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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. 

Recent evidence has revealed the emergence of a megadrought in southwestern North America since 2000. Megadroughts extend for at least 2 decades, making it challenging to identify such events until they are well established. Here, we examined tree-ring growth and stable isotope ratios in Pinus ponderosa at its driest niche edge to investigate whether trees growing near their aridity limit were sensitive to the megadrought climatic pre-conditions, and were capable of informing predictive efforts. During the decade before the megadrought, trees in four populations revealed increases in the cellulose δ13C content of earlywood, latewood, and false latewood, which, based on past studies are correlated with increased intrinsic water-use efficiency. However, radial growth and cellulose δ18O were not sensitive to pre-megadrought conditions. During the 2 decades preceding the megadrought, at all four sites, the changes in δ13C were caused by the high sensitivity of needle carbon and water exchange to drought trends in key winter months, and for three of the four sites during crucial summer months. Such pre-megadrought physiological sensitivity appears to be unique for trees near their arid range limit, as similar patterns were not observed in trees in ten reference sites located along a latitudinal gradient in the same megadrought domain, despite similar drying trends. Our results reveal the utility of tree-ring δ13C to reconstruct spatiotemporal patterns during the organizational phase of a megadrought, demonstrating that trees near the arid boundaries of a species’ distribution might be useful in the early detection of long-lasting droughts. 

Across forests, photosynthesis and woody growth respond to different climate cues. 

Abstract The US Southwest has been entrenched in a two‐decade‐long megadrought (MD), the most severe since 800 CE, which threatens the long‐term vitality and persistence of regional montane forests. Here, we report that in the face of record low winter precipitation and increasing atmospheric aridity, seasonal activity of the North American Monsoon (NAM) climate system brings sufficient precipitation during the height of the summer to alleviate extreme tree water stress. We studied seasonally resolved, tree‐ring stable carbon isotope ratios across a 57‐year time series (1960–2017) in 17 Ponderosa pine forests distributed across the NAM geographic domain. Our study focused on the isotope dynamics of latewood (LW), which is produced in association with NAM rains. During the MD, populations growing within the core region of the NAM operated at lower intrinsic and higher evaporative water‐use efficiencies (WUE_iand WUE_E, respectively), compared to populations growing in the periphery of the NAM domain, indicating less physiological water stress in those populations with access to NAM moisture. The disparities in water‐use efficiencies in periphery populations are due to a higher atmospheric vapor pressure deficit (VPD) and reduced access to summer soil moisture. The buffering advantage of the NAM, however, is weakening. We observed that since the MD, the relationship between WUE_iand WUE_Ein forests within the core NAM domain is shifting toward a drought response similar to forests on the periphery of the NAM. After correcting for past increases in the atmospheric CO₂concentration, we were able to isolate the LW time‐series responses to climate alone. This showed that the shift in the relation between WUE_iand WUE_Ewas driven by the extreme increases in MD‐associated VPD, with little advantageous influence on stomatal conductance from increases in atmospheric CO₂concentration. 

Abstract Tree‐ring carbon and oxygen isotope ratios have been used to understand past dynamics in forest carbon and water cycling. Recently, this has been possible for different parts of single growing seasons by isolating anatomical sections within individual annual rings. Uncertainties in this approach are associated with correlated climate legacies that can occur at a higher frequency, such as across successive seasons, or a lower frequency, such as across years. The objective of this study was to gain insight into how legacies affect cross‐correlation in the δ¹³C and δ¹⁸O isotope ratios in the earlywood (EW) and latewood (LW) fractions ofPinus ponderosatrees at thirteen sites across a latitudinal gradient influenced by the North American Monsoon (NAM) climate system. We observed that δ¹³C from EW and LW has significant positive cross‐correlations at most sites, whereas EW and LW δ¹⁸O values were cross‐correlated at about half the sites. Using combined statistical and mechanistic models, we show that cross‐correlations in both δ¹³C and δ¹⁸O can be largely explained by a low‐frequency (multiple‐year) mode that may be associated with long‐term climate change. We isolated, and statistically removed, the low‐frequency correlation, which resulted in greater geographical differentiation of the EW and LW isotope signals. The remaining higher‐frequency (seasonal) cross‐correlations between EW and LW isotope ratios were explored using a mechanistic isotope fractionation–climate model. This showed that lower atmospheric vapor pressure deficits associated with monsoon rain increase the EW‐LW differentiation for both δ¹³C and δ¹⁸O at southern sites, compared to northern sites. Our results support the hypothesis that dominantly unimodal precipitation regimes, such as near the northern boundary of the NAM, are more likely to foster cross‐correlations in the isotope signals of EW and LW, potentially due to greater sharing of common carbohydrate and soil water resource pools, compared to southerly sites with bimodal precipitation regimes.