptogam cover was reduced to near zero after 50 passes. Soil crusts redeveloped in just one to three years, and after five years the extensive bacteria and cryptogam cover left little visual evidence of disturbance. Surface irregularity remained low after five years, however, suggesting that recovery was incomplete.     

Species associations of oribatid mites in lichens on the island of Ailsa Craig,Firth of Clyde (Acarei: Cryptostigmata)

Species associations of oribatid mites in maritime lichens on the island of Ailsa Craig, Firth of Clyde, were investigated. The thirteen lichen species examined yielded a total of seventeen oribatid species assignable to two discrete associations. One was characteristic of foliose lichens on exposed rocks and the other was found in terricolous fruticose lichen on acid healthland. Morphology of the lichen thallus and altitude of the collection site were found to influence the abundance, occurrence and species diversity of the mites.     

Selection of microhabitat by the red-backed vole, Clethrionomys gapperi

Clethrionomys gapperi were captured in microhabitats with greater densities of overall cover than at noncapture or random sites within the study area. Variables describing cover density and distance from free water were selected in a discriminant function analysis to differentiate between vole capture and noncapture sites. Vole capture sites had greater amounts of cover within 4 dm above ground surface and were further from standing water than noncapture sites. The preferential use by C. gapperi of microhabitats with greater densities of cover is in agreement with laboratory and field assessments of habitat use reported in the literature.     

Predator-prey interactions between the digger wasp Clypeadon laticinctus and the harvester ant Pogonomyrmex occidentalis

Females of the digger wasp Clypeadon laticinctus prey exclusively on workers of the western harvester ant Pogonomyrmex occidentalis. There is considerable variability in the size of ant workers both within and between ant mounds, and wasps tend to select larger workers as prey. Worker ants were most likely to be outside the nest at surface temperatures of 25–35°C. Wasps were seen at ant mounds most frequently when surface temperatures were between 40 and 50°C. Wasps captured ants outside and inside ant nests. Predation attempts were equally likely to result in a successful capture in either location, but entering the ant nests was more time consuming. Approximately 63% of visits by wasps to ant mounds resulted in a successful prey capture. In general, ant mounds that were visually conspicuous or close to wasp nest aggregations were more likely to be visited by wasps. However, frequency of visitation and rates of predation at a given mound varied considerably from week to week. Agonistic interactions between female wasps involving chases and occasional fights were likely to occur whenever two wasps were simultaneously present at an ant mound, especially when the only access to prey was by entering the mound.     

Influence of cryptogamic crusts on moisture relationships of soils in Navajo National Monument, Arizona

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Community structure of Eleodes beetles (Coleoptera: Tenebrionidae) in the shortgrass steppe: scale-dependent uses of heterogeneity

Patterns in the community structure of darkling beetle (9 Eleodes spp., Coleoptera: Tenebrionidae) assemblages in the shortgrass steppe of north central Colorado were monitored by live pitfall trapping for 4 summers. There were significant correlations among weather (temperature and precipitation), species richness, and number of individuals per species captured; effects from weather conditions also displayed 1-month and 1-yr delayed effects. Population densities of 2 eleodid species were monitored by mark-recapture methods. Densities of these species varied relatively little among years and sites, although density was correlated with temperature and precipitation. Abiotic influences on both density and richness differed between 2 macrohabitat types (shortgrass upland, shrub floodplain). The 4 largest species were most abundant in the floodplain, whereas the smallest species was most common in the upland. Affinities with cactus and shrub microhabitats (and an avoidance of bare soil) were evident, although a preference for shaded microhabitats was not detected. These results do not conform well to previous explanations of why darkling beetle assemblages are spatially and temporally heterogeneous, which primarily focused on predation and thermoregulation. Therefore, an alternative mechanism concerning scale-dependent uses of heterogeneity and mobility is proposed to account for eleodid community patterns.     

Form and dispersion of Mima mounds in relation to slope steepness and aspect on the Columbia Plateau

Patterned ground consisting of Mima-type earth mounds and associated sorted stone circles and nets is widespread on the Columbia Plateau of western North America. Studies of the geometric relationships of mounds and stone nets to slope aspect and steepness were carried out at the Lawrence Memorial Grassland Preserve, north central Oregon in June 1987. Mound and moundfield characteristics were sampled on randomly chosen 1-ha plots on slopes of different aspect and steepness. Mounds were largest, most circular and symmetrical in form, and most fully encircled by beds of size-sorted stones on level sites. On slopes of increasing steepness, mounds decreased in size, showed increasing asymmetry and downslope elongation, and became connected into lines oriented up- and downslope. Encircling stone beds became more weakly developed or disappeared on upslope and downslope sides of the mounds, and the lateral beds developed downslope extensions that eventually merged with those of adjacent upslope and downslope mounds. These patterns are interpreted as reflecting changes in the manner of soil translocation by northern pocket gophers, Thomomys talpoides , due to their responses to tunneling on slopes and to the modification of the flow of water across the slope because of the presence of mounds.     

Site characteristics and habitat requirements of the endangered dwarf bear-claw poppy ( Arctomecon humilis Coville, Papaveraceae)

Arctomecon humilis Cov. is a narrow endemic, confined to gypsiferous substrates derived from the Shnabkaib Member of the Moenkopi Formation in southwestern Utah. The characteristics of seven A. humilis sites were studied to define habitat requirements of the species. Both physical and biotic aspects of the habitat were analyzed; geology, soil chemistry, and physical properties, as well as vascular and nonvascular plant communities, were studied. Chemical and physical properties of these soils vary considerably from those supporting adjacent desert-shrub communities. The dominant vascular species on A. humilis sites are shadscale and Mormon tea. It was found that A. humilis occurred in portions of the habitat where shrubs were relatively less dense. A soil-surface, cryptogamic community contributed 84% or more of the total living cover on sites that supported A. humilis . Species composition of the cryptogamic community was highly similar among sites. Likewise, composition of the cryptogamic cover was similar when random samples were compared with samples centered on A. humilis plants. The Purgatory Flat site, which does not support the poppy, seems inseparable from A. humilis sites, in respect to soil characteristics and composition of the associated plant cover. There is reason to believe an A. humilis population could be established at the Purgatory Flat site.