Accurately quantifying evapotranspiration (ET) is essential for modelling regional-scale ecosystem water balances. This study assembled an ET data set estimated from eddy flux and sapflow measurements for 13 ecosystems across a large climatic and management gradient from the United States, China, and Australia. Our objectives were to determine the relationships among monthly measured actual ET (ET), calculated FAO-56 grass reference ET (ET o ), measured precipitation (P), and leaf area index (LAI)-one associated key parameter of ecosystem structure. Results showed that the growing season ET from wet forests was generally higher than ET o while those from grasslands or woodlands in the arid and semi-arid regions were lower than ET o . Second, growing season ET was found to be converged to within š10% of P for most of the ecosystems examined. Therefore, our study suggested that soil water storage in the nongrowing season was important in influencing ET and water yield during the growing season. Lastly, monthly LAI, P, and ET o together explained about 85% of the variability of monthly ET. We concluded that the three variables LAI, P, and ET o , which were increasingly available from remote sensing products and weather station networks, could be used for estimating monthly regional ET dynamics with a reasonable accuracy. Such an empirical model has the potential to project the effects of climate and land management on water resources and carbon sequestration when integrated with ecosystem models.

Comparisons were made among Douglas-fir forest, aspen (broad leaf deciduous) forest and wheatgrass (C 3 ) grassland for ecosystem-level water-use efficiency (WUE). WUE was defined as the ratio of photosynthetic CO 2 assimilation rate and evapotranspiration (ET) rate. The ET data measured by eddy covariance were screened so that they overwhelmingly represented transpiration. The three sites used in this comparison spanned a range of vegetation (plant functional) types and environmental conditions within western Canada. When compared in the relative order Douglas-fir (located on Vancouver Island, BC), aspen (northern Saskatchewan), grassland (southern Alberta), the sites demonstrated a progressive decline in precipitation and a general increase in maximum air temperature and atmospheric saturation deficit (D max ) during the mid-summer. The average ( AE SD) WUE at the grassland site was 2.6 AE 0.7 mmol mol À1 , which was much lower than the average values observed for the two other sites (aspen: 5.4 AE 2.3, Douglasfir: 8.1 AE 2.4). The differences in WUE among sites were primarily because of variation in ET. The highest maximum ET rates were approximately 5, 3.2 and 2.7 mm day À1 for the grassland, aspen and Douglas-fir sites, respectively. There was a strong negative correlation between WUE and D max for all sites. We also made seasonal measurements of the carbon isotope ratio of ecosystem respired CO 2 (d R ) in order to test for the expected correlation between shifts in environmental conditions and changes to the ecosystemintegrated ratio of leaf intercellular to ambient CO 2 concentration (c i /c a ). There was a consistent increase in d R values in the grassland, aspen forest and Douglas-fir forest associated with a seasonal reduction in soil moisture. Comparisons were made between WUE measured using eddy covariance with that calculated based on D and d R measurements. There was excellent agreement between WUE values calculated using the two techniques. Our d R measurements indicated that c i /c a values were quite similar among the Douglas-fir, aspen and grassland sites, despite large variation in environmental conditions among sites. This implied that the shorter-lived grass species had relatively high c i /c a values for the D of their habitat. By contrast, the longer-lived Douglas-fir trees were more conservative in water-use with lower c i /c a values relative to their habitat D. This illustrates the interaction between biological and environmental characteristics influencing ecosystem-level WUE. The strong correlation we observed between the two independent measurements of WUE, indicates that the stable isotope composition of respired CO 2 is a useful ecosystem-scale tool to help study constraints to photosynthesis and acclimation of ecosystems to environmental stress.

Summary1. Drought-induced mortality and regional dieback of woody vegetation are reported from numerous locations around the world. Yet within any one site, predicting which species are most likely to survive global change-type drought is a challenge. 2. We studied the diversity of drought survival traits of a community of 15 woody plant species in a desert-chaparral ecotone. The vegetation was a mix of chaparral and desert shrubs, as well as endemic species that only occur along this margin. This vegetation boundary has large potential for drought-induced mortality because nearly all species are at the edge of their range. 3. Drought survival traits studied were vulnerability to drought-induced xylem cavitation, sapwood capacitance, deciduousness, photosynthetic stems, deep roots, photosynthetic responses to leaf water potential and hydraulic architecture. Drought survival strategies were evaluated as combinations of traits that could be effective in dealing with drought. 4. The large variation in seasonal predawn water potential of leaves and stem xylem ranged from À6Á82 to À0Á29 MPa and À6Á92 to À0Á27 MPa, respectively. The water potential at which photosynthesis ceases ranged from À9Á42 to À3Á44 MPa. Architecture was a determinant of hydraulic traits, with species supporting large leaf area per sapwood area exhibiting high rates of water transport, but also xylem that is vulnerable to drought-induced cavitation. Species with more negative midday leaf water potential during the growing season also showed access to deeper water sources based on hydrogen isotope analysis. 5. Drought survival mechanisms comprised of drought deciduousness, photosynthetic stems, tolerance of low minimum seasonal tissue water potential and vulnerability to drought-induced xylem cavitation thus varied orthogonally among species, and promote a diverse array of drought survival strategies in an arid ecosystem of considerable floristic complexity.

To ascertain whether browsing or hydrologic conditions influence the physiological performance of Salix and whether Salix and graminoids (Carex) use and possibly compete for similar water resources, we quantified the in situ seasonal patterns of plant water and carbon relations over three growing seasons. Our studies were designed to address the physiological factors which may be responsible for poor woody plant regeneration in montane riparian habitats of Rocky Mountain National Park, Colo. As these systems act to insure the delivery of fresh water to downstream users, the maintenance of their integrity is critical. We quantified plant water potentials, instantaneous rates of carbon fixation, leaf carbon isotope discrimination (Δ), leaf nitrogen content and water sources using stable isotopes of water (δO). The carbon and water relations of Salix were significantly affected by winter browsing by elk and in some cases by landscape position with regard to proximity to active streams. Winter browsing of Salix by elk significantly increased summer plant water potentials and integrative measures of gas exchange (Δ), though browsing did not consistently affect instantaneous rates of photosynthesis, leaf nitrogen or the sources of water used by Salix. No effect of experimental manipulations of surface water conditions on Salix physiology was observed, likely due to the mesic nature of our study period. Using a two-member linear mixing model, from δO values we calculated that Salix appears to rely on streams for approximately 80% of its water. In contrast, the graminoid Carex derives almost 50% of its water from rainfall, indicating divergent water source use by these two life forms. Based on these findings, winter browsing by elk improved Salix water balance possibly by altering the shoot to root ratio which in turn leads to higher water potentials and higher degrees of season-long gas exchange, while experimental damming had in general no effect on the physiological performance of Salix plants. In addition, as the water sources of Salix and Carex were significantly different, competition for water may not influence the growth, development, and regeneration of Salix. Thus, under the conditions of our study, herbivory had a positive effect on the physiological performance of Salix, but it is still unclear whether these changes in physiology transcend into improved Salix regeneration and survivorship. However, under drier environmental conditions such as lower snowpacks and lower stream flows, the browsing resistance of Salix and ecosystem regeneration may be greatly hindered because the reliance of Salix on stream water makes it vulnerable to changes in surface water and hydrological conditions.

This paper presents initial investigations of a new approach to monitor ecosystem processes in complex terrain on large scales. Metabolic processes in mountainous ecosystems are poorly represented in current ecosystem monitoring campaigns because the methods used for monitoring metabolism at the ecosystem scale (e.g., eddy covariance) require flat study sites. Our goal was to investigate the potential for using nocturnal down-valley winds (cold air drainage) for monitoring ecosystem processes in mountainous terrain from two perspectives: measurements of the isotopic composition of ecosystem-respired CO2 (delta13C(ER)) and estimates of fluxes of CO2 transported in the drainage flow. To test if this approach is plausible, we monitored the wind patterns, CO2 concentrations, and the carbon isotopic composition of the air as it exited the base of a young (approximately 40 yr-old) and an old (>450 yr-old) steeply sided Douglas-fir watershed. Nocturnal cold air drainage within these watersheds was strong, deep, and occurred on more than 80% of summer nights. The depth of cold air drainage rapidly increased to tower height or greater when the net radiation at the top of the tower approached zero. The carbon isotope composition of CO2 in the drainage system holds promise as an indicator of variation in basin-scale physiological processes. Although there was little vertical variation in CO2 concentration at any point in time, we found that the range of CO2 concentration over a single evening was sufficient to estimate delta 13C(ER) from Keeling plot analyses. The seasonal variation in delta 13C(ER) followed expected trends: during the summer dry season delta 13C(ER) became less negative (more enriched in 13C), but once rain returned in the fall, delta 13C(ER) decreased. However, we found no correlation between recent weather (e.g., vapor pressure deficit) and delta 13C(ER) either concurrently or with up to a one-week lag. Preliminary estimates suggest that the nocturnal CO2 flux advecting past the 28-m tower is a rather small fraction (<20%) of the watershed-scale respiration. This study demonstrates that monitoring the isotopic composition and CO2 concentration of cold air drainage at the base of a watershed provides a new tool for quantifying ecosystem metabolism in mountainous ecosystems on the basin scale.