Keirith A. Snyder

Precipitation pulses and carbon fluxes in semiarid and arid ecosystems

In the arid and semiarid regions of North America, discrete precipitation pulses are important triggers for biological activity. The timing and magnitude of these pulses may differentially affect the activity of plants and microbes, combining to influence the C balance of desert ecosystems. Here, we evaluate how a “pulse” of water influences physiological activity in plants, soils and ecosystems, and how characteristics, such as precipitation pulse size and frequency are important controllers of biological and physical processes in arid land ecosystems. We show that pulse size regulates C balance by determining the temporal duration of activity for different components of the biota. Microbial respiration responds to very small events, but the relationship between pulse size and duration of activity likely saturates at moderate event sizes. Photosynthetic activity of vascular plants generally increases following relatively larger pulses or a series of small pulses. In this case, the duration of physiological activity is an increasing function of pulse size up to events that are infrequent in these hydroclimatological regions. This differential responsiveness of photosynthesis and respiration results in arid ecosystems acting as immediate C sources to the atmosphere following rainfall, with subsequent periods of C accumulation should pulse size be sufficient to initiate vascular plant activity. Using the average pulse size distributions in the North American deserts, a simple modeling exercise shows that net ecosystem exchange of CO2 is sensitive to changes in the event size distribution representative of wet and dry years. An important regulator of the pulse response is initial soil and canopy conditions and the physical structuring of bare soil and beneath canopy patches on the landscape. Initial condition influences responses to pulses of varying magnitude, while bare soil/beneath canopy patches interact to introduce nonlinearity in the relationship between pulse size and soil water response. Building on this conceptual framework and developing a greater understanding of the complexities of these eco-hydrologic systems may enhance our ability to describe the ecology of desert ecosystems and their sensitivity to global change.

ECOHYDROLOGICAL IMPLICATIONS OF WOODY PLANT ENCROACHMENT

Increases in the abundance or density of woody plants in historically semiarid and arid grassland ecosystems have important ecological, hydrological, and socioeconomic implications. Using a simplified water-balance model, we propose a framework for con- ceptualizing how woody plant encroachment is likely to affect components of the water cycle within these ecosystems. We focus in particular on streamflow and the partitioning of evapotranspiration into evaporation and transpiration. On the basis of this framework, we suggest that streamflow and evaporation processes are affected by woody plant en- croachment in different ways, depending on the degree and seasonality of aridity and the availability of subsurface water. Differences in landscape physiography, climate, and runoff mechanisms mediate the influence of woody plants on hydrological processes. Streamflow is expected to decline as a result of woody plant encroachment in landscapes dominated by subsurface flow regimes. Similarly, encroachment of woody plants can be expected to produce an increase in the fractional contribution of bare soil evaporation to evapotrans- piration in semiarid ecosystems, whereas such shifts may be small or negligible in both subhumid and arid ecosystems. This framework for considering the effects of woody plant encroachment highlights important ecological and hydrological interactions that serve as a basis for predicting other ecological aspects of vegetation change—such as potential changes in carbon cycling within an ecosystem. In locations where woody plant encroach- ment results in increased plant transpiration and concurrently the availability of soil water is reduced, increased accumulation of carbon in soils emerges as one prediction. Thus, explicitly considering the ecohydrological linkages associated with vegetation change pro- vides needed information on the consequences of woody plant encroachment on water yield, carbon cycling, and other processes.

Water sources used by riparian trees varies among stream types on the San Pedro River, Arizona

Variation in the sources of water used by tree species has important ramifications for forest water balances. The fraction of tree transpiration water derived from the unsaturated soil zone and groundwater in a riparian forest was quantified for Populus fremontii, Salix gooddingii, and Prosopis velutinaacross a gradient of groundwater depth and streamflow regime on the San Pedro River in southeastern Arizona, US. The proportion of tree transpiration derived from different potential sources was determined using oxygen ( 18 O) and hydrogen (D) stable isotope analysis in conjunction with two- and three-compartment linear mixing models. Comparisons of 18 O and D of tree xylem water with that of potential water sources indicated that Salix gooddingii did not take up water in the upper soil layers during the summer rainy period, but instead used only groundwater, even at an ephemeral stream site where depth to groundwater exceeded 4 m. Populus fremontii, a dominant ‘phreatophyte’ in these semi-arid riparian ecosystems, also used mainly groundwater, but at the ephemeral stream site during the summer rainy season this species derived between 26 and 33% of its transpiration water from upper soil layers. Similarly, at the ephemeral stream site during the summer rainy period, Prosopis velutinaderived a greater fraction of its transpiration water from upper soil layers, than at a perennial stream site where groundwater depth was less than 2 m. Measurements of transpiration flux combined with stable isotope data revealed that Populus fremontiitranspired a greater quantity of water from upper soil layers at the ephemeral stream site than at the perennial stream site. These results imply that transpiration from groundwater and unsaturated soil layers by riparian vegetation may depend on the interaction between site conditions and species assemblage.

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

Tamarisk biocontrol in the western United States: ecological and societal implications

Tamarisk species (genus Tamarix), also commonly known as saltcedar, are among the most successful plant invaders in the western United States. At the same time, tamarisk has been cited as having enormous economic costs. Accordingly, local, state, and federal agencies have undertaken considerable efforts to eradicate this invasive plant and restore riparian habitats to pre-invasion status. Traditional eradication methods, including herbicide treatments, are now considered undesirable, because they are costly and often have unintended negative impacts on native species. A new biological control agent, the saltcedar leaf beetle (Diorhabda elongata), has been released along many watersheds in the western US, to reduce the extent of tamarisk cover in riparian areas. However, the use of this insect as a biological control agent may have unintended ecological, hydrological, and socioeconomic consequences that need to be anticipated by land managers and stakeholders undertaking restoration efforts. Here, we examine the possible ramifications of tamarisk control and offer recommendations to reduce potential negative impacts on valued riparian systems in the western US.

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.

Reconstructing relative humidity from plant δ18O and δD as deuterium deviations from the global meteoric water line

Cellulose delta18O and deltaD can provide insights on climates and hydrological cycling in the distant past and how these factors differ spatially. However, most studies of plant cellulose have used only one isotope, most commonly delta18O, resulting in difficulties partitioning variation in delta18O of precipitation vs. evaporative conditions that affect leaf water isotopic enrichment. Moreover, observations of pronounced diurnal differences from conventional steady-state model predictions of leaf water isotopic fractionation have cast some doubt on single isotope modeling approaches for separating precipitation and evaporation drivers of cellulose delta18O or deltaD. We explore a dual isotope approach akin to the concept of deuterium-excess (d), to establish deuterium deviations from the global meteoric water line in leaf water (deltad(l)) as driven by relative humidity (RH). To demonstrate this concept, we survey studies of leaf water delta18O and deltaD in hardwood vs. conifer trees. We then apply the concept to cellulose delta18O and deltaD using a mechanistic model of cellulose delta18O and deltaD to reconstruct deuterium deviations from the global meteoric water line (deltad(c)) in Quercus macrocarpa, Q. robur, and Pseudotsuga menziesii. For each species, deltad(c) showed strong correlations with RH across sites. deltad(c) agreed well with steady-state predictions for Q. macrocarpa, while for Q. robur, the relationship with RH was steeper than expected. The slope of deltad(c) vs. RH of P. menziesii was also close to steady-state predictions, but deltad(c) were more enriched than predicted. This is in agreement with our leaf water survey showing conifer deltad(l) was more enriched than predicted. Our data reveal that applications of this method should be appropriate for reconstructing RH from cellulose delta18O and deltaD after accounting for differences between hardwoods and conifers. Hence, deltad(c) should be useful for understanding variability in RH associated with past climatic cycles, across regional climates, or across complex terrain where climate modeling is challenging. Furthermore, deltad(c) and inferred RH values should help in constraining variation in source water delta18O.

Precipitation Regulates the Response of Net Ecosystem CO2 Exchange to Environmental Variation on United States Rangelands

Abstract Rangelands occupy 50% of Earths land surface and thus are important in the terrestrial carbon (C) cycle. For rangelands and other terrestrial ecosystems, the balance between photosynthetic uptake of carbon dioxide (CO2) and CO2 loss to respiration varies among years in response to interannual variation in the environment. Variability in CO2 exchange results from interannual differences in 1) environmental variables at a given point in the annual cycle (direct effects of the environment) and in 2) the response of fluxes to a given change in the environment because of interannual changes in biological factors that regulate photosynthesis and respiration (functional change). Functional change is calculated as the contribution of among-year differences in slopes of flux-environment relationships to the total variance in fluxes explained by the environment. Functional change complicates environmental-based predictions of CO2 exchange, yet its causes and contribution to flux variability remain poorly defined. We determine contributions of functional change and direct effects of the environment to interannual variation in net ecosystem exchange of CO2 (NEE) of eight rangeland ecosystems in the western United States (58 site-years of data). We predicted that 1) functional change is correlated with interannual change in precipitation on each rangeland and 2) the contribution of functional change to variance in NEE increases among rangelands as mean precipitation increases. Functional change explained 10–40% of the variance in NEE and accounted for more than twice the variance in fluxes of direct effects of environmental variability for six of the eight ecosystems. Functional change was associated with interannual variation in precipitation on most rangelands but, contrary to prediction, contributed proportionally more to variance in NEE on arid than more mesic ecosystems. Results indicate that we must account for the influence of precipitation on flux-environment relationships if we are to distinguish environmental from management effects on rangeland C balance.

Ecophysiological responses of salt cedar (Tamarix spp. L.) to the northern tamarisk beetle (Diorhabda carinulata Desbrochers) in a controlled environment.

The northern tamarisk beetle (Diorhabda carinulata Desbrochers) was released in several western states as a biocontrol agent to suppress Tamarix spp. L. which has invaded riparian ecosystems; however, effects of beetle herbivory on Tamarix physiology are largely undocumented and may have ecosystem ramifications. Herbivory by this insect produces discoloration of leaves and premature leaf drop in these ecosystems, yet the cause of premature leaf drop and the effects of this leaf drop are still unknown. Insect herbivory may change leaf photosynthesis and respiration and may affect a plant’s ability to regulate water loss and increase water stress. Premature leaf drop may affect plant tissue chemistry and belowground carbon allocation. We conducted a greenhouse experiment to understand how Tamarix responds physiologically to adult beetle and larvae herbivory and to determine the proximate cause of premature leaf drop. We hypothesized that plants experiencing beetle herbivory would have greater leaf and root respiration rates, greater photosynthesis, increased water stress, inefficient leaf nitrogen retranslocation, lower root biomass and lower total non-structural carbohydrates in roots. Insect herbivory reduced photosynthesis rates, minimally affected respiration rates, but significantly increased water loss during daytime and nighttime hours and this produced increased water stress. The proximate cause for premature leaf drop appears to be desiccation. Plants exposed to herbivory were inefficient in their retranslocation of nitrogen before premature leaf drop. Root biomass showed a decreasing trend in plants subjected to herbivory. Stress induced by herbivory may render these trees less competitive in future growing seasons.

Evaluating mountain meadow groundwater response to Pinyon-Juniper and temperature in a Great Basin watershed†

This research highlights development and application of an integrated hydrologic model (GSFLOW) to a semiarid, snow-dominated watershed in the Great Basin to evaluate Pinyon-Juniper (PJ) and temperature controls on mountain meadow shallow groundwater. The work used Google Earth Engine Landsat satellite and gridded climate archives for model evaluation. Model simulations across three decades indicated that the watershed operates on a threshold response to precipitation (P) > 400 mm y-1 to produce a positive yield (P-ET; 9%) resulting in stream discharge and a rebound in meadow groundwater levels during these wetter years. Observed and simulated meadow groundwater response to large P correlates with above average predicted soil moisture and with a normalized difference vegetation index (NDVI) threshold value > 0.3. A return to assumed pre-expansion PJ conditions or an increase in temperature to mid-21st century shifts yielded by only ±1% during the multi-decade simulation period; but changes of approximately ±4% occurred during wet years. Changes in annual yield were largely dampened by the spatial and temporal redistribution of evapotranspiration (ET) across the watershed. Yet, the influence of this redistribution and vegetation structural controls on snowmelt altered recharge to control water table depth in the meadow. Even a small-scale removal of PJ (0.5 km2) proximal to the meadow will promote a stable, shallow groundwater system resilient to droughts, while modest increases in temperature will produce a meadow susceptible to declining water levels and a community structure likely to move toward dry and degraded conditions. This article is protected by copyright. All rights reserved.