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J. Belnap

Publications and source records attributed to J. Belnap.

At least 37 records · Page 2Linked to original sources

Do soil characteristics or microhabitat determine field emergence and success of Bromus tectorum?

In southeastern Utah, Bromus tectorum occurs where Hilaria jamesii is dominant and rarely where Stipa hymenoides/S. comata dominate. To determine whether this distribution is due to soil characteristics or microhabitat, we transplanted H. jamesii soil to a Stipa site and vice versa during a severe drought (2001) and a wetter year (2002). Additionally, we planted B. tectorum under H. jamesii and Stipa canopies, with or without H. jamesii litter, and with or without herbivory. Bromus tectorum emergence and biomass in reciprocal transplants were similar at both sites; there were no site differences for all microhabitat treatments. Being under a plant canopy increased emergence in 2001 and decreased survival during 2002. Herbivory decreased emergence in 2001 and decreased survival during 2002. Litter increased emergence only under the canopy in 2001 but did not affect survival in 2002. Survival in 2001 was so low that biomass was unattainable; no microhabitat treatments affected biomass in 2002. We found that soil characteristics and microhabitat affected B. tectorum similarly in H. jamesii and Stipa patches, suggesting that these factors do not explain the association between B. tectorum and H. jamesii. However, these relationships may change during wet years when B. tectorum invasions most often occur. ?? 2006 Elsevier Ltd. All rights reserved.

Journal of Arid Environments

Heterogeneity of soil nutrients and subsurface biota in a dryland ecosystem

Dryland ecosystems have long been considered to have a highly heterogeneous distribution of nutrients and soil biota, with greater concentrations of both in soils under plants relative to interspace soils. We examined the distribution of soil resources in two plant communities (dominated by either the shrub Coleogyne ramosissima or the grass Stipa hymenoides) at two locations. Interspace soils were covered either by early successional biological soil crusts (BSCs) or by later successional BSCs (dominated by nitrogen (N)-fixing cyanobacteria and lichens). For each of the 8 plant type??crust type??locations, we sampled the stem, dripline, and 3 interspace distances around each of 3 plants. Soil analyses revealed that only available potassium (Kav) and ammonium concentrations were consistently greater under plants (7 of 8 sites and 6 of 8 sites, respectively). Nitrate and iron (Fe) were greater under plants at 4 sites, while all other nutrients were greater under plants at less than 50% of the sites. In contrast, calcium, copper, clay, phosphorus (P), and zinc were often greater in the interspace than under the plants. Soil microbial biomass was always greater under the plant compared to the interspace. The community composition of N-fixing bacteria was highly variable, with no distinguishable patterns among microsites. Bacterivorous nematodes and rotifers were consistently more abundant under plants (8 and 7 sites, respectively), and fungivorous and omnivorous nematodes were greater under plants at 5 of the 8 sites. Abundance of other soil biota was greater under plants at less than 50% of the sites, but highly correlated with the availability of N, P, Kav, and Fe. Unlike other ecosystems, the soil biota was only infrequently correlated with organic matter. Lack of plant-driven heterogeneity in soils of this ecosystem is likely due to (1) interspace soils covered with BSCs, (2) little incorporation of above-ground plant litter into soils, and/or (3) root deployment patterns. ?? 2007 Elsevier Ltd. All rights reserved.

Soil Biology and Biochemistry

Three distinct clades of cultured heterocystous cyanobacteria constitute the dominant N2-fixing members of biological soil crusts of the Colorado Plateau, USA

The identity of the numerically dominant N2-fixing bacteria in biological soil crusts of the Colorado Plateau region and two outlying areas was determined using multiple approaches, to link the environmental diversity of nifH gene sequences to cultured bacterial isolates from the regions. Of the nifH sequence-types detected in soil crusts of the Colorado Plateau, 89% (421/473) were most closely related to nifH signature sequences from cyanobacteria of the order Nostocales. N2-fixing cyanobacterial strains were cultured from crusts and their morphotypes, 16S rRNA gene and nifH gene sequences were characterized. The numerically dominant diazotrophs in the Colorado Plateau crusts fell within three clades of heterocystous cyanobacteria. Two clades are well-represented by phylogenetically and morphologically coherent strains, corresponding to the descriptions of Nostoc commune and Scytonema hyalinum, which are widely recognized as important N2-fixing components of soil crusts. A third, previously-overlooked clade was represented by a phylogenetically coherent but morphologically diverse group of strains that encompass the morphogenera Tolypothrix and Spirirestis. Many of the strains in each of these groups contained at least two nifH copies that represent different clusters in the nifH environmental survey. ?? 2007 Federation of European Microbiological Societies.

FEMS Microbiology Ecology

Effects of altered temperature and precipitation on desert protozoa associated with biological soil crusts

Biological soil crusts are diverse assemblages of bacteria, cyanobacteria, algae, fungi, lichens, and mosses that cover much of arid land soils. The objective of this study was to quantify protozoa associated with biological soil crusts and test the response of protozoa to increased temperature and precipitation as is predicted by some global climate models. Protozoa were more abundant when associated with cyanobacteria/lichen crusts than with cyanobacteria crusts alone. Amoebae, flagellates, and ciliates originating from the Colorado Plateau desert (cool desert, primarily winter precipitation) declined 50-, 10-, and 100-fold, respectively, when moved in field mesocosms to the Chihuahuan Desert (hot desert, primarily summer rain). However, this was not observed in protozoa collected from the Chihuahuan Desert and moved to the Sonoran desert (hot desert, also summer rain, but warmer than Chihuahuan Desert). Protozoa in culture began to encyst at 37??C. Cysts survived the upper end of daily temperatures (37-55??C), and could be stimulated to excyst if temperatures were reduced to 15??C or lower. Results from this study suggest that cool desert protozoa are influenced negatively by increased summer precipitation during excessive summer temperatures, and that desert protozoa may be adapted to a specific desert's temperature and precipitation regime. ?? 2006 by the International Society of Protistologists.

Journal of Eukaryotic Microbiology

The potential roles of biological soil crusts in dryland hydrologic cycles

Biological soil crusts (BSCs) are the dominant living cover in many drylands of the world. They possess many features that can influence different aspects of local hydrologic cycles, including soil porosity, absorptivity, roughness, aggregate stability, texture, pore formation, and water retention. The influence of biological soil crusts on these factors depends on their internal and external structure, which varies with climate, soil, and disturbance history. This paper presents the different types of biological soil crusts, discusses how crust type likely influences various aspects of the hydrologic cycle, and reviews what is known and not known about the influence of biological crusts on sediment production and water infiltration versus runoff in various drylands around the world. Most studies examining the effect of biological soil crusts on local hydrology are done by comparing undisturbed sites with those recently disturbed by the researchers. Unfortunately, this greatly complicates interpretation of the results. Applied disturbances alter many soil features such as soil texture, roughness, aggregate stability, physical crusting, porosity, and bulk density in ways that would not necessarily be the same if crusts were not naturally present. Combined, these studies show little agreement on how biological crusts affect water infiltration or runoff. However, when studies are separated by biological crust type and utilize naturally occurring differences among these types, results indicate that biological crusts in hyperarid regions reduce infiltration and increase runoff, have mixed effects in and regions, and increase infiltration and reduce runoff in semiarid cool and cold drylands. However, more studies are needed before broad generalizations can be made on how biological crusts affect infiltration and runoff. We especially need studies that control for sub-surface soil features such as bulk density, micro- and macropores, and biological crust structure. Unlike the mixed effects of biological crusts on infiltration and runoff among regions, almost all studies show that biological crusts reduce sediment production, regardless of crust or dryland type.

Hydrological Processes

Wind erodibility of soils at Fort Irwin, California (Mojave Desert), USA, before and after trampling disturbance: Implications for land management

Recently disturbed and 'control' (i.e. less recently disturbed) soils in the Mojave Desert were compared for their vulnerability to wind erosion, using a wind tunnel, before and after being experimentally trampled. Before trampling, control sites had greater cyanobacterial biomass, soil surface stability, threshold friction velocities (TFV, i.e. the wind speed required to move soil particles), and sediment yield than sites that had been more recently disturbed by military manoeuvres. After trampling, all sites showed a large drop in TFVs and a concomitant increase in sediment yield. Simple correlation analyses showed that the decline in TFVs and the rise in sediment yield were significantly related to cyanobacterial biomass (as indicated by soil chlorophyll a). However, chlorophyll a amounts were very low compared to chlorophyll a amounts found at cooler desert sites, where chlorophyll a is often the most important factor in determining TFV and sediment yield. Multiple regression analyses showed that other factors at Fort Irwin were more important than cyanobacterial biomass in determining the overall site susceptibility to wind erosion. These factors included soil texture (especially the fine, medium and coarse sand fractions), rock cover, and the inherent stability of the soil (as indicated by subsurface soil stability tests). Thus, our results indicate that there is a threshold of biomass below which cyanobacterial crusts are not the dominant factor in soil vulnerability to wind erosion. Most undisturbed soil surfaces in the Mojave Desert region produce very little sediment, but even moderate disturbance increases soil loss from these sites. Because current weathering rates and dust inputs are very low, soil formation rates are low as well. Therefore, soil loss in this region is likely to have long-term effects.

Earth Surface Processes and Landforms

Correlates of biological soil crust abundance across a continuum of spatial scales: Support for a hierarchical conceptual model

1. Desertification negatively impacts a large proportion of the global human population and > 30% of the terrestrial land surface. Better methods are needed to detect areas that are at risk of desertification and to ameliorate desertified areas. Biological soil crusts are an important soil lichen-moss-microbial community that can be used toward these goals, as (i) bioindicators of desertification damage and (ii) promoters of soil stability and fertility. 2. We identified environmental factors that correlate with soil crust occurrence on the landscape and might be manipulated to assist recovery of soil crusts in degraded areas. We conducted three studies on the Colorado Plateau, USA, to investigate the hypotheses that soil fertility [particularly phosphorus (P), manganese (Mn) and zinc (Zn)] and/or moisture limit soil crust lichens and mosses at four spatial scales. 3. In support of the soil fertility hypothesis, we found that lichen-moss crusts were positively correlated with several nutrients [Mn, Zn, potassium (K) and magnesium (Mg) were most consistent] at three of four spatial scales ranging from 3.5 cm2 in area to c. 800 km2. In contrast, P was negatively correlated with lichen-moss crusts at three scales. 4. Community composition varied with micro-aspect on ridges in the soil crust. Three micro-aspects [north-north-west (NNW), east-north-east (ENE) and TOP] supported greater lichen and moss cover than the warmer, windward and more xeric micro-aspects [west-south-west (WSW) and south-south-east (SSE)]. This pattern was poorly related to soil fertility; rather, it was consistent with the moisture limitation hypothesis. 5. Synthesis and application. Use of crusts as desertification bioindicators requires knowledge of a site's potential for crust cover in the absence of desertification. We present a multi-scale model of crust potential as a function of site properties. Future quantitative studies can use this model to guide sampling efforts. Also, our results suggest new directions in restoration research: enhancement of moisture residence time and fertilization with key nutrients (Mn, Zn, K and Mg). Re-establishment of soil crusts in desertified lands will help regain lost soil stability and fertility, and facilitate plant re-establishment. ?? 2006 British Ecological Society.

Journal of Applied Ecology

Direct effects of soil amendments on field emergence and growth of the invasive annual grass Bromus tectorum L. and the native perennial grass Hilaria jamesii (Torr.) Benth

Bromus tectorum L. is a non-native, annual grass that has invaded western North America. In SE Utah, B. tectorum generally occurs in grasslands dominated by the native perennial grass, Hilaria jamesii (Torr.) Benth. and rarely where the natives Stipa hymenoides Roem. and Schult. and S. comata Trin. & Rupr. are dominant. This patchy invasion is likely due to differences in soil chemistry. Previous laboratory experiments investigated using soil amendments that would allow B. tectorum to germinate but would reduce B. tectorum emergence without affecting H. jamesii. For this study we selected the most successful treatments (CaCl2, MgCl2, NaCl and zeolite) from a previous laboratory study and applied them in the field in two different years at B. tectorum-dominated field sites. All amendments except the lowest level of CaCl2 and zeolite negatively affected B. tectorum emergence and/or biomass. No amendments negatively affected the biomass of H. jamesii but NaCl reduced emergence. Amendment effectiveness depended on year of application and the length of time since application. The medium concentration of zeolite had the strongest negative effect on B. tectorum with little effect on H. jamesii. We conducted a laboratory experiment to determine why zeolite was effective and found it released large amounts of Na+, adsorbed Ca2+, and increased Zn2+, Fe2+, Mn2+, Cu2+, exchangeable Mg2+, exchangeable K, and NH 4+ in the soil. Our results suggest several possible amendments to control B. tectorum. However, variability in effectiveness due to abiotic factors such as precipitation and soil type must be accounted for when establishing management plans. ?? Springer 2006.

Plant and Soil

Carbon and nitrogen fixation differ between successional stages of biological soil crusts in the Colorado Plateau and Chihuahuan Desert

Biological soil crusts (cyanobacteria, mosses and lichens collectively) perform essential ecosystem services, including carbon (C) and nitrogen (N) fixation. Climate and land-use change are converting later successional soil crusts to early successional soil crusts with lower C and N fixation rates. To quantify the effect of such conversions on C and N dynamics in desert ecosystems we seasonally measured diurnal fixation rates in different biological soil crusts. We classified plots on the Colorado Plateau (Canyonlands) and Chihuahuan Desert (Jornada) as early (Microcoleus) or later successional (Nostoc/Scytonema or Placidium/Collema) and measured photosynthesis (Pn), nitrogenase activity (NA), and chlorophyll fluorescence (Fv/Fm) on metabolically active (moist) soil crusts. Later successional crusts typically had greater Pn, averaging 1.2-1.3-fold higher daily C fixation in Canyonlands and 2.4-2.8-fold higher in the Jornada. Later successional crusts also had greater NA, averaging 1.3-7.5-fold higher daily N fixation in Canyonlands and 1.3-25.0-fold higher in the Jornada. Mean daily Fv/Fm was also greater in later successional Canyonlands crusts during winter, and Jornada crusts during all seasons except summer. Together these findings indicate conversion of soil crusts back to early successional stages results in large reductions of C and N inputs into these ecosystems.

Journal of Arid Environments

Spatial modeling of biological soil crusts to support rangeland assessment and monitoring

Biological soil crusts are a diverse soil surface community, prevalent in semiarid regions, which function as ecosystem engineers and perform numerous important ecosystem services. Loss of crusts has been implicated as a factor leading to accelerated soil erosion and other forms of land degradation. To support assessment and monitoring efforts aimed at ensuring the sustainability of rangeland ecosystems, managers require spatially explicit information concerning potential cover and composition of biological soil crusts. We sampled low disturbance sites in Grand Staircase-Escalante National Monument (Utah, USA) to determine the feasibility of modeling the potential cover and composition of biological soil crusts in a large area. We used classification and regression trees to model cover of four crust types (light cyanobacterial, dark cyanobacterial, moss, lichen) and 1 cyanobacterial biomass proxy (chlorophyll a), based upon a parsimonious set of GIS (Geographic Information Systems) data layers (soil types, precipitation, and elevation). Soil type was consistently the best predictor, although elevation and precipitation were both invoked in the various models. Predicted and observed values for the dark cyanobacterial, moss, and lichen models corresponded moderately well (R 2 = 0.49, 0.64, 0.55, respectively). Cover of late successional crust elements (moss + lichen + dark cyanobacterial) was also successfully modeled (R2 = 0.64). We were less successful with models of light cyanobacterial cover (R2 = 0.22) and chlorophyll a (R2 = 0.09). We believe that our difficulty modeling chlorophyll a concentration is related to a severe drought and subsequent cyanobacterial mortality during the course of the study. These models provide the necessary reference conditions to facilitate the comparison between the actual cover and composition of biological soil crusts at a given site and their potential cover and composition condition so that sites in poor condition can be identified and management actions can be taken.

Rangeland Ecology and Management

Relevance of ammonium oxidation within biological soil crust communities

Thin, vertically structured topsoil communities that become ecologically important in arid regions (biological soil crusts or BSCs) are responsible for much of the nitrogen inputs into pristine arid lands. We studied N2 fixation and ammonium oxidation (AO) at subcentimetre resolution within BSCs from the Colorado Plateau. Pools of dissolved porewater nitrate/ nitrite, ammonium and organic nitrogen in wetted BSCs were high in comparison with those typical of aridosoils. They remained stable during incubations, indicating that input and output processes were of similar magnitude. Areal N2 fixation rates (6.5-48 ??mol C2H2 m-2 h -1) were high, the vertical distribution of N2 fixation peaking close to the surface if populations of heterocystous cyanobacteria were present, but in the subsurface if they were absent. Areal AO rates (19-46 ??mol N m-2 h-1) were commensurate with N2 fixation inputs. When considering oxygen availability, AO activity invariably peaked 2-3 mm deep and was limited by oxygen (not ammonium) supply. Most probable number (MPN)-enumerated ammonia-oxidizing bacteria (6.7-7.9 ?? 103 cells g-1 on average) clearly peaked at 2-3 mm depth. Thus, AO (hence nitrification) is a spatially restricted but important process in the nitrogen cycling of BSC, turning much of the biologically fixed nitrogen into oxidized forms, the fate of which remains to be determined.

Environmental Microbiology

Wildfire-resistant biological soil crusts and fire-induced loss of soil stability in Palouse prairies, USA

Frequent low-intensity fires are a natural component of the ecology of the Palouse prairies of northwestern North America. To study the effects of fire upon biological soil crusts (BSCs) occurring in these grasslands, we sampled three burned (in 2000) sites and three unburned sites in the Hell's Canyon area (OR, USA) ???1 year post-fire. We measured vascular plant and BSC cover, soil microbe pigmentation, texture and chemistry, and soil surface physical properties (stability and rugosity). Festuca idahoensis was two times more abundant in unburned plots (P=0.0006), and vascular plant and litter cover were generally higher in unburned plots. At the community scale, there was no difference in the lichen and moss species composition, suggesting much less drastic effects of fire on BSCs than reported in other systems. Soil surface stability (measured using slake value) was significantly lower in burned sites than unburned sites (median value=5 versus 6, P=0.008), a result which is likely due to the greater density of lichens and mosses encountered in the unburned plots. Soil microbe pigmentation was lower in burned plots (P=0.03), suggesting that the biomass of photosynthetic microbes had decreased; however, the presence of intra- and extracellular pigments in burned soils indicates that microorganisms were not eradicated. Pigments most strongly associated with cyanobacteria were more abundant in unburned sites, suggesting that cyanobacteria may have been more strongly impacted by the fire than other BSC components. Composition of nutrients and surface rugosity did not differ significantly between treatments. We hypothesize that Palouse prairie soil crusts are relatively resistant to wildfire because of low fire intensity and their occupation of space away from the vascular plant fuel load.

Applied Soil Ecology

Changes in plant functional groups, litter quality, and soil carbon and nitrogen mineralization with sheep grazing in an Inner Mongolian Grassland

This study reports on changes in plant functional group composition, litter quality, and soil C and N mineralization dynamics from a 9-year sheep grazing study in Inner Mongolia. Addressed are these questions: 1) How does increasing grazing intensity affect plant community composition? 2) How does increasing grazing intensity alter soil C and N mineralization dynamics? 3) Do changes in soil C and N mineralization dynamics relate to changes in plant community composition via inputs of the quality or quantity of litter? Grazing plots were set up near the Inner Mongolia Grassland Ecosystem Research Station (IMGERS) with 5 grazing intensities: 1.3, 2.7, 4.0, 5.3, and 6.7 sheep ha −1 ·yr −1 . Plant cover was lower with increasing grazing intensity, which was primarily due to a dramatic decline in grasses, Carex duriuscula , and Artemisia frigida . Changes in litter mass and percentage organic C resulted in lower total C in the litter layer at 4.0 and 5.3 sheep ha −1 ·yr −1 compared with 2.7 sheep ha −1 ·yr −1 . Total litter N was lower at 5.3 sheep ha −1 ·yr −1 compared with 2.7 sheep ha −1 ·yr −1 . Litter C:N ratios, an index of litter quality, were significantly lower at 4.0 sheep ha −1 ·yr −1 relative to 1.3 and 5.3 sheep ha −1 ·yr −1 . Cumulative C mineralized after 16 days decreased with increasing grazing intensity. In contrast, net N mineralization ( NH 4 NO − 3 ) after a 12-day incubation increased with increasing grazing intensity. Changes in C and N mineralization resulted in a narrowing of CO 2 -C:net N min ratios with increasing grazing intensity. Grazing explained 31% of the variability in the ratio of CO 2 -C:net N min . The ratio of CO 2 -C:net N min was positively correlated with litter mass. Furthermore, there was a positive correlation between litter mass and A. frigida cover. Results suggest that as grazing intensity increases, microbes become more C limited resulting in decreased microbial growth and demand for N.

Journal of Range Management

Response of desert biological soil crusts to alterations in precipitation frequency

Biological soil crusts, a community of cyanobacteria, lichens, and mosses that live on the soil surface, occur in deserts throughout the world. They are a critical component of desert ecosystems, as they are important contributors to soil fertility and stability. Future climate scenarios predict alteration of the timing and amount of precipitation in desert environments. Because biological soil crust organisms are only metabolically active when wet, and as soil surfaces dry quickly in deserts during late spring, summer, and early fall, the amount and timing of precipitation is likely to have significant impacts on the physiological functioning of these communities. Using the three dominant soil crust types found in the western United States, we applied three levels of precipitation frequency (50% below-average, average, and 50% above-average) while maintaining average precipitation amount (therefore changing both timing and size of applied events). We measured the impact of these treatments on photosynthetic performance (as indicated by dark-adapted quantum yield and chlorophyll a concentrations), nitrogenase activity, and the ability of these organisms to maintain concentrations of radiation-protective pigments (scytonemin, beta-carotene, echinenone, xanthophylls, and canthaxanthin). Increased precipitation frequency produced little response after 2.5 months exposure during spring (1 April-15 June) or summer (15 June-31 August). In contrast, most of the above variables had a large, negative response after exposure to increased precipitation frequency for 6 months spring-fall (1 April-31 October) treatment. The crusts dominated by the soil lichen Collema, being dark and protruding above the surface, dried the most rapidly, followed by the dark surface cyanobacterial crusts (Nostoc-Scytonema-Microcoleus), and then by the light cyanobacterial crusts (Microcoleus). This order reflected the magnitude of the observed response: crusts dominated by the lichen Collema showed the largest decline in quantum yield, chlorophyll a, and protective pigments; crusts dominated by Nostoc-Scytonema-Microcoleus showed an intermediate decline in these variables; and the crusts dominated by Microcoleus showed the least negative response. Most previous studies of crust response to radiation stress have been short-term laboratory studies, where organisms were watered and kept under moderate temperatures. Such conditions would give crust organisms access to ample carbon to respond to imposed stresses (e.g., production of UV-protective pigments, replacement of degraded chlorophyll). In contrast, our longer-term study showed that under field conditions of high air temperatures and frequent, small precipitation events, crust organisms appear unable to produce protective pigments in response to radiation stress, as they likely dried more quickly than when they received larger, less frequent events. Reduced activity time likely resulted in less carbon available to produce or repair chlorophyll a and/or protective pigments. Our findings may partially explain the global observation that soil lichen cover and richness declines as the frequency of summer rainfall increases. ?? Springer-Verlag 2003.

Oecologia

Roads as conduits for exotic plant invasions in a semiarid landscape

Roads are believed to be a major contributing factor to the ongoing spread of exotic plants. We examined the effect of road improvement and environmental variables on exotic and native plant diversity in roadside verges and adjacent semiarid grassland, shrubland, and woodland communities of southern Utah (U.S.A.). We measured the cover of exotic and native species in roadside verges and both the richness and cover of exotic and native species in adjacent interior communities (50 m beyond the edge of the road cut) along 42 roads stratified by level of road improvement (paved, improved surface, graded, and four-wheel-drive track). In roadside verges along paved roads, the cover of Bromus tectorum was three times as great (27%) as in verges along four-wheel-drive tracks (9%). The cover of five common exotic forb species tended to be lower in verges along four-wheel-drive tracks than in verges along more improved roads. The richness and cover of exotic species were both more than 50% greater, and the richness of native species was 30% lower, at interior sites adjacent to paved roads than at those adjacent to four-wheel-drive tracks. In addition, environmental variables relating to dominant vegetation, disturbance, and topography were significantly correlated with exotic and native species richness and cover. Improved roads can act as conduits for the invasion of adjacent ecosystems by converting natural habitats to those highly vulnerable to invasion. However, variation in dominant vegetation, soil moisture, nutrient levels, soil depth, disturbance, and topography may render interior communities differentially susceptible to invasions originating from roadside verges. Plant communities that are both physically invasible (e.g., characterized by deep or fertile soils) and disturbed appear most vulnerable. Decision-makers considering whether to build, improve, and maintain roads should take into account the potential spread of exotic plants.

Conservation Biology

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.

Microbial Ecology

Temporal variation in community composition, pigmentation, and Fv/Fm 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 (Fv/Fm) 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 Fv/Fm 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.

Microbial Ecology

Patterns of plant invasions: A case example in native species hotspots and rare habitats

Land managers require landscape-scale information on where exotic plant species have successfully established, to better guide research, control, and restoration efforts. We evaluated the vulnerability of various habitats to invasion by exotic plant species in a 100,000 ha area in the southeast corner of Grand Staircase-Escalante National Monument, Utah. For the 97 0.1-ha plots in 11 vegetation types, exotic species richness (log 10 ) was strongly negatively correlated to the cover of cryptobiotic soil crusts ( r = −0.47, P < 0.001), and positively correlated to native species richness ( r = 0.22, P < 0.03), native species cover ( r = 0.23, P < 0.05), and total nitrogen in the soil ( r = 0.40, P < 0.001). Exotic species cover was strongly positively correlated to exotic species richness ( r = 0.68, P < 0.001). Only 6 of 97 plots did not contain at least one exotic species. Exotic species richness was particularly high in locally rare, mesic vegetation types and nitrogen rich soils. Dry, upland plots ( n = 51) had less than half of the exotic species richness and cover compared to plots ( n = 45) in washes and lowland depressions that collect water intermittently. Plots dominated by trees had significantly greater native and exotic species richness compared to plots dominated by shrubs. For the 97 plots combined, 33% of the variance in exotic species richness could be explained by a positive relationship with total plant cover, and negative relationships with the cover of cryptobiotic crusts and bare ground. There are several reasons for concern: (1) Exotic plant species are invading hot spots of native plant diversity and rare/unique habitats. (2) The foliar cover of exotic species was greatest in habitats that had been invaded by several exotic species.(3) Continued disturbance of fragile cryptobiotic crusts by livestock, people, and vehicles may facilitate the further invasion of exotic plant species.

Biological Invasions