Biological soil crusts across disturbance­recovery scenarios: effect of grazing regime on community dynamics.

Grazing represents one of the most common disturbances in drylands worldwide, affecting both ecosystem structure and functioning. Despite the efforts to understand the nature and magnitude of grazing effects on ecosystem components and processes, contrasting results continue to arise. This is particularly remarkable for the biological soil crust (BSC) communities (i.e., cyanobacteria, lichens, and bryophytes), which play an important role in soil dynamics. Here we evaluated simultaneously the effect of grazing impact on BSC communities (resistance) and recovery after livestock exclusion (resilience) in a semiarid grassland of Central Mexico. In particular, we examined BSC species distribution, species richness, taxonomical group cover (i.e., cyanobacteria, lichen, bryophyte), and composition along a disturbance gradient with different grazing regimes (low, medium, high impact) and along a recovery gradient with differently aged livestock exclosures (short-, medium-, long-term exclusion). Differences in grazing impact and time of recovery from grazing both resulted in slight changes in species richness; however, there were pronounced shifts in species composition and group cover. We found we could distinguish four highly diverse and dynamic BSC species groups: (1) species with high resistance and resilience to grazing, (2) species with high resistance but low resilience, (3) species with low resistance but high resilience, and (4) species with low resistance and resilience. While disturbance resulted in a novel diversity configuration, which may profoundly affect ecosystem functioning, we observed that 10 years of disturbance removal did not lead to the ecosystem structure found after 27 years of recovery. These findings are an important contribution to our understanding of BCS dynamics from a species and community perspective placed in a land use change context.

Bromus tectorum invasion alters nitrogen dynamics in an undisturbed arid grassland ecosystem.

The nonnative annual grass Bromus tectorum has successfully replaced native vegetation in many arid and semiarid ecosystems. Initial introductions accompanied grazing and agriculture, making it difficult to separate the effects of invasion from physical disturbance. This study examined N dynamics in two recently invaded, undisturbed vegetation associations (C3 and C4). The response of these communities was compared to an invaded/ disturbed grassland. The invaded/disturbed communities had higher surface NH4+ input in spring, whereas there were no differences for surface input of NO3-. Soil inorganic N was dominated by NH4+, but invaded sites had greater subsurface soil NO3-. Invaded sites had greater total soil N at the surface four years post-invasion in undisturbed communities, but total N was lower in the invaded/disturbed communities. Soil delta15N increased with depth in the noninvaded and recently invaded communities, whereas the invaded/disturbed communities exhibited the opposite pattern. Enriched foliar delta15N values suggest that Bromus assimilated subsurface NO3-, whereas the native grasses were restricted to surface N. A Rayleigh distillation model accurately described decomposition patterns in the noninvaded communities where soil N loss is accompanied by increasing soil delta15N; however, the invaded/ disturbed communities exhibited the opposite pattern, suggesting redistribution of N within the soil profile. This study suggests that invasion has altered the mechanisms driving nitrogen dynamics. Bromus litter decomposition and soil NO3- concentrations were greater in the invaded communities during periods of ample precipitation, and NO3- leached from the surface litter, where it was assimilated by Bromus. The primary source of N input in these communities is a biological soil crust that is removed with disturbance, and the lack of N input by the biological soil crust did not balance N loss, resulting in reduced total N in the invaded/disturbed communities. Bromus produced a positive feedback loop by leaching NO3- from decomposing Bromus litter to subsurface soil layers, accessing that deepsoil N pool with deep roots and returning that N to the surface as biomass and subsequent litter. Lack of new inputs combined with continued loss will result in lower total soil N, evidenced by the lower total soil N in the invaded/disturbed communities.

Small-scale vertical distribution of bacterial biomass and diversity in biological soil crusts from arid lands in the Colorado plateau.

We characterized, at millimeter resolution, bacterial biomass, diversity, and vertical stratification of biological soil crusts in arid lands from the Colorado Plateau. Microscopic counts, extractable DNA, and plate counts of viable aerobic copiotrophs (VAC) revealed that the top centimeter of crusted soils contained atypically large bacterial populations, tenfold larger than those in uncrusted, deeper soils. The plate counts were not always consistent with more direct estimates of microbial biomass. Bacterial populations peaked at the immediate subsurface (1-2 mm) in light-appearing, young crusts, and at the surface (0-1 mm) in well-developed, dark crusts, which corresponds to the location of cyanobacterial populations. Bacterial abundance decreased with depth below these horizons. Spatially resolved DGGE fingerprints of Bacterial 16S rRNA genes demonstrated the presence of highly diverse natural communities, but we could detect neither trends with depth in bacterial richness or diversity, nor a difference in diversity indices between crust types. Fingerprints, however, revealed the presence of marked stratification in the structure of the microbial communities, probably a result of vertical gradients in physicochemical parameters. Sequencing and phylogenetic analyses indicated that most of the naturally occurring bacteria are novel types, with low sequence similarity (83-93%) to those available in public databases. DGGE analyses of the VAC populations indicated communities of lower diversity, with most types having sequences more than 94% similar to those in public databases. Our study indicates that soil crusts represent small-scale mantles of fertility in arid ecosystems, harboring vertically structured, little-known bacterial populations that are not well represented by standard cultivation methods.

Temporal variation in community composition, pigmentation, and F(v)/F(m) of desert cyanobacterial soil crusts.

Summers on the Colorado Plateau (USA) are typified by harsh conditions such as high temperatures, brief soil hydration periods, and high UV and visible radiation. We investigated whether community composition, physiological status, and pigmentation might vary in biological soil crusts as a result of such conditions. Representative surface cores were sampled at the ENE, WSW, and top microaspects of 20 individual soil crust pedicels at a single site in Canyonlands National Park, Utah, in spring and fall of 1999. Frequency of cyanobacterial taxa, pigment concentrations, and dark adapted quantum yield [F(v)/F(m)] were measured for each core. The frequency of major cyanobacterial taxa was lower in the fall compared to spring. The less-pigmented cyanobacterium Microcoleus vaginatus showed significant mortality when not in the presence of Nostoc spp. and Scytonema myochrous (Dillw.) Agardh. (both synthesizers of UV radiation-linked pigments) but had little or no mortality when these species were abundant. We hypothesize that the sunscreen pigments produced by Nostoc and Scytonema in the surface of crusts protect other, less-pigmented taxa. When fall and spring samples were compared, overall cyanobacterial frequency was lower in fall, while sunscreen pigment concentrations, chlorophyll a concentration, and F(v)/F(m) were higher in fall. The ratio of cyanobacterial frequency/chlorophyll a concentrations was 2-3 times lower in fall than spring. Because chlorophyll a is commonly used as a surrogate measure of soil cyanobacterial biomass, these results indicate that seasonality needs to be taken into consideration. In the fall sample, most pigments associated with UV radiation protection or repair were at their highest concentrations on pedicel tops and WSW microaspects, and at their lowest concentrations on ENE microaspects. We suggest that differential pigment concentrations between microaspects are induced by varying UV radiation dosage at the soil surface on these different microaspects.

Aeolian dust in Colorado Plateau soils: nutrient inputs and recent change in source.

Aeolian dust (windblown silt and clay) is an important component in arid-land ecosystems because it may contribute to soil formation and furnish essential nutrients. Few geologic surfaces, however, have been characterized with respect to dust-accumulation history and resultant nutrient enrichment. We have developed a combination of methods to identify the presence of aeolian dust in arid regions and to evaluate the roles of this dust in ecosystem processes. Unconsolidated sandy sediment on isolated surfaces in the Canyonlands region of the Colorado Plateau differs greatly in mineralogical and chemical composition from associated bedrock, mainly aeolian sandstone. Detrital magnetite in the surficial deposits produces moderately high values of magnetic susceptibility, but magnetite is absent in nearby bedrock. A component of the surficial deposits must be aeolian to account for the abundance of magnetite, which formed originally in far-distant igneous rocks. Particle-size analysis suggests that the aeolian dust component is typically as much as 20-30%. Dust inputs have enriched the sediments in many elements, including P, Mg, Na, K, and Mo, as well as Ca, at sites where bedrock lacks calcite cement. Soil-surface biologic crusts are effective dust traps that apparently record a change in dust sources over the past several decades. Some of the recently fallen dust may result from human disturbance of land surfaces that are far from the Canyonlands, such as the Mojave Desert. Some land-use practices in the study area have the potential to deplete soil fertility by means of wind-erosion removal of aeolian silt.

ENVIRONMENTAL AUDITING: Choosing Indicators of Natural Resource Condition: A Case Study in Arches National Park, Utah, USA

/ Heavy visitor use in many areas of the world have necessitated development of ways to assess visitation impacts. Arches National Park recently completed a Visitor Experience and Resource Protection (VERP) plan. Integral to this plan was developing a method to identify biological indicators that would both measure visitor impacts and response to management actions. The process used in Arches for indicator selection is outlined here as a model applicableto many areas facing similar challenges. The steps were: (1) Vegetation types most used by visitors were identified. Impacted and unimpacted areas in these types were sampled, comparing vegetation and soil factors. (2) Variables found to differ significantly between compared sites were used as potential indicators. (3) Site-specific criteria for indicators were developed, and potential indicators evaluated using these criteria. (4) Chosen indicators were further researched for ecological relevancy. (5) Final indicators were chosen, field tested, and monitoring sites designated. In Arches, indicators were chosen for monitoring annually (soil crust index, soil compaction, number of used social trails and soil aggregate stability) and every five years (vegetation cover and frequency; ground cover; soil chemistry; and plant tissue chemistry).KEY WORDS: Biological indicators; Recreation impacts; Recreation management; Desert soils; Trampling

Surface disturbances: Their role in accelerating desertification.

Maintaining soil stability and normal water and nutrient cycles in desert systems is critical to avoiding desertification. These particular ecosystem processes are threatened by trampling of livestock and people, and by off-road vehicle use. Soil compaction and disruption of cryptobiotic soil surfaces (composed of cyanobacteria, lichens, and mosses) can result in decreased water availability to vascular plants through decreased water infiltration and increased albedo with possible decreased precipitation. Surface disturbance may also cause accelerated soil loss through wind and water erosion and decreased diversity and abundance of soil biota. In addition, nutrient cycles can be altered through lowered nitrogen and carbon inputs and slowed decomposition of soil organic matter, resulting in lower nutrient levels in associated vascular plants. Some cold desert systems may be especially susceptible to these disruptions due to the paucity of surface-rooting vascular plants for soil stabilization, fewer nitrogen-fixing higher plants, and lower soil temperatures, which slow nutrient cycles. Desert soils may recover slowly from surface disturbances, resulting in increased vulnerability to desertification. Recovery from compaction and decreased soil stability is estimated to take several hundred years. Re-establishment rates for soil bacterial and fungal populations are not known. The nitrogen fixation capability of soil requires at least 50 years to recover. Recovery of crusts can be hampered by large amounts of moving sediment, and re-establishment can be extremely difficult in some areas. Given the sensitivity of these resources and slow recovery times, desertification threatens million of hectares of semiarid lands in the United States.