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  1. Globally, intrinsic water-use efficiency (iWUE) has risen dramatically over the past century in concert with increases in atmospheric CO 2 concentration. This increase could be further accelerated by long-term drought events, such as the ongoing multidecadal “megadrought” in the American Southwest. However, direct measurements of iWUE in this region are rare and largely constrained to trees, which may bias estimates of iWUE trends toward more mesic, high elevation areas and neglect the responses of other key plant functional types such as shrubs that are dominant across much of the region. Here, we found evidence that iWUE is increasing in the Southwest at one of the fastest rates documented due to the recent drying trend. These increases were particularly large across three common shrub species, which had a greater iWUE sensitivity to aridity than Pinus ponderosa , a common tree species in the western United States. The sensitivity of both shrub and tree iWUE to variability in atmospheric aridity exceeded their sensitivity to increasing atmospheric [CO 2 ]. The shift to more water-efficient vegetation would be, all else being equal, a net positive for plant health. However, ongoing trends toward lower plant density, diminished growth, and increasing vegetation mortality across the Southwest indicate that this increase in iWUE is unlikely to offset the negative impacts of aridification. 
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  2. null (Ed.)
    While tree rings have enabled interannual examination of the influence of climate on trees, this is not possible for most shrubs. Here, we leverage a multidecadal record of annual foliar carbon isotope ratio collections coupled with 39 y of survey data from two populations of the drought-deciduous desert shrub Encelia farinosa to provide insight into water-use dynamics and climate. This carbon isotope record provides a unique opportunity to examine the response of desert shrubs to increasing temperature and water stress in a region where climate is changing rapidly. Population mean carbon isotope ratios fluctuated predictably in response to interannual variations in temperature, vapor pressure deficit, and precipitation, and responses were similar among individuals. We leveraged the well-established relationships between leaf carbon isotope ratios and the ratio of intracellular to ambient CO 2 concentrations to calculate intrinsic water-use efficiency (iWUE) of the plants and to quantify plant responses to long-term environmental change. The population mean iWUE value increased by 53 to 58% over the study period, much more than the 20 to 30% increase that has been measured in forests [J. Peñuelas, J. G. Canadell, R. Ogaya, Glob. Ecol. Biogeogr. 20, 597–608 (2011)]. Changes were associated with both increased CO 2 concentration and increased water stress. Individuals whose lifetimes spanned the entire study period exhibited increases in iWUE that were very similar to the population mean, suggesting that there was significant plasticity within individuals rather than selection at the population scale. 
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  3. Summary

    While plant δ15N values have been applied to understand nitrogen (N) dynamics, uncertainties regarding intraspecific and temporal variability currently limit their application. We used a 28 yr record of δ15N values from two Mojave Desert populations ofEncelia farinosato clarify sources of population‐level variability.

    We leveraged > 3500 foliar δ15N observations collected alongside structural, physiological, and climatic data to identify plant and environmental contributors to δ15N values. Additional sampling of soils, roots, stems, and leaves enabled assessment of the distribution of soil N content and δ15N, intra‐plant fractionations, and relationships between soil and plant δ15N values.

    We observed extensive within‐population variability in foliar δ15N values and found plant age and foliar %N to be the strongest predictors of individual δ15N values. There were consistent differences between root, stem, and leaf δ15N values (spanningc. 3‰), but plant and bulk soil δ15N values were unrelated.

    Plant‐level variables played a strong role in influencing foliar δ15N values, and interannual relationships between climate and δ15N values were counter to previously recognized spatial patterns. This long‐term record provides insights regarding the interpretation of δ15N values that were not available from previous large‐scale syntheses, broadly enabling more effective application of foliar δ15N values.

     
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  4. Abstract

    The Craig–Gordon type (C–G) leaf water isotope enrichment models assume a homogeneous distribution of enriched water across the leaf surface, despite observations that Δ18O can become increasingly enriched from leaf base to tip. Datasets of this ‘progressive isotope enrichment’ are limited, precluding a comprehensive understanding of (a) the magnitude and variability of progressive isotope enrichment, and (b) how progressive enrichment impacts the accuracy of C–G leaf water model predictions. Here, we present observations of progressive enrichment in two conifer species that capture seasonal and diurnal variability in environmental conditions. We further examine which leaf water isotope models best capture the influence of progressive enrichment on bulk needle water Δ18O. Observed progressive enrichment was large and equal in magnitude across both species. The magnitude of this effect fluctuated seasonally in concert with vapour pressure deficit, but was static in the face of diurnal cycles in meteorological conditions. Despite large progressive enrichment, three variants of the C–G model reasonably successfully predicted bulk needle Δ18O. Our results thus suggest that the presence of progressive enrichment does not impact the predictive success of C–G models, and instead yields new insight regarding the physiological and anatomical mechanisms that cause progressive isotope enrichment.

     
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