Groundwater‐dependent ecosystems are often defined by the presence of deeply rooted phreatophytic plants. When connected to groundwater, phreatophytes in arid regions decouple ecosystem net primary productivity from precipitation, underscoring a disproportionately high biodiversity and exchange of resources relative to surrounding areas. However, groundwater‐dependent ecosystems are widely threatened due to the effects of water diversions, groundwater abstraction, and higher frequencies of episodic drought and heat waves. The resilience of these ecosystems to shifting ecohydrological–climatological conditions will depend largely on the capacity of dominant, phreatophytic plants to cope with dramatic reductions in water availability and increases in atmospheric water demand. This paper disentangles the broad range of hydraulic traits expressed by phreatophytic vegetation to better understand their capacity to survive or even thrive under shifting ecohydrological conditions. We focus on three elements of plant water relations: (a) hydraulic architecture (including root area to leaf area ratios and rooting depth), (b) xylem structure and function, and (c) stomatal regulation. We place the expression of these traits across a continuum of phreatophytic habits from obligate to semi‐obligate to semi‐facultative to facultative. Although many species occupy multiple phreatophytic niches depending on access to groundwater, we anticipate that populations are largely locally adapted to a narrow range of ecohydrological conditions regardless of gene flow across ecohydrological gradients. Consequently, we hypothesize that reductions in available groundwater and increases in atmospheric water demand will result in either (a) stand replacement of obligate phreatophytic species with more facultative species as a function of widespread mortality in highly groundwater‐dependent populations or (b) directional selection in semi‐obligate and semi‐facultative phreatophytes towards the expression of traits associated with highly facultative phreatophytes in the absence of species replacement. Anticipated shifts in the expression of hydraulic traits may have profound impacts on water cycling processes, species assemblages, and habitat structure of groundwater‐dependent woodlands and riparian forests.

1. The coordination of traits from individual organs to whole plants is under strong selection because of environmental constraints on resource acquisition and use. However, the tight coordination of traits may provide underlying mechanisms of how locally adapted plant populations can become maladapted because of climate change. 2. To better understand local adaptation in intraspecific trait coordination, we studied trait variability in the widely distributed foundation tree species, Populus fremontii using a common garden near the mid-elevational point of this species distribution. We examined 28 traits encompassing four spectra: phenology, leaf economic spectrum (LES), whole-tree architecture (Corner's Rule) and wood economic spectrum (WES). 3. Based on adaptive syndrome theory, we hypothesized that trait expression would be coordinated among and within trait spectra, reflecting local adaptation to either exposure to freeze-thaw conditions in genotypes sourced from highelevation populations or exposure to extreme thermal stress in genotypes sourced from low-elevation populations. 4. High-elevation genotypes expressed traits within the phenology and WES that limit frost exposure and tissue damage. Specifically, genotypes sourced from high elevations had later mean budburst, earlier mean budset, higher wood densities, higher bark fractions and smaller xylem vessels than their low-elevation counterparts. Conversely, genotypes sourced from low elevations expressed traits within the LES that prioritized hydraulic efficiency and canopy thermal regulation to cope with extreme heat exposure, including 40% smaller leaf areas, 67% higher stomatal densities and 34% higher mean theoretical maximum stomatal conductance. Lowelevation genotypes also expressed a lower stomatal control over leaf water potentials that subsequently dropped to pressures that could induce hydraulic failure.

Populus fremontii (Fremont cottonwood) is recognized as one of the most important foundation tree species in the southwestern USA and northern Mexico because of its ability to structure communities across multiple trophic levels, drive ecosystem processes and influence biodiversity via genetic-based functional trait variation. However, the areal extent of P. fremontii cover has declined dramatically over the last century due to the effects of surface water diversions, non-native species invasions and more recently climate change. Consequently, P. fremontii gallery forests are considered amongst the most threatened forest types in North America. In this paper, we unify four conceptual areas of genes to ecosystems research related to P. fremontii’s capacity to survive or even thrive under current and future environmental conditions: (i) hydraulic function related to canopy thermal regulation during heat waves; (ii) mycorrhizal mutualists in relation to resiliency to climate change and invasion by the non-native tree/shrub, Tamarix; (iii) phenotypic plasticity as a mechanism for coping with rapid changes in climate; and (iv) hybridization between P. fremontii and other closely related Populus species where enhanced vigour of hybrids may preserve the foundational capacity of Populus in the face of environmental change. We also discuss opportunities to scale these conceptual areas from genes to the ecosystem level via remote sensing. We anticipate that the exploration of these conceptual areas of research will facilitate solutions to climate change with a foundation species that is recognized as being critically important for biodiversity conservation and could serve as a model for adaptive management of arid regions in the southwestern USA and around the world.

Leaf carbon gain optimization in hot environments requires balancing leaf thermoregulation with avoiding excessive water loss via transpiration and hydraulic failure. The tradeoffs between leaf thermoregulation and transpirational water loss can determine the ecological consequences of heat waves that are increasing in frequency and intensity. We evaluated leaf thermoregulation strategies in warm‐ (>40°C maximum summer temperature) and cool‐adapted (<40°C maximum summer temperature) genotypes of the foundation tree species, Populus fremontii, using a common garden near the mid‐elevational point of its distribution. We measured leaf temperatures and assessed three modes of leaf thermoregulation: leaf morphology, midday canopy stomatal conductance and stomatal sensitivity to vapour pressure deficit. Data were used to parameterize a leaf energy balance model to estimate contrasts in midday leaf temperature in warm‐ and cool‐adapted genotypes. Warm‐adapted genotypes had 39% smaller leaves and 38% higher midday stomatal conductance, reflecting a 3.8°C cooler mean leaf temperature than cool‐adapted genotypes. Leaf temperatures modelled over the warmest months were on average 1.1°C cooler in warm‐ relative to cool‐adapted genotypes. Results show that plants adapted to warm environments are predisposed to tightly regulate leaf temperatures during heat waves, potentially at an increased risk of hydraulic failure.