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Coastal Tree-Ring Records for Paleoclimate and Paleoenvironmental Applications in North America

For more than a century, tree-ring research has identified relationships between climatic and ecological conditions and tree growth to describe past environments and constrain future ecosystem vulnerabilities. Tree-ring records are frequently used as environmental proxies that extend knowledge of past climate and ecology on millennial scales. Many of the most pressing global change questions facing North America concern the rate of climate change and vulnerability of ecosystems and society along the coast. The opportunities and applications in dendrochronology continue to grow with advancing methodologies, faster computational ability, and the cost-reduction of many chemical and anatomical analyses. Here, we propose that many pressing global change questions that affect coastal communities can be addressed using dendrochronological techniques. We review coastal tree-ring studies that demonstrate the utility and potential for future tree-ring studies in the northeastern, southeastern, northwestern, and southwestern North American coasts. Additionally, we show that tree-ring chronologies along the coast give insight into local and regional climate phenomena that are distinct from nearby, inland tree-ring chronologies of the same species. Lastly, we identify opportunities for coastal dendrochronology and encourage the collection of more tree-ring records that are directly impacted by coastal phenomena.

Quaternary Science Reviews

Fire history across forest types in the southern Beartooth Mountains, Wyoming

Fire is a critical ecosystem process that has played a key role in shaping forests throughout the Beartooth Mountains in northwestern Wyoming. The highly variable topography of the area provides ideal conditions to compare fire regimes across contiguous forest types, yet pyro-dendrochronological research in this area is limited. We reconstructed fire frequency, tree age structure, and post-fire tree growth response in the Clarks Fork Ranger District of the Shoshone National Forest to infer variations in historical fire behavior and stand effects. We collected fire-scarred trees and plot-based tree ages on plots ranging 0.5-5 km 2 in size across two forest types separated by 2 km: a lower-elevation forest of mixed Douglas-fir and lodgepole pine and a higher elevation treeline forest dominated by whitebark pine. Fires occurred in the lower-elevation forest in 1664, 1706, 1785, 1804, 1846, and 1900 with a mean fire return interval of 47 years. The fires in 1804 and 1900 were also recorded in the higher elevation forest, with significant tree mortality at high elevation in the 1900 fire. Both forests were multi-aged with little evidence of tree cohorts in response to severe, stand-replacing events. On average, tree growth increased after fires, with mean ring widths after fire 39% wider in Douglas-fir and 40% wider in lodgepole pine than pre-fire averages, suggesting that some tree mortality likely occurred in association with lower-elevation forest fires. Burns were more frequent in the lower-elevation forest and were occasionally able to spread into the upper-elevation whitebark stand. Although we suspect the transition of fires from low-to high-elevation occurred during drier years, we did not find any relationship between fire years and available climatic reconstructions via superposed epoch analysis. Regeneration during the 20 th Century in the whitebark forest documents recovery of this forest after the 1900 moderate-severity fire event. Finally, especially in the lower-elevation Douglas-fir forest, the period since the last recorded fire (1900) appears to be longer than any fire-free period in the historical record, suggesting that fire exclusion may be creating changes in landscape and patch-scale stand structures, which will likely impact future fire behavior, especially the extent of crown-replacing fire, in these forests.

Wyoming

Dendrochronology of a rare long-lived mediterranean shrub

Ceanothus verrucosus (CEVE) is a globally rare, long-lived, chaparral shrub endemic to coastal southern California (CA) and northern Mexico. There is concern for CEVE persistence because of habitat loss, fire, and climate change, yet little is known about basic features of the plant, including whether it contains annual rings, plant age, and climate–growth response. Growth-ring analysis was challenging because of semi-ring-porous structure, false, and missing rings. We successfully crossdated CEVE annual rings, primarily from Cabrillo National Monument, CA, using a nearby Pinus torreyana chronology. The oldest living individual had 116 rings; the oldest inner-ring date was 1873; and most of the plants established between 1894 and 1905, all older than previous estimates. CEVE mortality occurred during a dry period from the late 1940s through the early 1960s. Correlations between age and stem measurements were weak to moderate (r = 0.10 to 0.56) posing challenges for field-based estimates of plant ages, which are important for population modeling. Variability in CEVE ring width had a strong positive correlation with prior cool-season (October–April) precipitation, yet 2- to 7-day warm-season precipitation events were recorded as rare false rings in multiple years, indicating extreme plasticity in cambial phenology and growth response to moisture.

California

Characterizing historic streamflow to support drought planning in the upper Missouri River basin

This project combined tree-ring based paleo and modern climate and hydrologic research aimed at understanding the primary influences on drought risk and water reliability in basins critical for western U.S. water resources. New paleohydrologic datasets and analyses were developed and applied to contextualize future streamflow projections and address specific water management questions. These questions centered around optimizing future water management protocols for numerous objectives ranging from improving agricultural water allocation during drought while maintaining instream flows for aquatic ecosystem health, to the testing of operations across large river systems with complex infrastructure critical for downstream flood control, navigation, and hydropower generation. USGS scientists worked closely with the Bureau of Reclamation to estimate both past and future drought risk at key management locations throughout the Missouri basin, the Milk and St. Mary River system, and across the major managed river systems in the western United States. These efforts provided a roadmap for future water management strategies under changing climate and water supply conditions, which are detailed in Reclamation’s newly completed Missouri Headwaters Basin Study, the 2021 SECURE Water Act Report, and the forthcoming update of the St. Mary and Milk Rivers Basin Study. Among the major scientific findings to emerge was a new understanding of the long-term (1200-year) history of drought variability for the Missouri River, which highlighted the unusual severity of the early 2000s drought across the Rocky Mountain headwaters and adjacent high plains. By combining the extended drought record with extensive modern and paleoclimate records, we document how warming exacerbates severities of naturally occurring droughts, with recent decades defined by “hot” droughts and the 2000s (2001-2010) drought ranking as the most severe event in 1200 years. Increasingly severe drought events such as this strain already over-allocated water resources that multiple sectors of society depend heavily upon.

upper Missouri River basin

Streamflow reconstruction in the Upper Missouri River Basin using a novel Bayesian network model

A Bayesian model that uses the spatial dependence induced by the river network topology, and the leading principal components of regional tree-ring chronologies for paleo-streamflow reconstruction is presented. In any river basin, a convergent, dendritic network of tributaries comes together to form the main stem of a river. Consequently, it is natural to think of a spatial Markov process that recognizes this topological structure to develop a spatially consistent basin-scale streamflow reconstruction model that uses the information in streamflow and tree-ring chronology data to inform the reconstructed flows, while maintaining the space-time correlation structure of flows that is critical for water resource assessments and management. Given historical data from multiple streamflow gauges along a river, their tributaries in a watershed, and regional tree-ring chronologies, the model is fit and used to simultaneously reconstruct the full network of paleo-streamflow at all gauges in the basin progressing upstream to downstream along the river. The spatial network structure allows a substantial reduction in the uncertainty associated with paleo-streamflow as one proceeds downstream in the network and the spatial dependence structure increases the information content. Our application to eighteen streamflow gauges in the Upper Missouri River Basin shows that the mean adjusted-R2 for the basin is approximately 0.5 with good overall cross-validated skill as measured by five different skill metrics. A comparison with the traditional principal components regression shows that the spatial Bayesian model offers improvements, as downstream gauges are informed by the reconstruction of the upstream gauges, as well as the tree-ring chronologies.

Idaho, Montana, Wyoming

Potential flood events in the lower Missouri River basin over multiple centuries identified using tree-ring based multi-model streamflow reconstructions

The Missouri River basin (MRB), the largest river basin in the United States, presents major water management challenges due to its complex topology and extensive infrastructure designed to manage high annual flows. Severe hydroclimatic events, particularly floods in 1993, 2011, and 2019, have highlighted vulnerabilities, leading the U.S. Army Corps of Engineers to enhance flood risk assessments in 2023. This study focuses on the full MRB, using tree-ring based streamflow reconstructions to extend the Basin's historic streamflow perspective. We developed annual streamflow reconstructions for the Hermann, Missouri streamgage from water years 1360–1900 using K-nearest neighbor and principal component regression methods. These reconstructions incorporate 183 tree-ring chronologies (1360–2020) and modeled natural streamflow (1901–‍2020). Reconstructions were evaluated for consistency using statistical measures and wavelet analysis. Both methods produced similar results, confirming their reliability in reconstructing multi-century streamflows. This novel methodology, guided by wavelet analysis, was developed to identify potential flood events in the lower Missouri River basin (LMRB), resulting in the identification of 172 potential flood events from water years 1360–2020, with 126 events common to the two versions of the MRB reconstructions. The reconstructed streamflow records showed an increase in the number of events until the late-18th- or mid-19th-century. These findings provide insights into evolving flood characteristics in the LMRB. The novel methodology developed in this study can be used to identify potential flood events from tree-ring based annual streamflow reconstructions and generally be applicable to other river basins.

Colorado, Iowa, Kansas, Missouri, Nebraska, Wyomin

Investigating runoff efficiency in upper Colorado River streamflow over past centuries

With increasing concerns about the impact of warming temperatures on water resources, more attention is being paid to the relationship between runoff and precipitation, or runoff efficiency. Temperature is a key influence on Colorado River runoff efficiency, and warming temperatures are projected to reduce runoff efficiency. Here, we investigate the nature of runoff efficiency in the upper Colorado River (UCRB) basin over the past 400 years, with a specific focus on major droughts and pluvials, and to contextualize the instrumental period. We first verify the feasibility of reconstructing runoff efficiency from tree-ring data. The reconstruction is then used to evaluate variability in runoff efficiency over periods of high and low flow, and its correspondence to a reconstruction of late runoff season UCRB temperature variability. Results indicate that runoff efficiency has played a consistent role in modulating the relationship between precipitation and streamflow over past centuries, and that temperature has likely been the key control. While negative runoff efficiency is most common during dry periods, and positive runoff efficiency during wet years, there are some instances of positive runoff efficiency moderating the impact of precipitation deficits on streamflow. Compared to past centuries, the 20th century has experienced twice as many high flow years with negative runoff efficiency, likely due to warm temperatures. These results suggest warming temperatures will continue to reduce runoff efficiency in wet or dry years, and that future flows will be less than anticipated from precipitation due to warming temperatures.

Colorado River

The severity of the 2014–2015 snow drought in the Oregon Cascades in a multicentury context

The western United States (US) is a hotspot for snow drought. The Oregon Cascade Range is highly sensitive to warming and as a result has experienced the largest mountain snowpack losses in the western US since the mid-20th century, including a record-breaking snow drought in 2014–2015 that culminated in a state of emergency. While Oregon Cascade snowpacks serve as the state's primary water supply, short instrumental records limit water managers' ability to fully constrain long-term natural snowpack variability prior to the influence of ongoing and projected anthropogenic climate change. Here, we use annually-resolved tree-ring records to develop the first multi-century reconstruction of Oregon Cascade April 1st Snow Water Equivalent (SWE). The model explains 58% of observed snowpack variability and extends back to 1688 AD, nearly quintupling the length of the existing snowpack record. Our reconstruction suggests that only one other multiyear event in the last three centuries was as severe as the 2014–2015 snow drought. The 2015 event alone was more severe than nearly any other year in over three centuries. Extreme low-to-high snowpack “whiplash” transitions are a consistent feature throughout the reconstructed record. Multi-decadal intervals of persistent below-the-mean peak SWE are prominent features of pre-instrumental snowpack variability, but are generally absent from the instrumental period and likely not fully accounted for in modern water management. In the face of projected snow drought intensification and warming, our findings motivate adaptive management strategies that address declining snowpack and increasingly variable precipitation regimes.

Oregon

Historical fire regimes and contemporary fire effects within sagebrush habitats of Gunnison Sage-grouse

The historical role of fire in sagebrush ( Artemisia tridentata ) landscapes remains poorly understood, yet is important to inform management and conservation of obligate species such as the threatened Gunnison Sage-grouse (GUSG; Centrocercus minimus ). We reconstructed fire histories from tree-ring fire scars at sagebrush–forest ecotones (10 sites, 111 trees) to better understand the role of fire in sagebrush landscapes of the Upper Gunnison Basin (UGB), Colorado, and how fire may have changed following Euro-American settlement. We assessed likely influences of historical fire by surveying plant composition and structure at 100 sagebrush sites with and without recent (2001–2020) fires. Tree-ring fire scars revealed a history of repeated low-severity fire at sagebrush–forest ecotones until 1892, followed by over a century without fire. Between 1684 and 1892, the mean fire interval (MFI) among sites averaged 49.6 years (ranging from 18.2 to 119 years). Fire over this period occurred synchronously at two or more sites on average every 23.6 years, potentially indicative of spread between sites. Most (70%) of the historical fires burned in the early growing season, consistent with times of strong wind. Recent burns exhibited reductions in sagebrush cover (5% vs. 25% in unburned sites) and concomitant increases in herbaceous cover (55% vs. 40%). These differences declined over time but persisted for at least two decades. Burned sites were dominated by native perennial grasses, forbs, and resprouting shrub species. Historically, such openings may have served as seasonal GUSG habitat. Our results indicate that parts of the UGB sagebrush landscapes were characterized historically by frequent fire and dynamic vegetation mosaics that included open, grassy patches. These findings support the use of prescribed fire to restore and maintain this ecological process and vegetation heterogeneity. However, the contemporary context for fire has changed and now includes substantially reduced, Endangered Species Act (ESA)-listed GUSG populations, increased risk of non-native plant invasion, and climate warming. These circumstances highlight new risks, information needs, and opportunities for key knowledge co-production via management–research partnerships.

Colorado

Paleohydrological context for recent floods and droughts in the Fraser River Basin, British Columbia, Canada

The recent intensification of floods and droughts in the Fraser River Basin (FRB) of British Columbia has had profound cultural, ecological, and economic impacts that are expected to be exacerbated further by anthropogenic climate change. In part due to short instrumental runoff records, the long-term stationarity of hydroclimatic extremes in this major North American watershed remains poorly understood, highlighting the need to use high-resolution paleoenvironmental proxies to inform on past streamflow. Here we use a network of tree-ring proxy records to develop 11 subbasin-scale, complementary flood- and drought-season reconstructions, the first of their kind. The reconstructions explicitly target management-relevant flood and drought seasons within each basin, and are examined in tandem to provide an expanded assessment of extreme events across the FRB with immediate implications for water management. We find that past high flood-season flows have been of greater magnitude and occurred in more consecutive years than during the observational record alone. Early 20th century low flows in the drought season were especially severe in both duration and magnitude in some subbasins relative to recent dry periods. Our Fraser subbasin-scale reconstructions provide long-term benchmarks for the natural flood and drought variability prior to anthropogenic forcing. These reconstructions demonstrate that the instrumental streamflow records upon which current management is based likely underestimate the full natural magnitude, duration, and frequency of extreme seasonal flows in the FRB, as well as the potential severity of future anthropogenically forced events.

British Columbia

Networks of tree-ring based streamflow reconstructions for the Pacific Northwest, U.S.A

Water resources in the Pacific Northwest (PNW) are characterized by significant interannual to interdecadal variation. Paleo-proxy reconstructions such as those derived from tree-rings provide longer-term context and supplement information on this expected range of variability, which can improve planning, management, and response related to extreme events and hydrologic change. However, existing paleo-proxy reconstructions have yet to address the potential for pronounced within- and among-basin variations in the PNW due to a lack of spatial coverage. Here we develop methodologically consistent reconstructions for 36 gages in the PNW, including the Columbia and Snake River drainages, as well as key coastal watersheds. These reconstructions extend back at least to the 1500s coefficient of efficiency. Reconstruction skill is relatively high (mean R 2 = 0.63), and snowpack- or winter precipitation-sensitive chronologies from high-elevation sites provide important contributions to reconstruction skill. At the whole-region scale, reconstructed variability indicates evidence for drier and wetter years, more persistent decadal variability, and correspondingly longer episodes of deficit and surplus compared to instrumental records. Within the region, this expanded range of extremes appears especially prevalent in the Snake River and southern PNW watersheds. Regionally, cumulative deficits in the early 1500s and mid 1600s rival those of the early 20 th century, though persistence and timing vary widely among basins. These reconstructions suggest that considering within-region variability will be key for water management and planning under climate change, and that sub-regional adaptation strategies are likely to be advantageous.

Pacific Northwest

Patterns and drivers of cottonwood mortality in the middle Rio Grande, New Mexico, USA

Riparian ecosystems are some of the most valuable and vulnerable on the planet. Riparian tree mortality is increasing in the western United States, where altered streamflows are combining with warming climate. Between 2011 and 2013, one third of an extensive stand of Populus deltoides var. wislizeni (Rio Grande cottonwood) died along the middle Rio Grande on the Pueblo of Santa Ana in New Mexico. Mortality coincided with a severe drought that followed a decade of decreasing streamflow, but it was heterogeneous, with adjacent patches of dead and live trees. The goal of this research was to determine the drivers of mortality to provide insights into future risks of die-off and potential management interventions. We compared tree age, competition, tree-ring widths, sediment particle size and climate influences between live and dead forest patches in a nested plot design. Live and dead trees had similar age, stand density and particle sizes of shallow sediments. Tree-ring widths had the highest correlations with July–September streamflow (1932–2013). All trees had declining ring growth since 1992, coinciding with declining late summer streamflow. An accelerated decline in growth began in 2002, corresponding to recent warmer droughts. Trees that died had lower ring growth 3 years prior to death and in the mid-1900s. Dead trees also had coarser deep sediments 2.4–3.7 m below ground, suggesting that reduced water holding capacity was an important factor for mortality. Water management to increase streamflow during the late summer, especially during times of extended drought, could reduce mortality risk in the face of projected increasingly warm droughts.

New Mexico

Patterns and correlates of giant sequoia foliage dieback during California’s 2012–2016 hotter drought

Hotter droughts – droughts in which unusually high temperatures exacerbate the effects of low precipitation – are expected to increase in frequency and severity in coming decades, challenging scientists and managers to identify which parts of forested landscapes may be most vulnerable. In 2014, in the middle of California’s historically unprecedented 2012–2016 hotter drought, we noticed apparently drought-induced foliage dieback in giant sequoias ( Sequoiadendron giganteum Lindl. [Buchholz]) in Sequoia and Kings Canyon national parks, California. Characteristics of the dieback were consistent with a controlled process of drought-induced senescence: younger (distal) shoots remained green while older (proximal) shoots were preferentially shed. As part of an ongoing interdisciplinary effort to understand and map sequoia vulnerability to hotter droughts, we reviewed historical records for evidence of previous foliage dieback events, surveyed dieback along trail corridors in eight sequoia groves, and analyzed tree-ring data from a high- and a low-foliage-dieback area. In sharp contrast to the greatly elevated mortality of other coniferous species found at low and middle elevations, we estimate that <1% of sequoias died during the drought. Foliage dieback was notably elevated in 2014 – the most severe single drought year in our 122-year record – but much lower in subsequent years. We found no historical records of similar foliage dieback during previous droughts. Dieback in 2014 was highly variable both within and among groves, ranging from virtually no dieback in some areas to nearly 50% in others. Dieback was highest (1) at low elevations, probably due to higher temperatures, reduced snowpack, and earlier snowmelt; (2) in areas of low adult sequoia densities, which likely reflect intrinsically more stressful sites; and (3) on steep slopes, probably reflecting reduced water availability. Average sequoia ring widths were narrower at the high-dieback than the low-dieback tree-ring site, but for reasons that remain unclear the sites did not differ in their proportional ring-width responses to past droughts. Collectively, our results suggest that giant sequoia vulnerability to hotter droughts may be spatially quite variable, and that at least some of that variability can be explained by metrics related to site water balance. Future research will focus on integrating our results with physiological and remote-sensing data, including tracking sequoias as they recover from the drought.

California

Investigating vegetation responses to underground nuclear explosions through integrated analyses

Vegetation has the potential to respond to underground nuclear explosions, yet these links have not been fully explored. Given the lack of previously described signatures, the changes in vegetation are possibly subtle. The integration of multiple different data streams is potentially a useful approach to improve signal detection. Here, we investigate whether semi-arid vegetation growth patterns responded to eight legacy underground nuclear tests at the Nevada National Security Site in southern Nevada, USA. We tested for spatial and temporal changes in vegetation cover, tree growth patterns, and tree leaf spectral properties using ground-based measurements, including those from tree-rings and hyperspectral surface vegetation reflectance, as well as space-based measurements of Normalized Difference Vegetation Index (NDVI) from Landsat. Multiple data streams suggest a localized (<1.2 km) spatial pattern whereby tree growth is enhanced closer to the source of the underground test relative to sites further away. We also observed a more regional (>1.2–9 km) pattern whereby tree growth is suppressed coincident with a drought beginning 1 year before the 1989 tests, but continuing in the 5 years following the tests, which is anomalous relative to what is expected based on the response of tree growth to previous droughts. Quantification of the relative effects of the tests on vegetation remains a challenge due to the coincident drought and the potential for other disturbances to have impacted tree growth at this time, but the integration of these data reveals a more nuanced growth response than any other one data set indicates alone.

Nevada

LiDAR and paleoseismology solve earthquake mystery in the Pacific Northwest, USA

One of the largest historical earthquakes in the U.S. Pacific Northwest occurred on December 15, 1872 near the south end of Lake Chelan. Lack of recognized surface deformation suggested that the earthquake occurred on a blind, perhaps deep, fault. New LiDAR data revealed a NW-side-up scarp along the north side of Spencer Canyon near Entiat, Washington. Landslides triggered during the earthquake impounded small ponds in Spencer Canyon; the larger of the two landslides obliterated a portion of the scarp. Tree-ring counts show that the oldest trees on each landslide are 130 and 128 years old, and lend credence to the idea that the earthquake triggered the landslides. Trenches across the scarp exposed a NW-dipping thrust fault offsetting young soils and Mesozoic bedrock. Radiocarbon and tree ring data shows that the last fault movement was between 1856 and 1873 CE, and was most likely during the 1872 CE earthquake.

Idaho, Montana, Oregon, Washington

Tree rings reveal unmatched 2nd century drought in the Colorado River Basin

The ongoing 22 year drought in the Upper Colorado River Basin (UCRB) has been extremely severe, even in the context of the longest available tree-ring reconstruction of annual flow at Lees Ferry, Arizona, dating back to 762 CE. While many southwestern drought assessments have been limited to the past 1200 years, longer paleorecords of moisture variability do exist for the UCRB. Here, gridded drought-atlas data in the UCRB domain along with naturalized streamflow data from the instrumental period (1906–2021) are used in a K nearest neighbor (KNN) nonparametric algorithm to develop a streamflow reconstruction for the Lees Ferry gage starting in 1 CE. The reconstruction reveals a 2nd century drought unmatched in severity by the current drought or by well-documented medieval period droughts in the UCRB. Although data are sparse, analysis of individual long tree ring records and other paleoclimatic data also support the occurrence of an exceptional 2nd century drought.

Arizona, Colorado, New Mexico, Utah, Wyoming

Tree-ring derived avalanche frequency and climate associations in a high-latitude, maritime climate

Snow avalanches are a natural hazard in mountainous areas worldwide with severe impacts that include fatalities, damage to infrastructure, disruption to commerce, and landscape disturbance. Understanding long-term avalanche frequency patterns, and associated climate and weather influences, improves our understanding of how climate change may affect avalanche activity. We used dendrochronological techniques to evaluate the historical frequency of large magnitude avalanches (LMAs) in the high-latitude climate of southeast Alaska, United States. We collected 434 cross sections throughout six avalanche paths near Juneau, Alaska. This resulted in 2706 identified avalanche growth disturbances between 1720 and 2018, which allowed us to reconstruct 82 years with LMA activity across three sub-regions. By combining this tree-ring-derived avalanche data set with a suite of climate and atmospheric variables and applying a generalized linear model to fit a binomial regression, we found that February and March precipitation and the Oceanic Niño Index (ONI) were significant predictors of LMA activity in the study area. Specifically, LMA activity occurred during winters with substantial February and March precipitation and neutral or negative (cold) ONI values, while years not characterized by LMAs occur more frequently during warm winters (positive ONI values). Our examination of the climate-avalanche relationship in southeast Alaska sheds light on important climate variables and physical processes associated with LMA years. These results can be used to inform long-term infrastructure planning and avalanche mitigation operations in an urban area, such as Juneau, where critical infrastructure is subject to substantial avalanche hazard.

Alaska

Seasonal and species-level water-use strategies and groundwater dependence in dryland riparian woodlands during extreme drought

Drought-induced groundwater decline and warming associated with climate change are primary threats to dryland riparian woodlands. We used the extreme 2012–2019 drought in southern California as a natural experiment to assess how differences in water-use strategies and groundwater dependence may influence the drought susceptibility of dryland riparian tree species with overlapping distributions. We analyzed tree-ring stable carbon and oxygen isotopes collected from two cottonwood species ( Populus trichocarpa and P . fremontii ) along the semi-arid Santa Clara River. We also modeled tree source water δ 18 O composition to compare with observed source water δ 18 O within the floodplain to infer patterns of groundwater reliance. Our results suggest that both species functioned as facultative phreatophytes that used shallow soil moisture when available but ultimately relied on groundwater to maintain physiological function during drought. We also observed apparent species differences in water-use strategies and groundwater dependence related to their regional distributions. P . fremontii was constrained to more arid river segments and ostensibly used a greater proportion of groundwater to satisfy higher evaporative demand. P . fremontii maintained ∆ 13 C at pre-drought levels up until the peak of the drought, when trees experienced a precipitous decline in ∆ 13 C. This response pattern suggests that trees prioritized maintaining photosynthetic processes over hydraulic safety, until a critical point. In contrast, P . trichocarpa showed a more gradual and sustained reduction in ∆ 13 C, indicating that drought conditions induced stomatal closure and higher water use efficiency. This strategy may confer drought avoidance for P . trichocarpa while increasing its susceptibility to anticipated climate warming.

California