Interactions between beech and oak seedlings can modify the effects of hotter droughts and the onset of hydraulic failure.

Mixing species with contrasting resource use strategies could reduce forest vulnerability to extreme events. Yet, how species diversity affects seedling hydraulic responses to heat and drought, including mortality risk, is largely unknown. Using open-top chambers, we assessed how, over several years, species interactions (monocultures vs mixtures) modulate heat and drought impacts on the hydraulic traits of juvenile European beech and pubescent oak. Using modeling, we estimated species interaction effects on timing to drought-induced mortality and the underlying mechanisms driving these impacts. We show that mixtures mitigate adverse heat and drought impacts for oak (less negative leaf water potential, higher stomatal conductance, and delayed stomatal closure) but enhance them for beech (lower water potential and stomatal conductance, narrower leaf safety margins, faster tree mortality). Potential underlying mechanisms include oak's larger canopy and higher transpiration, allowing for quicker exhaustion of soil water in mixtures. Our findings highlight that diversity has the potential to alter the effects of extreme events, which would ensure that some species persist even if others remain sensitive. Among the many processes driving diversity effects, differences in canopy size and transpiration associated with the stomatal regulation strategy seem the primary mechanisms driving mortality vulnerability in mixed seedling plantations.

Predicting key water stress indicators of Eucalyptus viminalis and Callitris rhomboidea using high-resolution visible to short-wave infrared spectroscopy.

Drought is one of the main factors contributing to tree mortality worldwide and drought events are set to become more frequent and intense in the face of a changing climate. Quantifying water stress of forests is crucial in predicting and understanding their vulnerability to drought-induced mortality. Here, we explore the use of high-resolution spectroscopy in predicting water stress indicators of two native Australian tree species, Callitris rhomboidea and Eucalyptus viminalis. Specific spectral features and indices derived from leaf-level spectroscopy were assessed as potential proxies to predict leaf water potential (Ψleaf), equivalent water thickness (EWT) and fuel moisture content (FMC) in a dedicated laboratory experiment. New spectral indices were identified that enabled very high confidence linear prediction of Ψleaf for both species (R2 > 0.85) with predictive capacity increasing when accounting for a breakpoint in the relationships using segmented regression (E. viminalis, R2 > 0.89; C. rhomboidea, R2 > 0.87). EWT and FMC were also linearly predicted to a high accuracy (E. viminalis, R2 > 0.90; C. rhomboidea, R2 > 0.80). This study highlights the potential of spectroscopy as a tool for predicting measures of plant water noninvasively, enabling broader applications for monitoring and managing plant water stress.

Leaf water relations determine the trade-off between ozone resistance and stomatal functionality in urban tree species.

Urban trees possess different capacities to mitigate ozone (O3) pollution through stomatal uptake. Stomatal closure protects trees from oxidative damage but limits their growth. To date, it is unclear how plant hydraulic function affect stomatal behaviour and determine O3 resistance. We assessed gas exchange and hydraulic traits in three subtropical urban tree species, Celtis sinensis, Quercus acutissima, and Q. nuttallii, under nonfiltered ambient air (NF) and elevated O3 (NF60). NF60 decreased photosynthetic rate (An) and stomatal conductance (gs) only in Q. acutissima and Q. nuttallii. Maintained An in C. sinensis suggested high O3 resistance and was attributed to higher leaf capacitance at the full turgor. However, this species exhibited a reduced stomatal sensitivity to vapour pressure deficit and an increased minimal gs under NF60. Such stomatal dysfunction did not decrease intrinsic water use efficiency (WUE) due to a tight coupling of An and gs. Conversely, Q. acutissima and Q. nuttallii showed maintained stomatal sensitivity and increased WUE, primarily correlated with gs and leaf water relations, including relative water content and osmotic potential at turgor loss point. Our findings highlight a trade-off between O3 resistance and stomatal functionality, with efficient stomatal control reducing the risk of hydraulic failure under combined stresses.

The sound of lichens: ultrasonic acoustic emissions during desiccation question cavitation events in the hyphae.

Lichens are a mutualistic symbiosis between a fungus and one or more photosynthetic partners. They are photosynthetically active during desiccation down to relative water contents (RWCs) as low as 30% (on dry mass). Experimental evidence suggests that during desiccation, the photobionts have a higher hydration level than the surrounding fungal pseudo-tissues. Explosive cavitation events in the hyphae might cause water movements towards the photobionts. This hypothesis was tested in two foliose lichens by measurements of ultrasonic acoustic emissions (UAEs), a method commonly used in vascular plants but never in lichens, and by measurements of PSII efficiency, water potential, and RWC. Thallus structural changes were characterized by low-temperature scanning electron microscopy. The thalli were silent between 380% and 30% RWCs, when explosive cavitation events should cause movements of liquid water. Nevertheless, the thalli emitted UAEs at ~5% RWC. Accordingly, the medullary hyphae were partially shrunken at ~15% RWC, whereas they were completely shrunken at <5% RWC. These results do not support the hypothesis of hyphal cavitation and suggest that the UAEs originate from structural changes at hyphal level. The shrinking of hyphae is proposed as an adaptation to avoid cell damage at very low RWCs.

Drought response strategies of vascular epiphytes in isolated pasture trees in a Costa Rican tropical montane landscape.

PREMISE: Vascular epiphytes of tropical montane cloud forests are vulnerable to climate change, particularly as cloud bases elevate and reduce atmospheric inputs to the system. However, studies have generally focused on epiphytes in contiguous forests, with little research being done on epiphytes on isolated pasture trees. We investigated water relations of pasture-tree epiphytes at three sites located below and above the elevation of the average cloud base in Monteverde, Costa Rica. METHODS: We measured sap velocity and four microclimate variables in both the dry and wet season of 2018. We also measured functional traits, including pressure volume (PV) curves, predawn/midday water potential, and various lab-based water relations traits. We used linear mixed models to assess the correlation between microclimate and sap velocity in both seasons and ANOVA to assess the variation in PV curve and water potential variables. RESULTS: The turgor loss point generally increased from the wettest to driest site. However, this trend was driven primarily by the increasing prevalence of leaf succulence at drier sites. Microclimatic variables correlated strongly with sap velocity in the wet season, but low soil moisture availability caused this correlation to break down during the dry season. CONCLUSIONS: Our results emphasize the vulnerability of cloud forest epiphytes to rising cloud bases. This vulnerability may be more severe in pasture trees that lack the potential buffer of surrounding forest, but additional research that directly compares the canopy microclimate conditions between forest and pasture trees is needed to confirm this possibility.

Incorporating pressure-volume traits into the leaf economics spectrum.

In recent years, attempts have been made in linking pressure-volume parameters and the leaf economics spectrum to expand our knowledge of the interrelationships among leaf traits. We provide theoretical and empirical evidence for the coordination of the turgor loss point and associated traits with net CO2 assimilation (An ) and leaf mass per area (LMA). We measured gas exchange, pressure-volume curves and leaf structure in 45 ferns and angiosperms, and explored the anatomical and chemical basis of the key traits. We propose that the coordination observed between mass-based An , capacitance and the turgor loss point (πtlp ) emerges from their shared link with leaf density (one of the components of LMA) and, specially, leaf saturated water content (LSWC), which in turn relates to cell size and nitrogen and carbon content. Thus, considering the components of LMA and LSWC in ecophysiological studies can provide a broader perspective on leaf structure and function.

Limits of thermal and hydrological tolerance in a foundation tree species (Populus fremontii) in the desert southwestern United States.

Populus fremontii is among the most dominant, and ecologically important riparian tree species in the western United States and can thrive in hyper-arid riparian corridors. Yet, P. fremontii forests have rapidly declined over the last decade, particularly in places where temperatures sometimes exceed 50°C. We evaluated high temperature tolerance of leaf metabolism, leaf thermoregulation, and leaf hydraulic function in eight P. fremontii populations spanning a 5.3°C mean annual temperature gradient in a well-watered common garden, and at source locations throughout the lower Colorado River Basin. Two major results emerged. First, despite having an exceptionally high Tcrit (the temperature at which Photosystem II is disrupted) relative to other tree taxa, recent heat waves exceeded Tcrit , requiring evaporative leaf cooling to maintain leaf-to-air thermal safety margins. Second, in midsummer, genotypes from the warmest locations maintained lower midday leaf temperatures, a higher midday stomatal conductance, and maintained turgor pressure at lower water potentials than genotypes from more temperate locations. Taken together, results suggest that under well-watered conditions, P. fremontii can regulate leaf temperature below Tcrit along the warm edge of its distribution. Nevertheless, reduced Colorado River flows threaten to lower water tables below levels needed for evaporative cooling during episodic heat waves.

Xylem resistance to cavitation increases during summer in Pinus halepensis.

Cavitation resistance has often been viewed as a relatively static trait, especially for stems of forest trees. Meanwhile, other hydraulic traits, such as turgor loss point (Ψtlp ) and xylem anatomy, change during the season. In this study, we hypothesized that cavitation resistance is also dynamic, changing in coordination with Ψtlp . We began with a comparison of optical vulnerability (OV), microcomputed tomography (µCT) and cavitron methods. All three methods significantly differed in the slope of the curve,Ψ12 and Ψ88 , but not in Ψ50 (xylem pressures that cause 12%, 88%, 50% cavitation, respectively). Thus, we followed the seasonal dynamics (across 2 years) of Ψ50 in Pinus halepensis under Mediterranean climate using the OV method. We found that Ψ50 is a plastic trait with a reduction of approximately 1 MPa from the end of the wet season to the end of the dry season, in coordination with the dynamics of the midday xylem water potential (Ψmidday ) and the Ψtlp . The observed plasticity enabled the trees to maintain a stable positive hydraulic safety margin and avoid cavitation during the long dry season. Seasonal plasticity is vital for understanding the actual risk of cavitation to plants and for modeling species' ability to tolerate harsh environments.

Rapid leaf xylem acclimation diminishes the chances of embolism in grapevines.

Under most conditions tight stomatal regulation in grapevines (Vitis vinifera) avoids xylem embolism. The current study evaluated grapevine responses to challenging scenarios that might lead to leaf embolism and consequential leaf damage. We hypothesized that embolism would occur if the vines experienced low xylem water potential (Ψx) shortly after bud break or later in the season under a combination of extreme drought and heat. We subjected vines to two potentially dangerous environments: (i) withholding irrigation from a vineyard grown in a heatwave-prone environment, and (ii) subjecting potted vines to terminal drought 1 month after bud break. In the field experiment, a heatwave at the beginning of August resulted in leaf temperatures over 45 °C. However, effective stomatal response maintained the xylem water potential (Ψx) well above the embolism threshold, and no leaf desiccation was observed. In the pot experiment, leaves of well-watered vines in May were relatively vulnerable to embolism with 50% embolism (P50) at -1.8 MPa. However, when exposed to drought, these leaves acclimated their leaf P50 by 0.65 MPa in less than a week and before reaching embolism values. When dried to embolizing Ψx, the leaf damage proportion matched (percentage-wise) the leaf embolism level. Our findings indicate that embolism and leaf damage are usually avoided by the grapevines' efficient stomatal regulation and rapid acclimation of their xylem vulnerability.

The ecosystem wilting point defines drought response and recovery of a Quercus-Carya forest.

Soil and atmospheric droughts increasingly threaten plant survival and productivity around the world. Yet, conceptual gaps constrain our ability to predict ecosystem-scale drought impacts under climate change. Here, we introduce the ecosystem wilting point (ΨEWP ), a property that integrates the drought response of an ecosystem's plant community across the soil-plant-atmosphere continuum. Specifically, ΨEWP defines a threshold below which the capacity of the root system to extract soil water and the ability of the leaves to maintain stomatal function are strongly diminished. We combined ecosystem flux and leaf water potential measurements to derive the ΨEWP of a Quercus-Carya forest from an "ecosystem pressure-volume (PV) curve," which is analogous to the tissue-level technique. When community predawn leaf water potential (Ψpd ) was above ΨEWP (=-2.0 MPa), the forest was highly responsive to environmental dynamics. When Ψpd fell below ΨEWP , the forest became insensitive to environmental variation and was a net source of carbon dioxide for nearly 2 months. Thus, ΨEWP is a threshold defining marked shifts in ecosystem functional state. Though there was rainfall-induced recovery of ecosystem gas exchange following soaking rains, a legacy of structural and physiological damage inhibited canopy photosynthetic capacity. Although over 16 growing seasons, only 10% of Ψpd observations fell below ΨEWP , the forest is commonly only 2-4 weeks of intense drought away from reaching ΨEWP , and thus highly reliant on frequent rainfall to replenish the soil water supply. We propose, based on a bottom-up analysis of root density profiles and soil moisture characteristic curves, that soil water acquisition capacity is the major determinant of ΨEWP , and species in an ecosystem require compatible leaf-level traits such as turgor loss point so that leaf wilting is coordinated with the inability to extract further water from the soil.

Aridity-dependent sequence of water potentials for stomatal closure and hydraulic dysfunctions in woody plants.

The sequence of physiological events during drought strongly impacts plants' overall performance. Here, we synthesized the global data of stomatal and hydraulic traits in leaves and stems of 202 woody species to evaluate variations in the water potentials for key physiological events and their sequence along the climatic gradient. We found that the seasonal minimum water potential, turgor loss point, stomatal closure point, and leaf and stem xylem vulnerability to embolism were intercorrelated and decreased with aridity, indicating that water stress drives trait co-selection. In xeric regions, the seasonal minimum water potential occurred at lower water potential than turgor loss point, and the subsequent stomatal closure delayed embolism formation. In mesic regions, however, the seasonal minimum water potential did not pose a threat to the physiological functions, and stomatal closure occurred even at slightly more negative water potential than embolism. Our study demonstrates that the sequence of water potentials for physiological dysfunctions of woody plants varies with aridity, that is, xeric species adopt a more conservative sequence to prevent severe tissue damage through tighter stomatal regulation (isohydric strategy) and higher embolism resistance, while mesic species adopt a riskier sequence via looser stomatal regulation (anisohydric strategy) to maximize carbon uptake at the cost of hydraulic safety. Integrating both aridity-dependent sequence of water potentials for physiological dysfunctions and gap between these key traits into the hydraulic framework of process-based vegetation models would improve the prediction of woody plants' responses to drought under global climate change.

Leaf habit affects the distribution of drought sensitivity but not water transport efficiency in the tropics.

Considering the global intensification of aridity in tropical biomes due to climate change, we need to understand what shapes the distribution of drought sensitivity in tropical plants. We conducted a pantropical data synthesis representing 1117 species to test whether xylem-specific hydraulic conductivity (KS ), water potential at leaf turgor loss (ΨTLP ) and water potential at 50% loss of KS (ΨP50 ) varied along climate gradients. The ΨTLP and ΨP50 increased with climatic moisture only for evergreen species, but KS did not. Species with high ΨTLP and ΨP50 values were associated with both dry and wet environments. However, drought-deciduous species showed high ΨTLP and ΨP50 values regardless of water availability, whereas evergreen species only in wet environments. All three traits showed a weak phylogenetic signal and a short half-life. These results suggest strong environmental controls on trait variance, which in turn is modulated by leaf habit along climatic moisture gradients in the tropics.

Drought acclimation of Quercus ilex leaves improves tolerance to moderate drought but not resistance to severe water stress.

Increasing temperature and drought can result in leaf dehydration and defoliation even in drought-adapted tree species such as the Mediterranean evergreen Quercus ilex L. The stomatal regulation of leaf water potential plays a central role in avoiding this phenomenon and is constrained by a suite of leaf traits including hydraulic conductance and vulnerability, hydraulic capacitance, minimum conductance to water vapour, osmotic potential and cell wall elasticity. We investigated whether the plasticity in these traits may improve leaf tolerance to drought in two long-term rainfall exclusion experiments in Mediterranean forests. Osmotic adjustment was observed to lower the water potential at turgor loss in the rainfall-exclusion treatments, thus suggesting a stomatal closure at more negative water potentials and a more anisohydric behaviour in drier conditions. Conversely, leaf hydraulic conductance and vulnerability did not exhibit any plasticity between treatments so the hydraulic safety margins were narrower in the rainfall-exclusion treatments. The sequence of leaf responses to seasonal drought and dehydration was conserved among treatments and sites but trees were more likely to suffer losses of turgor and hydraulic functioning in the rainfall-exclusion treatments. We conclude that leaf plasticity might help the trees to tolerate moderate drought but not to resist severe water stress.

Projections of leaf turgor loss point shifts under future climate change scenarios.

Predicting the consequences of climate change is of utmost importance to mitigate impacts on vulnerable ecosystems; plant hydraulic traits are particularly useful proxies for predicting functional disruptions potentially occurring in the near future. This study assessed the current and future regional patterns of leaf water potential at turgor loss point (Ψtlp ) by measuring and projecting the Ψtlp of 166 vascular plant species (159 angiosperms and 7 gymnosperms) across a large climatic range spanning from alpine to Mediterranean areas in NE Italy. For angiosperms, random forest models predicted a consistent shift toward more negative values in low-elevation areas, whereas for gymnosperms the pattern was more variable, particularly in the alpine sector (i.e., Alps and Prealps). Simulations were also developed to evaluate the number of threatened species under two Ψtlp plasticity scenarios (low vs. high plasticity), and it was found that in the worst-case scenario approximately 72% of the angiosperm species and 68% of gymnosperms within a location were at risk to exceed their physiological plasticity. The different responses to climate change by specific clades might produce reassembly in natural communities, undermining the resilience of natural ecosystems to climate change.

Trading water for carbon in the future: Effects of elevated CO2 and warming on leaf hydraulic traits in a semiarid grassland.

The effects of climate change on plants and ecosystems are mediated by plant hydraulic traits, including interspecific and intraspecific variability of trait phenotypes. Yet, integrative and realistic studies of hydraulic traits and climate change are rare. In a semiarid grassland, we assessed the response of several plant hydraulic traits to elevated CO2 (+200 ppm) and warming (+1.5 to 3°C; day to night). For leaves of five dominant species (three graminoids and two forbs), and in replicated plots exposed to 7 years of elevated CO2 , warming, or ambient climate, we measured: stomatal density and size, xylem vessel size, turgor loss point, and water potential (pre-dawn). Interspecific differences in hydraulic traits were larger than intraspecific shifts induced by elevated CO2 and/or warming. Effects of elevated CO2 were greater than effects of warming, and interactions between treatments were weak or not detected. The forbs showed little phenotypic plasticity. The graminoids had leaf water potentials and turgor loss points that were 10% to 50% less negative under elevated CO2 ; thus, climate change might cause these species to adjust their drought resistance strategy away from tolerance and toward avoidance. The C4 grass also reduced allocation of leaf area to stomata under elevated CO2 , which helps explain observations of higher soil moisture. The shifts in hydraulic traits under elevated CO2 were not, however, simply due to higher soil moisture. Integration of our results with others' indicates that common species in this grassland are more likely to adjust stomatal aperture in response to near-term climate change, rather than anatomical traits; this contrasts with apparent effects of changing CO2 on plant anatomy over evolutionary time. Future studies should assess how plant responses to drought may be constrained by the apparent shift from tolerance (via low turgor loss point) to avoidance (via stomatal regulation and/or access to deeper soil moisture).

Correlations between leaf economics, mechanical resistance and drought tolerance across 41 cycad species.

BACKGROUND AND AIMS: We conducted a comprehensive analysis of the functional traits of leaves (leaflets) of cycads. The aim of this study was to clarify the functional divergence between the earlier origin Cycadaceae and the later differentiated Zamiaceae, and the differences in trait associations between cycads and angiosperms. METHODS: We selected 20 Cycadaceae species and 21 Zamiaceae species from the same cycad garden in South China, and measured their leaf structure, economic traits, mechanical resistance (Fp) and leaf water potential at the turgor loss point (πtlp). In addition, we compiled a dataset of geographical distribution along with climatic variables for these cycad species, and some leaf traits of tropical-sub-tropical angiosperm woody species from the literature for comparison. KEY RESULTS: The results showed significantly contrasting leaf trait syndromes between the two families, with Zamiaceae species exhibiting thicker leaves, higher carbon investments and greater Fp than Cycadaceae species. Leaf thickness (LT) and πtlp were correlated with mean climatic variables in their native distribution ranges, indicating their evolutionary adaptation to environmental conditions. Compared with the leaves of angiosperms, the cycad leaves were thicker and tougher, and more tolerant to desiccation. Greater Fp was associated with a higher structural investment in both angiosperms and cycads; however, cycads showed lower Fp at a given leaf mass per area or LT than angiosperms. Enhancement of Fp led to more negative πtlp in angiosperms, but the opposite trend was observed in cycads. CONCLUSIONS: Our results reveal that variations in leaf traits of cycads are mainly influenced by taxonomy and the environment of their native range. We also demonstrate similar leaf functional associations in terms of economics, but different relationships with regard to mechanics and drought tolerance between cycads and angiosperms. This study expands our understanding of the ecological strategies and likely responses of cycads to future climate change.

Root pressure-volume curve traits capture rootstock drought tolerance.

BACKGROUND AND AIMS: Living root tissues significantly constrain plant water uptake under drought, but we lack functional traits to feasibly screen diverse plants for variation in the drought responses of these tissues. Water stress causes roots to lose volume and turgor, which are crucial to root structure, hydraulics and growth. Thus, we hypothesized that root pressure-volume (p-v) curve traits, which quantify the effects of water potential on bulk root turgor and volume, would capture differences in rootstock drought tolerance. METHODS: We used a greenhouse experiment to evaluate relationships between root p-v curve traits and gas exchange, whole-plant hydraulic conductance and biomass under drought for eight grapevine rootstocks that varied widely in drought performance in field trials (101-14, 110R, 420A, 5C, 140-Ru, 1103P, Ramsey and Riparia Gloire), grafted to the same scion variety (Vitis vinifera 'Chardonnay'). KEY RESULTS: The traits varied significantly across rootstocks, and droughted vines significantly reduced root turgor loss point (πtlp), osmotic potential at full hydration (πo) and capacitance (C), indicating that roots became less susceptible to turgor loss and volumetric shrinkage. Rootstocks that retained a greater root volume (i.e. a lower C) also maintained more gas exchange under drought. The rootstocks that previous field trials have classified as drought tolerant exhibited significantly lower πtlp, πo and C values in well-watered conditions, but significantly higher πo and πtlp values under water stress, than the varieties classified as drought sensitive. CONCLUSIONS: These findings suggest that acclimation in root p-v curve traits improves gas exchange in persistently dry conditions, potentially through impacts on root hydraulics or root to shoot chemical signalling. However, retaining turgor and volume in previously unstressed roots, as these roots deplete wet soil to moderately negative water potentials, could be more important to drought performance in the deep, highly heterogenous rooting zones which grapevines develop under field conditions.

Hydraulic prediction of drought-induced plant dieback and top-kill depends on leaf habit and growth form.

Hydraulic failure caused by severe drought contributes to aboveground dieback and whole-plant death. The extent to which dieback or whole-plant death can be predicted by plant hydraulic traits has rarely been tested among species with different leaf habits and/or growth forms. We investigated 19 hydraulic traits in 40 woody species in a tropical savanna and their potential correlations with drought response during an extreme drought event during the El Niño-Southern Oscillation in 2015. Plant hydraulic trait variation was partitioned substantially by leaf habit but not growth form along a trade-off axis between traits that support drought tolerance versus avoidance. Semi-deciduous species and shrubs had the highest branch dieback and top-kill (complete aboveground death) among the leaf habits or growth forms. Dieback and top-kill were well explained by combining hydraulic traits with leaf habit and growth form, suggesting integrating life history traits with hydraulic traits will yield better predictions.

Beyond leaf habit: generalities in plant function across 97 tropical dry forest tree species.

Leaf habit has been hypothesized to define a linkage between the slow-fast plant economic spectrum and the drought resistance-avoidance trade-off in tropical forests ('slow-safe vs fast-risky'). However, variation in hydraulic traits as a function of leaf habit has rarely been explored for a large number of species. We sampled leaf and branch functional traits of 97 tropical dry forest tree species from four sites to investigate whether patterns of trait variation varied consistently in relation to leaf habit along the 'slow-safe vs fast-risky' trade-off. Leaf habit explained from 0% to 43.69% of individual trait variation. We found that evergreen and semi-deciduous species differed in their location along the multivariate trait ordination when compared to deciduous species. While deciduous species showed consistent trait values, evergreen species trait values varied as a function of the site. Last, trait values varied in relation to the proportion of deciduous species in the plant community. We found that leaf habit describes the strategies that define drought avoidance and plant economics in tropical trees. However, leaf habit alone does not explain patterns of trait variation, which suggests quantifying site-specific or species-specific uncertainty in trait variation as the way forward.

Mesophyll photosynthetic sensitivity to leaf water potential in Eucalyptus: a new dimension of plant adaptation to native moisture supply.

Photosynthetic sensitivity to drought is a fundamental constraint on land-plant evolution and ecosystem function. However, little is known about how the sensitivity of photosynthesis to nonstomatal limitations varies among species in the context of phylogenetic relationships. Using saplings of 10 Eucalyptus species, we measured maximum CO2 -saturated photosynthesis using A-ci curves at several different leaf water potentials (ψleaf ) to quantify mesophyll photosynthetic sensitivity to ψleaf (MPS), a measure of how rapidly nonstomatal limitations to carbon uptake increase with declining ψleaf . MPS was compared to the macroclimatic moisture availability of the species' native habitats, while accounting for phylogenetic relationships. We found that species native to mesic habitats have greater MPS but higher maximum photosynthetic rates during non-water-stressed conditions, revealing a trade-off between maximum photosynthesis and drought sensitivity. Species with lower turgor loss points have lower MPS, indicating coordination among photosynthetic and water-relations traits. By accounting for phylogenetic relationships among closely related species, we provide the first compelling evidence that MPS in Eucalyptus evolved in an adaptive fashion with climatically determined moisture availability, opening the way for further study of this poorly explored dimension of plant adaptation to drought.