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Dryland riparian woodlands are considered to be locally buffered from droughts by shallow and stable groundwater levels. However, climate change is causing more frequent and severe drought events, accompanied by warmer temperatures, collectively threatening the persistence of these groundwater dependent ecosystems through a combination of increasing evaporative demand and decreasing groundwater supply. We conducted a dendro-isotopic analysis of radial growth and seasonal (semi-annual) carbon isotope discrimination (Δ13 C) to investigate the response of riparian cottonwood stands to the unprecedented California-wide drought from 2012 to 2019, along the largest remaining free-flowing river in Southern California. Our goals were to identify principal drivers and indicators of drought stress for dryland riparian woodlands, determine their thresholds of tolerance to hydroclimatic stressors, and ultimately assess their vulnerability to climate change. Riparian trees were highly responsive to drought conditions along the river, exhibiting suppressed growth and strong stomatal closure (inferred from reduced Δ13 C) during peak drought years. However, patterns of radial growth and Δ13 C were quite variable among sites that differed in climatic conditions and rate of groundwater decline. We show that the rate of groundwater decline, as opposed to climate factors, was the primary driver of site differences in drought stress, and trees showed greater sensitivity to temperature at sites subjected to faster groundwater decline. Across sites, higher correlation between radial growth and Δ13 C for individual trees, and higher inter-correlation of Δ13 C among trees were indicative of greater drought stress. Trees showed a threshold of tolerance to groundwater decline at 0.5 m year-1 beyond which drought stress became increasingly evident and severe. For sites that exceeded this threshold, peak physiological stress occurred when total groundwater recession exceeded ~3 m. These findings indicate that drought-induced groundwater decline associated with more extreme droughts is a primary threat to dryland riparian woodlands and increases their susceptibility to projected warmer temperatures.

Tree-ring carbon isotope discrimination (Δ13C) and oxygen isotopes (δ18O) collected from white pine (Pinus strobus) trees adjacent to Lake Superior show potential to produce the first winter-specific paleoclimate reconstruction with inter-annual resolution for this region. Isotopic signatures from 1976 to 2015 were strongly linked to antecedent winter minimum temperatures (Tmin), Lake Superior peak ice cover, and regional to continental-scale atmospheric winter pressure variability including the North American Dipole. The immense thermal inertia of Lake Superior underlies the unique connection between winter conditions and tree-ring Δ13C and δ18O signals from the following growing season in trees located near the lake. By combining these signals, we demonstrate feasibility to reconstruct variability in Tmin, ice cover, and continental-scale atmospheric circulation patterns (r ≥ 0.65, P

Summary: We compiled hydrogen and oxygen stable isotope compositions (δ2H and δ18O) of leaf water from multiple biomes to examine variations with environmental drivers. Leaf water δ2H was more closely correlated with δ2H of xylem water or atmospheric vapour, whereas leaf water δ18O was more closely correlated with air relative humidity. This resulted from the larger proportional range for δ2H of meteoric waters relative to the extent of leaf water evaporative enrichment compared with δ18O. We next expressed leaf water as isotopic enrichment above xylem water (Δ2H and Δ18O) to remove the impact of xylem water isotopic variation. For Δ2H, leaf water still correlated with atmospheric vapour, whereas Δ18O showed no such correlation. This was explained by covariance between air relative humidity and the Δ18O of atmospheric vapour. This is consistent with a previously observed diurnal correlation between air relative humidity and the deuterium excess of atmospheric vapour across a range of ecosystems. We conclude that 2H and 18O in leaf water do indeed reflect the balance of environmental drivers differently; our results have implications for understanding isotopic effects associated with water cycling in terrestrial ecosystems and for inferring environmental change from isotopic biomarkers that act as proxies for leaf water. [ABSTRACT FROM AUTHOR]

Climate warming in recent decades has negatively impacted forest health in the western United States. Here, we report on potential early warning signals (EWS) for drought‐related mortality derived from measurements of tree‐ring growth (ring width index; RWI) and carbon isotope discrimination (∆13C), primarily focused on ponderosa pine (Pinus ponderosa). Sampling was conducted in the southern Sierra Nevada Mountains, near the epicenter of drought severity and mortality associated with the 2012–2015 California drought and concurrent outbreak of western pine beetle (Dendroctonus brevicomis). At this site, we found that widespread mortality was presaged by five decades of increasing sensitivity (i.e., increased explained variation) of both tree growth and ∆13C to Palmer Drought Severity Index (PDSI). We hypothesized that increasing sensitivity of tree growth and ∆13C to hydroclimate constitute EWS that indicate an increased likelihood of widespread forest mortality caused by direct and indirect effects of drought. We then tested these EWS in additional ponderosa pine‐dominated forests that experienced varying mortality rates associated with the same California drought event. In general, drier sites showed increasing sensitivity of RWI to PDSI over the last century, as well as higher mortality following the California drought event compared to wetter sites. Two sites displayed evidence that thinning or fire events that reduced stand basal area effectively reversed the trend of increasing hydroclimate sensitivity. These comparisons indicate that reducing competition for soil water and/or decreasing bark beetle host tree density via forest management—particularly in drier regions—may buffer these forests against drought stress and associated mortality risk. EWS such as these could provide land managers more time to mitigate the extent or severity of forest mortality in advance of droughts. Substantial efforts at deploying additional dendrochronological research in concert with remote sensing and forest modeling will aid in forecasting of forest responses to continued climate warming. [ABSTRACT FROM AUTHOR]

Rising atmospheric [CO2], ca, is expected to affect stomatal regulation of leaf gas-exchange of woody plants, thus influencing energy fluxes as well as carbon (C), water, and nutrient cycling of forests. Researchers have proposed various strategies for stomatal regulation of leaf gas-exchange that include maintaining a constant leaf internal [CO2], ci, a constant drawdown in CO2 ( ca − ci), and a constant ci/ ca. These strategies can result in drastically different consequences for leaf gas-exchange. The accuracy of Earth systems models depends in part on assumptions about generalizable patterns in leaf gas-exchange responses to varying ca. The concept of optimal stomatal behavior, exemplified by woody plants shifting along a continuum of these strategies, provides a unifying framework for understanding leaf gas-exchange responses to ca. To assess leaf gas-exchange regulation strategies, we analyzed patterns in ci inferred from studies reporting C stable isotope ratios (δ13C) or photosynthetic discrimination (∆) in woody angiosperms and gymnosperms that grew across a range of ca spanning at least 100 ppm. Our results suggest that much of the ca-induced changes in ci/ ca occurred across ca spanning 200 to 400 ppm. These patterns imply that ca − ci will eventually approach a constant level at high ca because assimilation rates will reach a maximum and stomatal conductance of each species should be constrained to some minimum level. These analyses are not consistent with canalization toward any single strategy, particularly maintaining a constant ci. Rather, the results are consistent with the existence of a broadly conserved pattern of stomatal optimization in woody angiosperms and gymnosperms. This results in trees being profligate water users at low ca, when additional water loss is small for each unit of C gain, and increasingly water-conservative at high ca, when photosystems are saturated and water loss is large for each unit C gain. [ABSTRACT FROM AUTHOR]

The article offers the author's insights on a study published in the periodical "Tree Physiology," focusing on the difference between stem water and leaf cellulose. The author discusses the steady state models of leaf water and cellulose, the oxygen isotope composition of cellulose, the climatic signals imprinted in tree rings, and the impact of environmental variation in the stomatal conductance and photosynthetic assimilation rates.

Cellulose δ18O and δD 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 δ18O, resulting in difficulties partitioning variation in δ18O 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 δ18O or δD. 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 (Δdl) as driven by relative humidity (RH). To demonstrate this concept, we survey studies of leaf water δ18O and δD in hardwood vs. conifer trees. We then apply the concept to cellulose δ18O and δD using a mechanistic model of cellulose δ18O and δD to reconstruct deuterium deviations from the global meteoric water line (δdc) in Quercus macrocarpa, Q. robur, and Pseudotsuga menziesii. For each species, Δdc showed strong correlations with RH across sites. Δdc agreed well with steady-state predictions for Q. macrocarpa, while for Q. robur, the relationship with RH was steeper than expected. The slope of Δdc vs. RH of P. menziesii was also close to steady-state predictions, but δdc were more enriched than predicted. This is in agreement with our leaf water survey showing conifer δdl was more enriched than predicted. Our data reveal that applications of this method should be appropriate for reconstructing RH from cellulose δ18O and δD after accounting for differences between hardwoods and conifers. Hence, δdc 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, Δdc and inferred RH values should help in constraining variation in source water δ18O. [ABSTRACT FROM AUTHOR]

Swiss needle cast ( SNC) is a fungal disease of Douglas-fir ( Pseudotsuga menziesii) that has recently become prevalent in coastal areas of the Pacific Northwest. We used growth measurements and stable isotopes of carbon and oxygen in tree-rings of Douglas-fir and a non-susceptible reference species (western hemlock, Tsuga heterophylla) to evaluate their use as proxies for variation in past SNC infection, particularly in relation to potential explanatory climate factors. We sampled trees from an Oregon site where a fungicide trial took place from 1996 to 2000, which enabled the comparison of stable isotope values between trees with and without disease. Carbon stable isotope discrimination (Δ13 C) of treated Douglas-fir tree-rings was greater than that of untreated Douglas-fir tree-rings during the fungicide treatment period. Both annual growth and tree-ring Δ13 C increased with treatment such that treated Douglas-fir had values similar to co-occurring western hemlock during the treatment period. There was no difference in the tree-ring oxygen stable isotope ratio between treated and untreated Douglas-fir. Tree-ring Δ13 C of diseased Douglas-fir was negatively correlated with relative humidity during the two previous summers, consistent with increased leaf colonization by SNC under high humidity conditions that leads to greater disease severity in following years. [ABSTRACT FROM AUTHOR]

Tree-ring characteristics are commonly used to reconstruct climate variables, but divergence from the assumption of a single biophysical control may reduce the accuracy of these reconstructions. Here, we present data from bur oaks ( Quercus macrocarpa Michx.) sampled within and beyond the current species bioclimatic envelope to identify the primary environmental controls on ring-width indices ( RWIs) and carbon stable isotope discrimination (Δ13 C) in tree-ring cellulose. Variation in Δ13 C and RWI was more strongly related to leaf-to-air vapour pressure deficit ( VPD) at the centre and western edge of the range compared with the northern and wettest regions. Among regions, Δ13 C of tree-ring cellulose was closely predicted by VPD and light responses of canopy-level Δ13 C estimated using a model driven by eddy flux and meteorological measurements ( R2 = 0.96, P = 0.003). RWI and Δ13 C were positively correlated in the drier regions, while they were negatively correlated in the wettest region. The strength and direction of the correlations scaled with regional VPD or the ratio of precipitation to evapotranspiration. Therefore, the correlation strength between RWI and Δ13 C may be used to infer past wetness or aridity from paleo wood by determining the degree to which carbon gain and growth have been more limited by moisture or light. [ABSTRACT FROM AUTHOR]

Despite experimental evidence showing that elevated CO2 levels increase growth in most plants, the isolation of a signal consistent with anthropogenically caused increases in atmospheric CO2 from the dendrochronological record has shown mixed results. Our extensive sets of tree ring data from the Ozark Mountains in Missouri showed that since 1850, Quercus velutina Lam., Quercus coccinea Muench., and Pinus echinata Mill, trees increased in stem growth coincidently with increases in atmospheric CO2. Those long-term increases in radial growth appear unrelated to historical disturbance levels for the region, to long-term changes in relevant climatic variables, or to productivity of sites sampled for the purpose of creating a time sequence of tree ring growth. It is still unclear what the potential role of nitrogen deposition might have been for tree growth. We cross-dated a large number of increment cores and aligned the ring width data by pith date for accurate age constant assessments of growth over the past 150 years. Thus, we circumvented changes in growth trend associated with differences in physiological functioning during development, as well as the need for statistical detrending that removes an unknown degree of long-term .environmental signal, the so called segment length curse that applies to standard dendrochronological investigations. When the positive relationship between CO2 and ring width was examined at different ages, an ontogenetic decline in the rate of growth stimulation was found. Specifically, both the pooled Quercus spp. and P. echinata were characterized by a negative exponential pattern of response over a developmental sequence through age 50. Further knowledge of an intrinsic decline in CO2 sensitivity with tree age or size such as this may be important for increased accuracy in estimating terrestrial carbon stocks across successional landscapes. [ABSTRACT FROM AUTHOR]

We compiled hydrogen and oxygen stable isotope compositions (δ 2 H and δ 18 O) of leaf water from multiple biomes to examine variations with environmental drivers. Leaf water δ 2 H was more closely correlated with δ 2 H of xylem water or atmospheric vapour, whereas leaf water δ 18 O was more closely correlated with air relative humidity. This resulted from the larger proportional range for δ 2 H of meteoric waters relative to the extent of leaf water evaporative enrichment compared with δ 18 O. We next expressed leaf water as isotopic enrichment above xylem water (Δ 2 H and Δ 18 O) to remove the impact of xylem water isotopic variation. For Δ 2 H, leaf water still correlated with atmospheric vapour, whereas Δ 18 O showed no such correlation. This was explained by covariance between air relative humidity and the Δ 18 O of atmospheric vapour. This is consistent with a previously observed diurnal correlation between air relative humidity and the deuterium excess of atmospheric vapour across a range of ecosystems. We conclude that 2 H and 18 O in leaf water do indeed reflect the balance of environmental drivers differently; our results have implications for understanding isotopic effects associated with water cycling in terrestrial ecosystems and for inferring environmental change from isotopic biomarkers that act as proxies for leaf water., (© 2022 The Authors New Phytologist © 2022 New Phytologist Foundation.)

To test tree growth sensitivity to temperature under different ambient CO 2 concentrations, we determined stem radial growth rates as they relate to variation in temperature during the last deglacial period, and compare these to modern tree growth rates as they relate to spatial variation in temperature across the modern species distributional range. Paleo oaks were sampled from Northern Missouri, USA and compared to a pollen-based, high-resolution paleo temperature reconstruction from Northern Illinois, USA. Growth data were from 53 paleo bur oak log cross sections collected in Missouri. These oaks were preserved in river and stream sediments and were radiocarbon-dated to a period of rapid climate change during the last deglaciation (10.5 and 13.3 cal kyr BP). Growth data from modern bur oaks were obtained from increment core collections paired with USDA Forest Service Forest Inventory and Analysis data collected across the Great Plains, Midwest, and Upper Great Lakes regions. For modern oaks growing at an average [CO 2 ] of 330 ppm, growth sensitivity to temperature (i.e., the slope of growth rate versus temperature) was about twice that of paleo oaks growing at an average [CO 2 ] of 230 ppm. These data help to confirm that leaf-level predictions that photosynthesis and thus growth will be more sensitive to temperature at higher [CO 2 ] in mature trees-suggesting that tree growth forest productivity will be increasingly sensitive to temperature under projected global warming and high-[CO 2 ] conditions.

Rising atmospheric [CO2 ], ca , is expected to affect stomatal regulation of leaf gas-exchange of woody plants, thus influencing energy fluxes as well as carbon (C), water, and nutrient cycling of forests. Researchers have proposed various strategies for stomatal regulation of leaf gas-exchange that include maintaining a constant leaf internal [CO2 ], ci , a constant drawdown in CO2 (ca  - ci ), and a constant ci /ca . These strategies can result in drastically different consequences for leaf gas-exchange. The accuracy of Earth systems models depends in part on assumptions about generalizable patterns in leaf gas-exchange responses to varying ca . The concept of optimal stomatal behavior, exemplified by woody plants shifting along a continuum of these strategies, provides a unifying framework for understanding leaf gas-exchange responses to ca . To assess leaf gas-exchange regulation strategies, we analyzed patterns in ci inferred from studies reporting C stable isotope ratios (δ(13) C) or photosynthetic discrimination (∆) in woody angiosperms and gymnosperms that grew across a range of ca spanning at least 100 ppm. Our results suggest that much of the ca -induced changes in ci /ca occurred across ca spanning 200 to 400 ppm. These patterns imply that ca  - ci will eventually approach a constant level at high ca because assimilation rates will reach a maximum and stomatal conductance of each species should be constrained to some minimum level. These analyses are not consistent with canalization toward any single strategy, particularly maintaining a constant ci . Rather, the results are consistent with the existence of a broadly conserved pattern of stomatal optimization in woody angiosperms and gymnosperms. This results in trees being profligate water users at low ca , when additional water loss is small for each unit of C gain, and increasingly water-conservative at high ca , when photosystems are saturated and water loss is large for each unit C gain., (© 2015 John Wiley & Sons Ltd.)

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.

Swiss needle cast (SNC) is a fungal disease of Douglas-fir (Pseudotsuga menziesii) that has recently become prevalent in coastal areas of the Pacific Northwest. We used growth measurements and stable isotopes of carbon and oxygen in tree-rings of Douglas-fir and a non-susceptible reference species (western hemlock, Tsuga heterophylla) to evaluate their use as proxies for variation in past SNC infection, particularly in relation to potential explanatory climate factors. We sampled trees from an Oregon site where a fungicide trial took place from 1996 to 2000, which enabled the comparison of stable isotope values between trees with and without disease. Carbon stable isotope discrimination (Δ(13)C) of treated Douglas-fir tree-rings was greater than that of untreated Douglas-fir tree-rings during the fungicide treatment period. Both annual growth and tree-ring Δ(13)C increased with treatment such that treated Douglas-fir had values similar to co-occurring western hemlock during the treatment period. There was no difference in the tree-ring oxygen stable isotope ratio between treated and untreated Douglas-fir. Tree-ring Δ(13)C of diseased Douglas-fir was negatively correlated with relative humidity during the two previous summers, consistent with increased leaf colonization by SNC under high humidity conditions that leads to greater disease severity in following years., (© 2013 John Wiley & Sons Ltd.)

Co-occurring species often have different strategies for tolerating daily cycles of water stress. One underlying parameter that can link together the suite of traits that enables a given strategy is wood density. Here we compare hydraulic traits of two pioneer species from a tropical forest in Panama that differ in wood density: Miconia argentea and Anacardium excelsum. As hypothesized, the higher wood density of Miconia was associated with smaller diameter vessels and fibres, more water stress-resistant leaves and stems, and roughly half the capacitance of the lower wood density Anacardium. However, the scaling of hydraulic parameters such as the increases in leaf area and measures of hydraulic conductivity with stem diameter was remarkably similar between the two species. The collection of traits exhibited by Miconia allowed it to tolerate more water stress than Anacardium, which relied more heavily on its capacitance to buffer daily water potential fluctuations. This work demonstrates the importance of examining a range of hydraulic traits throughout the plant and highlights the spectrum of possible strategies for coping with daily and seasonal water stress cycles., (© 2011 Blackwell Publishing Ltd.)