Jayne Belnap

Water pulses and biogeochemical cycles in arid and semiarid ecosystems

The episodic nature of water availability in arid and semiarid ecosystems has significant consequences on belowground carbon and nutrient cycling. Pulsed water events directly control belowground processes through soil wet-dry cycles. Rapid soil microbial response to incident moisture availability often results in almost instantaneous C and N mineralization, followed by shifts in C/N of microbially available substrate, and an offset in the balance between nutrient immobilization and mineralization. Nitrogen inputs from biological soil crusts are also highly sensitive to pulsed rain events, and nitrogen losses, particularly gaseous losses due to denitrification and nitrate leaching, are tightly linked to pulses of water availability. The magnitude of the effect of water pulses on carbon and nutrient pools, however, depends on the distribution of resource availability and soil organisms, both of which are strongly affected by the spatial and temporal heterogeneity of vegetation cover, topographic position and soil texture. The ‘inverse texture hypothesis’ for net primary production in water-limited ecosystems suggests that coarse-textured soils have higher NPP than fine-textured soils in very arid zones due to reduced evaporative losses, while NPP is greater in fine-textured soils in higher rainfall ecosystems due to increased water-holding capacity. With respect to belowground processes, fine-textured soils tend to have higher water-holding capacity and labile C and N pools than coarse-textured soils, and often show a much greater flush of N mineralization. The result of the interaction of texture and pulsed rainfall events suggests a corollary hypothesis for nutrient turnover in arid and semiarid ecosystems with a linear increase of N mineralization in coarse-textured soils, but a saturating response for fine-textured soils due to the importance of soil C and N pools. Seasonal distribution of water pulses can lead to the accumulation of mineral N in the dry season, decoupling resource supply and microbial and plant demand, and resulting in increased losses via other pathways and reduction in overall soil nutrient pools. The asynchrony of resource availability, particularly nitrogen versus water due to pulsed water events, may be central to understanding the consequences for ecosystem nutrient retention and long-term effects on carbon and nutrient pools. Finally, global change effects due to changes in the nature and size of pulsed water events and increased asynchrony of water availability and growing season will likely have impacts on biogeochemical cycling in water-limited ecosystems.

Biological soil crusts : structure, function, and management

The volume comprises 33 chapters and is divided into the following eight Parts:Part I: Taxonomic Composition, Ecology and Biogeography of Soil-Crust Communities * Part II: Heterotrophic Components of Biological Soil Crusts * Part III: Structure of Biological Soil Crusts: Microscale to Landscape * Part IV: Biological Soil Crusts as an Ecosystem Component: Carbon and Nitrogen Acquisition and Interactions with Vascular Plants * Part V: Soil Stability and Hydrology as Influenced by Soil Crusts * Part VI: Disturbance to Biological Soil Crusts: Resistance, Resilience and Restoration * Part VII: Monitoring and Management of Biological Soil Crusts * Part VIII: Conclusions.

EXOTIC PLANT INVASION ALTERS NITROGEN DYNAMICS IN AN ARID GRASSLAND

The introduction of nonnative plant species may decrease ecosystem stability by altering the availability of nitrogen (N) for plant growth. Invasive species can impact N availability by changing litter quantity and quality, rates of N 2-fixation, or rates of N loss. We quantified the effects of invasion by the annual grass Bromus tectorum on N cycling in an arid grassland on the Colorado Plateau (USA). The invasion occurred in 1994 in two community types in an undisturbed grassland. This natural experiment allowed us to measure the immediate responses following invasion without the confounding effects of previous disturbance. Litter biomass and the C:N and lignin:N ratios were measured to determine the effects on litter dynamics. Long-term soil incubations (415 d) were used to measure potential microbial respiration and net N mineralization. Plant-available N was quantified for two years in situ with ion-exchange resin bags, and potential changes in rates of gaseous N loss were estimated by measuring denitrification enzyme activity. Bromus invasion significantly increased litter biomass, and Bromus litter had significantly greater C:N and lignin:N ratios than did native species. The change in litter quantity and chemistry decreased potential rates of net N mineralization in sites with Bromusby decreasing nitrogen available for microbial activity. Inorganic N was 50% lower on Hilaria sites with Bromus during the spring of 1997, but no differences were observed during 1998. The contrasting differences between years are likely due to moisture availability; spring precipitation was 15% greater than average during 1997, but 52% below average during spring of 1998. Bromus may cause a short-term decrease in N loss by decreasing substrate availability and denitrification enzyme activity, but N loss is likely to be greater in invaded sites in the long term because of increased fire frequency and greater N volatilization during fire. We hypothesize that the introduction of Bromus in conjunction with land-use change has es- tablished a series of positive feedbacks that will decrease N availability and alter species composition.

Comparison of Soil Bacterial Communities in Rhizospheres of Three Plant Species and the Interspaces in an Arid Grassland

ABSTRACT Soil bacteria are important contributors to primary productivity and nutrient cycling in arid land ecosystems, and their populations may be greatly affected by changes in environmental conditions. In parallel studies, the composition of the total bacterial community and of members of the Acidobacterium division were assessed in arid grassland soils using terminal restriction fragment length polymorphism (TRF, also known as T-RFLP) analysis of 16S rRNA genes amplified from soil DNA. Bacterial communities associated with the rhizospheres of the native bunchgrasses Stipa hymenoides and Hilaria jamesii, the invading annual grass Bromus tectorum, and the interspaces colonized by cyanobacterial soil crusts were compared at three depths. When used in a replicated field-scale study, TRF analysis was useful for identifying broad-scale, consistent differences in the bacterial communities in different soil locations, over the natural microscale heterogeneity of the soil. The compositions of the total bacterial community and Acidobacterium division in the soil crust interspaces were significantly different from those of the plant rhizospheres. Major differences were also observed in the rhizospheres of the three plant species and were most apparent with analysis of the Acidobacterium division. The total bacterial community and the Acidobacterium division bacteria were affected by soil depth in both the interspaces and plant rhizospheres. This study provides a baseline for monitoring bacterial community structure and dynamics with changes in plant cover and environmental conditions in the arid grasslands.

SOIL BIOTA IN AN UNGRAZED GRASSLAND: RESPONSE TO ANNUAL GRASS (BROMUS TECTORUM) INVASION

Bromus tectorum is an exotic annual grass that currently dominates many western U.S. semi-arid ecosystems, and the effects of this grass on ecosystems in general, and soil biota specifically, are unknown. Bromus recently invaded two ungrazed and unburned perennial bunchgrass communities in southeastern Utah. This study compared the soil food-web structure of the two native grassland associations (Stipa [S] and Hilaria [H]), with and without the presence of Bromus. Perennial grass and total vascular-plant cover were higher in S than in H plots, while quantities of ground litter were similar. Distribution of live and dead plant material was highly clumped in S and fairly homogenous in H. Soil food-web structure was different between H and S, with lower trophic levels more abundant in H and higher trophic levels more abundant in S. In Bromus-invaded plots, the quantity of ground litter was 2.2 times higher in Hilaria–Bromus (HB) than in H plots, and 2.8 times higher in Stipa–Bromus (SB) than in S plots. Soil...

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.

MICROENVIRONMENTS AND MICROSCALE PRODUCTIVITY OF CYANOBACTERIAL DESERT CRUSTS1

We used microsensors to characterize physicochemical microenvironments and photosynthesis occurring immediately after water saturation in two desert soil crusts from southeastern Utah, which were formed by the cyanobacteria Microcoleus vaginatus Gomont, Nostoc spp., and Scytonema sp. The light fields within the crusts presented steep vertical gradients in magnitude and spectral composition. Near‐surface light‐trapping zones were formed due to the scattering nature of the sand particles, but strong light attenuation resulted in euphotic zones only ca. 1 mm deep, which were progressively enriched in longer wavelengths with depth. Rates of gross photosynthesis (3.4–9.4 mmol O2·m−2·h−1) and dark respiration (0.81–3.1 mmol O−2·m−2·h−1) occurring within 1 to several mm from the surface were high enough to drive the formation of marked oxygen microenvironments that ranged from oxygen supersaturation to anoxia. The photosynthetic activity also resulted in localized pH values in excess of 10, 2–3 units above the soil pH. Differences in metabolic parameters and community structure between two types of crusts were consistent with a successional pattern, which could be partially explained on the basis of the microenvironments. We discuss the significance of high metabolic rates and the formation of microenvironments for the ecology of desert crusts, as well as the advantages and limitations of microsensor‐based methods for crust investigation.

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.

Response of Colorado River runoff to dust radiative forcing in snow

The waters of the Colorado River serve 27 million people in seven states and two countries but are overallocated by more than 10% of the river’s historical mean. Climate models project runoff losses of 7–20% from the basin in this century due to human-induced climate change. Recent work has shown however that by the late 1800s, decades prior to allocation of the river’s runoff in the 1920s, a fivefold increase in dust loading from anthropogenically disturbed soils in the southwest United States was already decreasing snow albedo and shortening the duration of snow cover by several weeks. The degree to which this increase in radiative forcing by dust in snow has affected timing and magnitude of runoff from the Upper Colorado River Basin (UCRB) is unknown. Here we use the Variable Infiltration Capacity model with postdisturbance and predisturbance impacts of dust on albedo to estimate the impact on runoff from the UCRB across 1916–2003. We find that peak runoff at Lees Ferry, Arizona has occurred on average 3 wk earlier under heavier dust loading and that increases in evapotranspiration from earlier exposure of vegetation and soils decreases annual runoff by more than 1.0 billion cubic meters or ∼5% of the annual average. The potential to reduce dust loading through surface stabilization in the deserts and restore more persistent snow cover, slow runoff, and increase water resources in the UCRB may represent an important mitigation opportunity to reduce system management tensions and regional impacts of climate change.

A Classification of Ecological Boundaries

Abstract Ecologists use the term boundary to refer to a wide range of real and conceptual structures. Because imprecise terminology may impede the search for general patterns and theories about ecological boundaries, we present a classification of the attributes of ecological boundaries to aid in communication and theory development. Ecological boundaries may differ in their origin and maintenance, their spatial structure, their function, and their temporal dynamics. A classification system based on these attributes should help ecologists determine whether boundaries are truly comparable. This system can be applied when comparing empirical studies, comparing theories, and testing theoretical predictions against empirical results.