K. R. Hultine

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.

Seasonal variations in moisture use in a piñon-juniper woodland

In water-limited environments of the intermountain region of North America, summer precipitation may play a role in the structure and function of aridland communities and ecosystems. This study examined the potential reliance on summer precipitation of two widespread, coexisting woody species in the southwestern United States, Pinus edulis Englmn. (Colorado piñon) and Juniperus osteosperma (Torr) Little (Utah juniper). The current distributions of P. edulis and J. osteosperma are highly suggestive of different dependencies on summer rainfall. We hypothesized that P. edulis was dependent on summer precipitation, utilizing summer precipitation even during extremely dry summers, whereas J. osteosperma was not dependent, using summer precipitation only when amounts were above some minimum threshold. Using sap flux and stable isotopic methods to assess seasonal water sources and water use efficiency, we examined the response of these two species to seasonal variations in moisture at a site located near the northern limits of the North American monsoon. Both sap flux and isotopic results indicated that P. edulis was responsive to summer rain, while J. osteosperma was not. Following summer rain events, sap flux density increased in P. edulis for several days, but not in J. osteosperma. Isotopic evidence indicated that P. edulis took up summer-derived moisture to a greater extent than J. osteosperma. Values of the natural abundance stable isotope ratio of carbon of leaf soluble carbohydrates increased over the summer for P. edulis, indicative of assimilation at higher water use efficiency, but were invariant for J. osteosperma. Our results supported the hypothesis that P. edulis and J. osteosperma are differentially sensitive to summer precipitation and are discussed in the light of potential changes in the seasonality of precipitation associated with climate change.

Wood anatomy constrains stomatal responses to atmospheric vapor pressure deficit in irrigated, urban trees.

Plant transpiration is strongly constrained by hydraulic architecture, which determines the critical threshold for cavitation. Because species vary greatly in vulnerability to cavitation, hydraulic limits to transpiration and stomatal conductance have not generally been incorporated into ecological and climate models. We measured sap flow, leaf transpiration, and vulnerability to cavitation of a variety of tree species in a well-irrigated but semi-arid urban environment in order to evaluate the generality of stomatal responses to high atmospheric vapor pressure deficit (D). We found evidence of broad patterns of stomatal responses to humidity based on systematic differences in vulnerability to cavitation. Ring-porous taxa consistently had vulnerable xylem and showed strong regulation of transpiration in response to D, while diffuse-porous taxa were less vulnerable and transpiration increased nearly linearly with D. These results correspond well to patterns in the distribution of the taxa, such as the prevalence of diffuse-porous species in riparian ecosystems, and also provide a means of representing maximum transpiration rates at varying D in broad categories of trees.

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.

Calibration of thermal dissipation sap flow probes for ring- and diffuse-porous trees

Thermal dissipation probes (the Granier method) are routinely used in forest ecology and water balance studies to estimate whole-tree transpiration. This method utilizes an empirically derived equation to measure sap flux density, which has been reported as independent of wood characteristics. However, errors in calculated sap flux density may occur when large gradients in sap velocity occur along the sensor length or when sensors are inserted into non-conducting wood. These may be conditions routinely associated with ring-porous species, yet there are few cases in which the original calibration has been validated for ring-porous species. We report results from laboratory calibration measurements conducted on excised stems of four ring-porous species and two diffuse-porous species. Our calibration results for ring-porous species were considerably different compared with the original calibration equation. Calibration equation coefficients obtained in this study differed by as much as two to almost three orders of magnitude when compared with the original equation of Granier. Coefficients also differed between ring-porous species across all pressure gradient conditions considered; however, no differences between calibration slopes were observed for data collected within the range of expected in situ pressure gradients. In addition, dye perfusions showed that in three of the four ring-porous species considered, active sapwood was limited to the outermost growth ring. In contrast, our calibration results for diffuse-porous species showed generally good agreement with the empirically derived Granier calibration, and dye perfusions showed that active sapwood was associated with many annual growth rings. Our results suggest that the original calibration of Granier is not universally applicable to all species and xylem types and that previous estimates of absolute rates of water use for ring-porous species obtained using the original calibration coefficients may be associated with substantial error.

Population structure, physiology and ecohydrological impacts of dioecious riparian tree species of western North America

The global water cycle is intimately linked to vegetation structure and function. Nowhere is this more apparent than in the arid west where riparian forests serve as ribbons of productivity in otherwise mostly unproductive landscapes. Dioecy is common among tree species that make up western North American riparian forests. There are intrinsic physiological differences between male and female dioecious riparian trees that may influence population structure (i.e., the ratio of male to female trees) and impact ecohydrology at large scales. In this paper, we review the current literature on sex ratio patterns and physiology of dioecious riparian tree species. Then develop a conceptual framework of the mechanisms that underlie population structure of dominant riparian tree species. Finally, we identify linkages between population structure and ecohydrological processes such as evapotranspiration and streamflow. A more thorough understanding of the mechanisms that underlie population structure of dominant riparian tree species will enable us to better predict global change impacts on vegetation and water cycling at multiple scales.

Ecophysiology of riparian cottonwood and willow before, during, and after two years of soil water removal

Riparian cottonwood/willow forest assemblages are highly valued in the southwestern United States for their wildlife habitat, biodiversity, and watershed protection. Yet these forests are under considerable threat from climate change impacts on water resources and land-use activities to support human enterprise. Stream diversions, groundwater pumping, and extended drought have resulted in the decline of cottonwood/willow forests along many riparian corridors in the Southwest and, in many cases, the replacement of these forests with less desirable invasive shrubs and trees. Nevertheless, ecophysiological responses of cottonwood and willow, along with associated ecohydrological feedbacks of soil water depletion, are not well understood. Ecophysiological processes of mature Fremont cottonwood and coyote willow stands were examined over four consecutive growing seasons (2004-2007) near Salt Lake City, Utah, USA. The tree stands occurred near the inlet of a reservoir that was drained in the spring of 2005 and remained empty until mid-summer of 2006, effectively removing the primary water source for most of two growing seasons. Stem sap flux density (Js) in cottonwood was highly correlated with volumetric soil moisture (theta) in the upper 60 cm and decreased sevenfold as soil moisture dropped from 12% to 7% after the reservoir was drained. Conversely, Js in willow was marginally correlated with 0 and decreased by only 25% during the same period. Opposite patterns emerged during the following growing season: willow had a lower whole-plant conductance (kt) in June and higher leaf carbon isotope ratios (delta13C) than cottonwood in August, whereas k(t) and delta13C were otherwise similar between species. Water relations in both species recovered quickly from soil water depletion, with the exception that sapwood area to stem area (As:Ast) was significantly lower in both species after the 2007 growing season compared to 2004. Results suggest that cottonwood has a greater sensitivity to interannual reductions in water availability, while willow is more sensitive to longer periods of soil water depletion. These data shed light on the linkage between soil water deficits and ecophysiological processes of threatened riparian forests given potential land-use and long-term drought impacts on freshwater resources.

Effect of gender on sap‐flux‐scaled transpiration in a dominant riparian tree species: Box elder (Acer negundo)

[1] Acer negundo is a dioecious riparian tree species with a spatial segregation of the sexes along soil moisture gradients. Females are typically more common in wet sites along streams (typically F/M � 1.6), whereas males are more common in drier sites away from streams (typically F/M � 0.6). Spatial segregation between sexes may develop because of the higher reproductive cost in females compared to males. If so, female Acer negundo trees would be under stronger selection to maximize resource uptake, and would therefore likely occur at greater frequencies in high resources sites (i.e., along streamsides), and increase rates of resource acquisition (i.e., water and nutrients). The spatial segregation of the sexes leads to the hypothesis that male and female individuals have varying influence on ecosystem evapotranspiration. To address this, stem sap flux was measured on mature streamside (� 1 m from stream channel) and nonstreamside (>1 m from stream channel) male and female Acer negundo trees occurring in Red Butte Canyon near Salt Lake City, Utah, during the 2004 growing season. Despite having similar predawn and midday water potentials, sap flux density was 76% higher in streamside female trees than in males (P < 0.0001), while sap flux density was 19% greater in nonstreamside female trees compared to males (P < 0.0001). Mean daily sap flux density of all A. negundo populations was highly correlated with mean daily vapor pressure deficit (P < 0.0001), and was moderately correlated with mean daily photosynthetic active radiation (P = 0.0263). At the watershed scale, nonstreamside male and female A. negundo trees contributed 20 and 21% respectively to the estimated 1.7 mm d � 1 transpiration flux from dominant riparian vegetation away from streamsides (estimated from scaled sap flux measurements of all dominant riparian tree species in Red Butte Canyon). Male and female A. negundo trees contributed 31 and 46% respectively of the estimated 8.0 mm d � 1 transpiration flux from dominant riparian vegetation adjacent to the stream channel. Results from this investigation show that the population structure of dioecious riparian trees has direct consequences on ecosystem ET, particularly along stream margins. Shifts in population structure therefore, may have profound impacts on several ecohydrological processes including stream discharge, biogeochemical cycling, and ecosystem productivity.

Elevated stream inorganic nitrogen impacts on a dominant riparian tree species: Results from an experimental riparian stream system

water from 5 mmol l � 1 to about 100 mmol l � 1 , while the second plot received no additional nitrogen. Relative stem sap flux density (Js) did not vary between plots until near the conclusion of the 2006 growing season, when trees in the fertilized plot showed a steep upswing in Js relative to trees in the control plot. Sap flux in 2007 increased consistently by 0.4% day � 1 in the fertilized plot relative to the control plot over a 75-day period, before leveling off near the conclusion of the growing season. At the onset of the experiment, leaf nitrogen per unit mass and leaf nitrogen per unit area were significantly higher in the control plot, and leaf C:N ratios were lower. In 2007, however, differences in leaf chemistry disappeared, suggesting that leaf nitrogen increased in the fertilized trees relative to the control trees. Stem diametergrowth in 2007 was 15% greater in the fertilized trees, although there were no differences in either canopy radial or canopy height growth throughout the experiment. Results from this investigation suggest that increases in stream inorganic nitrogen affect water use, litter quality, and productivity of dominant riparian vegetation. These effects may have important feedbacks on several ecohydrological processes.