Sean M. Schaeffer

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.

Microbial control over carbon cycling in soil

A major thrust of terrestrial microbial ecology is focused on understanding when and how the composition of the microbial community affects the functioning of biogeochemical processes at the ecosystem scale (meters-to-kilometers and days-to-years). While research has demonstrated these linkages for physiologically and phylogenetically “narrow” processes such as trace gas emissions and nitrification, there is less conclusive evidence that microbial community composition influences the “broad” processes of decomposition and organic matter (OM) turnover in soil. In this paper, we consider how soil microbial community structure influences C cycling. We consider the phylogenetic level at which microbes form meaningful guilds, based on overall life history strategies, and suggest that these are associated with deep evolutionary divergences, while much of the species-level diversity probably reflects functional redundancy. We then consider under what conditions it is possible for differences among microbes to affect process dynamics, and argue that while microbial community structure may be important in the rate of OM breakdown in the rhizosphere and in detritus, it is likely not important in the mineral soil. In mineral soil, physical access to occluded or sorbed substrates is the rate-limiting process. Microbial community influences on OM turnover in mineral soils are based on how organisms allocate the C they take up – not only do the fates of the molecules differ, but they can affect the soil system differently as well. For example, extracellular enzymes and extracellular polysaccharides can be key controls on soil structure and function. How microbes allocate C may also be particularly important for understanding the long-term fate of C in soil – is it sequestered or not?

Seasonal estimates of riparian evapotranspiration using remote and in situ measurements

In many semi-arid basins during extended periods when surface snowmelt or storm runoff is absent, groundwater constitutes the primary water source for human habitation, agriculture and riparian ecosystems. Utilizing regional groundwater models in the management of these water resources requires accurate estimates of basin boundary conditions. A critical groundwater boundary condition that is closely coupled to atmospheric processes and is typically known with little certainty is seasonal riparian evapotranspiration (ET). This quantity can often be a significant factor in the basin water balance in semi-arid regions yet is very difficult to estimate over a large area. Better understanding and quantification of seasonal, large-area riparian ET is a primary objective of the Semi-Arid Land-Surface-Atmosphere (SALSA) Program. To address this objective, a series of interdisciplinary experimental campaigns were conducted in 1997 in the San Pedro Basin in southeastern Arizona. The riparian system in this basin is primarily made up of three vegetation communities: mesquite (Prosopis velutina), sacaton grasses (Sporobolus wrightii), and a cottonwood (Populus fremontii)/willow (Salix goodingii) forest gallery. Micrometeorological measurement techniques were used to estimate ET from the mesquite and grasses. These techniques could not be utilized to estimate fluxes from the cottonwood/willow (C/W) forest gallery due to the height (20‐30 m) and non-uniform linear nature of the forest gallery. Short-term (2‐4 days) sap flux measurements were made to estimate canopy transpiration over several periods of the riparian growing season. Simultaneous remote sensing measurements were used to spatially extrapolate tree and stand measurements. Scaled C/W stand level sap flux estimates were utilized to calibrate a Penman‐Monteith model to enable temporal extrapolation between synoptic measurement periods. With this model and set of measurements, seasonal riparian vegetation water use estimates for the riparian corridor were obtained. To validate these models, a 90-day pre-monsoon water balance over a 10 km section of the river was carried out. All components of the water balance, including riparian ET, were

Transpiration of cottonwood/willow forest estimated from sap flux

Cottonwood/willow forests in the American Southwest consist of discrete, even-aged vegetation patches arranged in narrow strips along active and abandoned stream channels of alluvial flood plains. We used the heat-pulse velocity technique in this study to estimate transpiration in 12 such forest patches along a perennially flowing reach of the San Pedro River in southeastern Arizona, USA during five periods from April to October 1997. Transpiration per unit sapwood area was consistently higher for the larger cottonwood trees found on outer secondary channels compared to that of smaller cottonwood trees along the active channel, but statistically significant differences were found only in August and October. Conversely, transpiration per unit sapwood area in willow was markedly higher for trees along the primary channel than for those few larger trees that were sampled on the outer margins of the forest. Average daily transpiration at the canopy scale among the patches in July was 4:8 0:7 mm per day and ranged from 5:7 0:6 mm per day in young forest patches adjacent to the primary stream channel to 3:1 0:6 mm per day in more successionally advanced patches on secondary channels. Differences in our estimates of transpiration between forest patches along primary and secondary stream channels were related to differences in the ratio of sapwood area to ground area of the forest patches, and leaf area index. Estimates of transpiration from this forest type, and projections of transpiration and groundwater flux over larger areas on the San Pedro River, should take into account structural variation in these forests that relate to population dynamics of dominant trees.

Seasonal and episodic moisture controls on plant and microbial contributions to soil respiration

Moisture inputs drive soil respiration (SR) dynamics in semi-arid and arid ecosystems. However, determining the contributions of root and microbial respiration to SR, and their separate temporal responses to periodic drought and water pulses, remains poorly understood. This study was conducted in a pine forest ecosystem with a Mediterranean-type climate that receives seasonally varying precipitation inputs from both rainfall (in the winter) and fog-drip (primarily in the summer). We used automated SR measurements, radiocarbon SR source partitioning, and a water addition experiment to understand how SR, and its separate root and microbial sources, respond to seasonal and episodic changes in moisture. Seasonal changes in SR were driven by surface soil water content and large changes in root respiration contributions. Superimposed on these seasonal patterns were episodic pulses of precipitation that determined the short-term SR patterns. Warm season precipitation pulses derived from fog-drip, and rainfall following extended dry periods, stimulated the largest SR responses. Microbial respiration dominated these SR responses, increasing within hours, whereas root respiration responded more slowly over days. We conclude that root and microbial respiration sources respond differently in timing and magnitude to both seasonal and episodic moisture inputs. These findings have important implications for the mechanistic representation of SR in models and the response of dry ecosystems to changes in precipitation patterns.

Responses of soil nitrogen dynamics in a Mojave Desert ecosystem to manipulations in soil carbon and nitrogen availability

We investigated the effects of changes in soil C and N availability on N mineralization, nitrification, denitrification, NH3 volatilization, and soil respiration in the Mojave Desert. Results indicate a C limitation to microbial N cycling. Soils from underneath the canopies of Larrea tridentata (DC.) Cov., Pleuraphis rigida Thurber, and Lycium spp. exhibited higher rates of CO2 flux, lower rates of NH3 volatilization, and a decrease in inorganic N (NH4+-N and NO3--N) with C addition. In addition to C limitation, soils from plant interspaces also exhibited a N limitation. Soils from all locations had net immobilization of N over the course of a 15-day laboratory incubation. However, soils from interspaces had lower rates of net nitrification and potential denitrification compared to soils from under plant canopies. The response to changes in C availability appears to be a short-term increase in microbial immobilization of inorganic N. Under controlled conditions, and over a longer time period, the effects of C and N availability appear to give way to larger differences due to spatial location. These findings have implications for ecosystems undergoing changes in soil C and N availability due to such processes as desertification, exotic species invasions, or elevated atmospheric CO2 concentration.

Preface paper to the Semi-Arid Land-Surface-Atmosphere (SALSA) Program special issue.

The Semi-Arid Land-Surface-Atmosphere Program (SALSA) is a multi-agency, multi-national research effort that seeks to evaluate the consequences of natural and human-induced environmental change in semi-arid regions. The ultimate goal of SALSA is to advance scientific understanding of the semi-arid portion of the hydrosphere-biosphere interface in order to provide reliable information for environmental decision making. SALSA approaches this goal through a program of long-term, integrated observations, process research, modeling, assessment, and information management that is sustained by cooperation among scientists and information users. In this preface to the SALSA special issue, general program background information and the critical nature of semi-arid regions is presented. A brief description of the Upper San Pedro River Basin, the initial location for focused SALSA research follows. Several overarching research objectives under which much of the interdisciplinary research contained in the special issue was undertaken are discussed. Principal methods, primary research sites and data collection used by numerous investigators during 1997-1999 are then presented. Scientists from about 20 US, five European (four French and one Dutch), and three Mexican agencies and institutions have collaborated closely to make the research leading to this special issue a reality. The SALSA Program has served as a model of interagency cooperation by breaking new ground in the approach to large scale interdisciplinary science with relatively limited resources.

Alterations of nitrogen dynamics under elevated carbon dioxide in an intact Mojave Desert ecosystem: Evidence from nitrogen-15 natural abundance

We examined soil and vegetation N isotopic composition (δ15N) and soil inorganic N availability in an intact Mojave desert ecosystem to evaluate potential effects of elevated atmospheric CO2 on N cycling. Vegetation from the dominant perennial shrub Larrea tridentata under elevated CO2 was enriched in 15N. Over a 7-month sampling period, Larrea δ15N values increased from 5.7±0.1‰ to 9.0±1.1‰ with elevated CO2; under ambient conditions, δ15N values of shrubs increased from 4.9±0.3‰ to 6.6±0.7‰. No difference was found in soil δ15N under elevated and ambient CO2. Soil δ15N values under the drought deciduous shrubs Lycium spp. were greatest (7.2±0.3‰), and soil under the C4 perennial bunchgrass Pleuraphis rigida had the lowest values (4.5±0.2‰). Several mechanisms could explain the enrichment in 15N of vegetation with elevated CO2. Results suggest that microbial activity has increased with elevated CO2, enriching pools of plant-available N and decreasing N availability. This hypothesis is supported by a significant reduction of plant-available N under elevated CO2. This indicates that exposure to elevated CO2 has resulted in significant perturbations to the soil N cycle, and that plant δ15N may be a useful tool for interpreting changes in the N cycle in numerous ecosystems.

Pulse additions of soil carbon and nitrogen affect soil nitrogen dynamics in an arid Colorado Plateau shrubland

Biogeochemical cycles in arid and semi-arid ecosystems depend upon the ability of soil microbes to use pulses of resources. Brief periods of high activity generally occur after precipitation events that provide access to energy and nutrients (carbon and nitrogen) for soil organisms. To better understand pulse-driven dynamics of microbial soil nitrogen (N) cycling in an arid Colorado Plateau ecosystem, we simulated a pulsed addition of labile carbon (C) and N in the field under the canopies of the major plant species in plant interspaces. Soil microbial activity and N cycling responded positively to added C while NH4+–N additions resulted in an accumulation of soil NO3−. Increases in microbial activity were reflected in higher rates of respiration and N immobilization with C addition. When both C and N were added to soils, N losses via NH3 volatilization decreased. There was no effect of soil C or N availability on microbial biomass N suggesting that the level of microbial activity (respiration) may be more important than population size (biomass) in controlling short-term dynamics of inorganic and labile organic N. The effects of C and N pulses on soil microbial function and pools of NH4+–N and labile organic N were observed to last only for the duration of the moisture pulse created by treatment addition, while the effect on the NO3−–N pool persisted after soils dried to pre-pulse moisture levels. We observed that increases in available C lead to greater ecosystem immobilization and retention of N in soil microbial biomass and also lowered rates of gaseous N loss. With the exception of trace gas N losses, the lack of interaction between available C and N on controlling N dynamics, and the subsequent reduction in plant available N with C addition has implications for the competitive relationships between plants species, plants and microbes, or both.

Soil-plant N processes in a High Arctic ecosystem, NW Greenland are altered by long-term experimental warming and higher rainfall.

Rapid temperature and precipitation changes in High Arctic tundra ecosystems are altering the biogeochemical cycles of carbon (C) and nitrogen (N), but in ways that are difficult to predict. The challenge grows from the uncertainty of N cycle responses and the extent to which shifts in soil N are coupled with the C cycle and productivity of tundra systems. We used a long-term (since 2003) experiment of summer warming and supplemental summer water additions to a High Arctic ecosystem in NW Greenland, and applied a combination of discrete sampling and in situ soil core incubations to measure C and N pools and seasonal microbial processes that might control plant-available N. We hypothesized that elevated temperature and increased precipitation would stimulate microbial activity and net inorganic N mineralization, thereby increasing plant N-availability through the growing season. While we did find increased N mineralization rates under both global change scenarios, water addition also significantly increased net nitrification rates, loss of NO3 (-) -N via leaching, and lowered rates of labile organic N production. We also expected the chronic warming and watering would lead to long-term changes in soil N-cycling that would be reflected in soil δ(15) N values. We found that soil δ(15) N decreased under the different climate change scenarios. Our results suggest that temperature accelerates biological processes and existing C and N transformations, but moisture increases soil hydraulic connectivity and so alters the pathways, and changes the fate of the products of C and N transformations. In addition, our findings indicate that warmer, wetter High Arctic tundra will be cycling N and C in ways that may transform these landscapes in part leading to greater C sequestration, but simultaneously, N losses from the upper soil profile that may be transported to depth dissolved in water and or transported off site in lateral flow.