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Plants respond to rapid environmental change in ways that depend on both their genetic identity and their phenotypic plasticity, impacting their survival as well as associated ecosystems. However, genetic and environmental effects on phenotype are difficult to quantify across large spatial scales and through time. Leaf hyperspectral reflectance offers a potentially robust approach to map these effects from local to landscape levels. Using a handheld field spectrometer, we analyzed leaf-level hyperspectral reflectance of the foundation tree species Populus fremontii in wild populations and in three 6-year-old experimental common gardens spanning a steep climatic gradient. First, we show that genetic variation among populations and among clonal genotypes is detectable with leaf spectra, using both multivariate and univariate approaches. Spectra predicted population identity with 100% accuracy among trees in the wild, 87%-98% accuracy within a common garden, and 86% accuracy across different environments. Multiple spectral indices of plant health had significant heritability, with genotype accounting for 10%-23% of spectral variation within populations and 14%-48% of the variation across all populations. Second, we found gene by environment interactions leading to population-specific shifts in the spectral phenotype across common garden environments. Spectral indices indicate that genetically divergent populations made unique adjustments to their chlorophyll and water content in response to the same environmental stresses, so that detecting genetic identity is critical to predicting tree response to change. Third, spectral indicators of greenness and photosynthetic efficiency decreased when populations were transferred to growing environments with higher mean annual maximum temperatures relative to home conditions. This result suggests altered physiological strategies further from the conditions to which plants are locally adapted. Transfers to cooler environments had fewer negative effects, demonstrating that plant spectra show directionality in plant performance adjustments. Thus, leaf reflectance data can detect both local adaptation and plastic shifts in plant physiology, informing strategic restoration and conservation decisions by enabling high resolution tracking of genetic and phenotypic changes in response to climate change., (© 2024 The Author(s). Plant, Cell & Environment published by John Wiley & Sons Ltd.)

Increasing heatwaves are threatening forest ecosystems globally. Leaf thermal regulation and tolerance are important for plant survival during heatwaves, though the interaction between these processes and water availability is unclear. Genotypes of the widely distributed foundation tree species Populus fremontii were studied in a controlled common garden during a record summer heatwave-where air temperature exceeded 48 °C. When water was not limiting, all genotypes cooled leaves 2 to 5 °C below air temperatures. Homeothermic cooling was disrupted for weeks following a 72-h reduction in soil water, resulting in leaf temperatures rising 3 °C above air temperature and 1.3 °C above leaf thresholds for physiological damage, despite the water stress having little effect on leaf water potentials. Tradeoffs between leaf thermal safety and hydraulic safety emerged but, regardless of water use strategy, all genotypes experienced significant leaf mortality following water stress. Genotypes from warmer climates showed greater leaf cooling and less leaf mortality after water stress in comparison with genotypes from cooler climates. These results illustrate how brief soil water limitation disrupts leaf thermal regulation and potentially compromises plant survival during extreme heatwaves, thus providing insight into future scenarios in which ecosystems will be challenged with extreme heat and unreliable soil water access., Competing Interests: Competing interests statement:The authors declare no competing interest.

The succulent plant syndrome is defined by the coordination of traits that enhance internal water storage within plant tissues. Although distributed globally in different habitats, succulent plants are thought to have evolved to avoid drought in arid regions, due to trait modifications that decrease tissue water deficits. We evaluated the evolution and the ecological significance of the succulent strategy at a global scale by comparing the climatic niche of species displaying succulence within the Core Caryophyllales with their non-succulent relatives. We assembled and curated a worldwide dataset of 201K georeferenced records belonging to 5447 species within 28 families, and analyzed the climatic niche of species along with their origin and evolutionary trajectories using ecological niche modeling, phylogenetic regression, divergence dates and ancestral state estimation. Results indicate the Core Caryophyllales have inhabited drylands since their origin in the Early Cretaceous. However, the succulent syndrome appeared and diversified during later geologic periods. The climatic niche space of succulents is narrower than non-succulent relatives, but no climate niche separation was detected between groups. Our results support alternative interpretations on the environmental and ecological forces that spurred the origin and diversification of the succulent plant syndrome and the radiation of rich succulent lineages., (© The Author(s) 2024. Published by Oxford University Press on behalf of the Society for Experimental Biology. All rights reserved. For commercial re-use, please contact reprints@oup.com for reprints and translation rights for reprints. All other permissions can be obtained through our RightsLink service via the Permissions link on the article page on our site—for further information please contact journals.permissions@oup.com.)

Background and Scope: Plant functional traits are the result of natural selection to optimize carbon gain, leading to a broad spectrum of traits across environmental gradients. Among plant traits, leaf water storage capacity is paramount for plant drought resistance. We explored whether leaf-succulent taxa follow trait correlations similar to those of non-leaf-succulent taxa to evaluate whether both are similarly constrained by relationships between leaf water storage and climate., Methods: We tested the relationships among three leaf traits related to water storage capacity and resource use strategies in 132 species comprising three primary leaf types: succulent, sclerophyllous, and leaves with rapid returns on water investment, referred to as fast return. Correlation coefficients among specific leaf area (SLA), water mass per unit of area (WMA), and saturated water content (SWC) were tested, along with relationships between leaf trait spectra and aridity determined from species occurrence records., Results: Both SWC and WMA at a given SLA were ~10-fold higher in succulent leaves than in non-succulent leaves. While SWC actually increased with SLA in non-succulent leaves, no relationship was detected between SWC and SLA in succulent leaves, although WMA decreased with SLA in all leaf types. A principal component analysis (PCA) revealed that succulent taxa occupied a widely different mean trait space than either fast-return (P < 0.0001) or sclerophyllous (P < 0.0001) taxa along the first PCA axis, which explained 63 % of mean trait expression among species. However, aridity only explained 12 % of the variation in PCA1 values. This study is among the first to establish a structural leaf trait spectrum in succulent leaf taxa and quantify contrasts in leaf water storage among leaf types relative to specific leaf area., Conclusions: Trait coordination in succulent leaf taxa may not follow patterns similar to those of widely studied non-succulent taxa., (© The Author(s) 2024. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For commercial re-use, please contact reprints@oup.com for reprints and translation rights for reprints. All other permissions can be obtained through our RightsLink service via the Permissions link on the article page on our site—for further information please contact journals.permissions@oup.com.)

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 T crit (the temperature at which Photosystem II is disrupted) relative to other tree taxa, recent heat waves exceeded T crit , 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 T crit 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., (© 2023 The Authors New Phytologist © 2023 New Phytologist Foundation.)

Background: The plant family Cactaceae provides some of the most striking examples of adaptive evolution, expressing undeniably the most spectacular New World radiation of succulent plants distributed across arid and semi-arid regions of the Americas. Cacti are widely regarded for their cultural, economic and ecological value, yet they are also recognized as one of the most threatened and endangered taxonomic groups on the planet., Scope: This paper reviews current threats to species of cacti that have distributions in arid to semi-arid subtropical regions. Our review focuses primarily on four global change forces: (1) increases in atmospheric CO2 concentrations; (2) increases in mean annual temperatures and heat waves; (3) increases in the duration, frequency and intensity of droughts; and (4) and increases in competition and wildfire frequency from invasion by non-native species. We provide a broad range of potential priorities and solutions for stemming the extinction risk of cacti species and populations., Conclusions: Mitigating ongoing and emerging threats to cacti will require not only strong policy initiatives and international cooperation, but also new and creative approaches to conservation. These approaches include determining species at risk from climate extremes, enhancing habitat quality after disturbance, approaches and opportunities for ex situ conservation and restoration, and the potential use of forensic tools for identifying plants that have been removed illegally from the wild and sold on open markets., (© The Author(s) 2023. Published by Oxford University Press on behalf of the Annals of Botany Company.)

Plant water use theory has largely been developed within a plant-performance paradigm that conceptualizes water use in terms of value for carbon gain and that sits within a neoclassical economic framework. This theory works very well in many contexts but does not consider other values of water to plants that could impact their fitness. Here, we survey a range of alternative hypotheses for drivers of water use and stomatal regulation. These hypotheses are organized around relevance to extreme environments, population ecology, and community ecology. Most of these hypotheses are not yet empirically tested and some are controversial (e.g. requiring more agency and behavior than is commonly believed possible for plants). Some hypotheses, especially those focused around using water to avoid thermal stress, using water to promote reproduction instead of growth, and using water to hoard it, may be useful to incorporate into theory or to implement in Earth System Models., (© 2023 The Authors New Phytologist © 2023 New Phytologist Foundation.)

Selection on quantitative traits by heterogeneous climatic conditions can lead to substantial trait variation across a species range. In the context of rapidly changing environments, however, it is equally important to understand selection on trait plasticity. To evaluate the role of selection in driving divergences in traits and their associated plasticities within a widespread species, we compared molecular and quantitative trait variation in Populus fremontii (Fremont cottonwood), a foundation riparian distributed throughout Arizona. Using SNP data and genotypes from 16 populations reciprocally planted in three common gardens, we first performed Q ST -F ST analyses to detect selection on traits and trait plasticity. We then explored the environmental drivers of selection using trait-climate and plasticity-climate regressions. Three major findings emerged: (1) There was significant genetic variation in traits expressed in each of the common gardens and in the phenotypic plasticity of traits across gardens, both of which were heritable. (2) Based on Q ST -F ST comparisons, there was evidence of selection in all traits measured; however, this result varied from no effect in one garden to highly significant in another, indicating that detection of past selection is environmentally dependent. We also found strong evidence of divergent selection on plasticity across environments for two traits. (3) Traits and/or their plasticity were often correlated with population source climate (R 2 up to .77 and .66, respectively). These results suggest that steep climate gradients across the Southwest have played a major role in shaping the evolution of divergent phenotypic responses in populations and genotypes now experiencing climate change., (© 2022 John Wiley & Sons Ltd.)

Successful conservation of threatened species and ecosystems in a rapidly changing world requires scientifically sound decision-making tools that are readily accessible to conservation practitioners. Physiological applications that examine how plants and animals interact with their environment are now widely used when planning, implementing and monitoring conservation. Among these tools, stable-isotope physiology is a potentially powerful, yet under-utilized cornerstone of current and future conservation efforts of threatened and endangered plants. We review the underlying concepts and theory of stable-isotope physiology and describe how stable-isotope applications can support plant conservation. We focus on stable isotopes of carbon, hydrogen, oxygen and nitrogen to address plant ecophysiological responses to changing environmental conditions across temporal scales from hours to centuries. We review examples from a broad range of plant taxa, life forms and habitats and provide specific examples where stable-isotope analysis can directly improve conservation, in part by helping identify resilient, locally adapted genotypes or populations. Our review aims to provide a guide for practitioners to easily access and evaluate the information that can be derived from stable-isotope signatures, their limitations and how stable isotopes can improve conservation efforts., (© The Author(s) 2022. Published by Oxford University Press and the Society for Experimental Biology.)

Responses of terrestrial ecosystems to climate change have been explored in many regions worldwide. While continued drying and warming may alter process rates and deteriorate the state and performance of ecosystems, it could also lead to more fundamental changes in the mechanisms governing ecosystem functioning. Here we argue that climate change will induce unprecedented shifts in these mechanisms in historically wetter climatic zones, towards mechanisms currently prevalent in dry regions, which we refer to as 'dryland mechanisms'. We discuss 12 dryland mechanisms affecting multiple processes of ecosystem functioning, including vegetation development, water flow, energy budget, carbon and nutrient cycling, plant production and organic matter decomposition. We then examine mostly rare examples of the operation of these mechanisms in non-dryland regions where they have been considered irrelevant at present. Current and future climate trends could force microclimatic conditions across thresholds and lead to the emergence of dryland mechanisms and their increasing control over ecosystem functioning in many biomes on Earth., (© 2022. Springer Nature Limited.)

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., (© 2022 John Wiley & Sons Ltd.)

Rainfall timing, frequency, and quantity is rapidly changing in dryland regions, altering dryland plant communities. Understanding dryland plant responses to future rainfall scenarios is crucial for implementing proactive management strategies, particularly in light of land cover changes concurrent with climate change. One such change is woody plant encroachment, an increasing abundance of woody plants in areas formerly dominated by grasslands or savannas. Continued woody plant encroachment will depend, in part, on seedling capacity to establish and thrive under future climate conditions. Seedling performance is primarily impacted by soil moisture conditions governed by precipitation amount (quantity) and frequency. We hypothesized that (H1) seedling performance would be enhanced by both greater soil moisture and pulse frequency, such that seedlings with similar mean soil moisture would perform best under high pulse frequency. Alternatively, (H2) mean soil moisture would have greater influence than pulse frequency, such that a given pulse frequency would have little influence on seedling performance. The hypotheses were tested with Prosopis velutina, a shrub native to the United States that has encroached throughout its range and is invasive in other continents. Seedlings were grown in a greenhouse under two soil moisture treatments, each which was maintained by two pulse frequency treatments. Contrary to H1, mean soil moisture had greater impact than pulse frequency on seedling growth, photosynthetic gas exchange, leaf chemistry, and biomass allocation. These results indicate that P. velutina seedlings may be more responsive to rainfall amount than frequency, at least within the conditions seedlings experienced in this experimental manipulation., (© 2022. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

Efforts to maintain the function of critical ecosystems under climate change often begin with foundation species. In the southwestern United States, cottonwood trees support diverse communities in riparian ecosystems that are threatened by rising temperatures. Genetic variation within cottonwoods shapes communities and ecosystems, but these effects may be modified by phenotypic plasticity, where genotype traits change in response to environmental conditions. Here, we investigated plasticity in Fremont cottonwood (Populus fremontii) leaf litter traits as well as the consequences of plasticity for riparian ecosystems. We used three common gardens each planted with genotypes from six genetically divergent populations spanning a 12°C temperature gradient, and a decomposition experiment in a common stream environment. We found that leaf litter area, specific leaf area, and carbon to nitrogen ratio (C:N) were determined by interactions between genetics and growing environment, as was the subsequent rate of litter decomposition. Most of the genetic variation in leaf litter traits appeared among rather than within source populations with distinct climate histories. Source populations from hotter climates generally produced litter that decomposed more quickly, but plasticity varied the magnitude of this effect. We also found that hotter growing conditions reduced the variation in litter traits produced across genotypes, homogenizing the litter inputs to riparian ecosystems. All genotypes in the hottest garden produced comparatively small leaves that decomposed quickly and supported lower abundances of aquatic invertebrates, whereas the same genotypes in the coldest garden produced litter with distinct morphologies and decomposition rates. Our results suggest that plastic responses to climate stress may constrict the expression of genetic variation in predictable ways that impact communities and ecosystems. Understanding these interactions between genetic and environmental variation is critical to our ability to plan for the role of foundation species when managing and restoring riparian ecosystems in a warming world., (© 2021 by the Ecological Society of America.)

Premise: Plants rely on pools of internal nonstructural carbohydrates (NSCs: soluble sugars plus starch) to support metabolism, growth, and regrowth of tissues damaged from disturbance such as foliage herbivory. However, impacts of foliage herbivory on the quantity and composition of NSC pools in long-lived woody plants are currently unclear. We implemented a controlled defoliation experiment on mature Tamarix spp.-a dominant riparian woody shrub/tree that has evolved with intense herbivory pressure-to test two interrelated hypotheses: (1) Repeated defoliation disproportionately impacts aboveground versus belowground NSC storage. (2) Defoliation disproportionately impacts starch versus soluble sugar storage., Methods: Hypotheses were tested by transplanting six Tamarix seedlings into each of eight cylinder mesocosms (2 m diameter, 1 m in depth). After 2.5 years, plants in four of the eight mesocosms were mechanically defoliated repeatedly over a single growing season, and all plants were harvested in the following spring., Results: Defoliation had no impact on either above- or belowground soluble sugar pools. However, starch in defoliated plants dropped to 55% and 26% in stems and roots, respectively, relative to control plants, resulting in an over 2-fold higher soluble sugar to starch ratio in defoliated plants., Conclusions: The results suggest that defoliation occurring over a single growing season does not impact immediate plant functions such as osmoregulation, but depleted starch could limit future fitness, particularly where defoliation occurs over multiple years. These results improve our understanding of how woody plants cope with episodic defoliation caused by foliage herbivory and other disturbances., (© 2021 Botanical Society of America.)

Environmental change and biodiversity loss are but two of the complex challenges facing conservation practitioners and policy makers. Relevant and robust scientific knowledge is critical for providing decision-makers with the actionable evidence needed to inform conservation decisions. In the Anthropocene, science that leads to meaningful improvements in biodiversity conservation, restoration and management is desperately needed. Conservation Physiology has emerged as a discipline that is well-positioned to identify the mechanisms underpinning population declines, predict responses to environmental change and test different in situ and ex situ conservation interventions for diverse taxa and ecosystems. Here we present a consensus list of 10 priority research themes. Within each theme we identify specific research questions (100 in total), answers to which will address conservation problems and should improve the management of biological resources. The themes frame a set of research questions related to the following: (i) adaptation and phenotypic plasticity; (ii) human-induced environmental change; (iii) human-wildlife interactions; (iv) invasive species; (v) methods, biomarkers and monitoring; (vi) policy, engagement and communication; (vii) pollution; (viii) restoration actions; (ix) threatened species; and (x) urban systems. The themes and questions will hopefully guide and inspire researchers while also helping to demonstrate to practitioners and policy makers the many ways in which physiology can help to support their decisions., (© The Author(s) 2021. Published by Oxford University Press and the Society for Experimental Biology.)

The tall (>4 m), charismatic and threatened columnar cacti, pasacana [ Echinopsis atacamensis (Vaupel) Friedrich & G.D. Rowley)], grows on the Bolivian Altiplano and provides environmental and economic value to these extremely cold, arid and high-elevation (~4000 m) ecosystems. Yet very little is known about their growth rates, ages, demography and climate sensitivity. Using radiocarbon in spine dating time series, we quantitatively estimate the growth rate (5.8 and 8.3 cm yr -1 ) and age of these cacti (up to 430 years). These data and our field measurements yield a survivorship curve that suggests precipitation on the Altiplano is important for this species' recruitment. Our results also reveal a relationship between nighttime temperatures on the Altiplano and the variation in oxygen isotope values in spines (δ 18 O). The annual δ 18 O minimums from 58 years of in-series spine tissue from pasacana on the Altiplano provides at least decadal proxy records of temperature ( r  = 0.58; P  < 0.0001), and evidence suggests that there are longer records connecting modern Altiplano temperatures to sea-surface temperatures (SSTs) in the Atlantic Ocean. While the role of Atlantic SSTs on the South American Summer Monsoon (SASM) and precipitation on the Bolivian Altiplano is well described, the impact of SSTs on Altiplano temperatures is disputed. Understanding the modern impact of SSTs on temperature on the Altiplano is important to both understand the impact of future climate change on pasacana cactus and to understand past climate changes on the Altiplano. This is the best quantitative evidence to date of one of the oldest known cactus in the world, although there are likely many older cacti on the Altiplano, or elsewhere, that have not been sampled yet. Together with growth, isotope and age data, this information should lead to better management and conservation outcomes for this threatened species and the Altiplano ecosystem., (© The Author(s) 2021. Published by Oxford University Press and the Society for Experimental Biology.)

The COVID-19 pandemic and associated public health measures have had unanticipated effects on ecosystems and biodiversity. Conservation physiology and its mechanistic underpinnings are well positioned to generate robust data to inform the extent to which the Anthropause has benefited biodiversity through alterations in disturbance-, pollution- and climate change-related emissions. The conservation physiology toolbox includes sensitive biomarkers and tools that can be used both retroactively (e.g. to reconstruct stress in wildlife before, during and after lockdown measures) and proactively (e.g. future viral waves) to understand the physiological consequences of the pandemic. The pandemic has also created new risks to ecosystems and biodiversity through extensive use of various antimicrobial products (e.g. hand cleansers, sprays) and plastic medical waste. Conservation physiology can be used to identify regulatory thresholds for those products. Moreover, given that COVID-19 is zoonotic, there is also opportunity for conservation physiologists to work closely with experts in conservation medicine and human health on strategies that will reduce the likelihood of future pandemics (e.g. what conditions enable disease development and pathogen transfer) while embracing the One Health concept. The conservation physiology community has also been impacted directly by COVID-19 with interruptions in research, training and networking (e.g. conferences). Because this is a nascent discipline, it will be particularly important to support early career researchers and ensure that there are recruitment pathways for the next generation of conservation physiologists while creating a diverse and inclusive community. We remain hopeful for the future and in particular the ability of the conservation physiology community to deliver relevant, solutions-oriented science to guide decision makers particularly during the important post-COVID transition and economic recovery., (© The Author(s) 2021. Published by Oxford University Press and the Society for Experimental Biology.)

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., (© The Author(s) 2020. Published by Oxford University Press and the Society for Experimental Biology.)

Non-structural carbohydrates (NSCs) are necessary for plant growth and affected by plant water status, but the temporal dynamics of water stress impacts on NSC are not well understood. We evaluated how seasonal NSC concentrations varied with plant water status (predawn xylem water potential, Ψ) and air temperature (T) in the evergreen desert shrub Larrea tridentata. Aboveground sugar and starch concentrations were measured weekly or monthly for ~1.5 years on 6-12 shrubs simultaneously instrumented with automated stem psychrometers; leaf photosynthesis (A net ) was measured monthly for 1 year. Leaf sugar increased during the dry, premonsoon period, associated with lower Ψ (greater water stress) and high T. Leaf sugar accumulation coincided with declines in leaf starch and stem sugar, suggesting the prioritization of leaf sugar during low photosynthetic uptake. Leaf starch was strongly correlated with A net and peaked during the spring and monsoon seasons, while stem starch remained relatively constant except for depletion during the monsoon. Recent photosynthate appeared sufficient to support spring growth, while monsoon growth required the remobilization of stem starch reserves. The coordinated responses of different NSC fractions to water status, photosynthesis, and growth demands suggest that NSCs serve multiple functions under extreme environmental conditions, including severe drought., (© 2020 John Wiley & Sons Ltd.)

Applying physiological tools, knowledge and concepts to understand conservation problems (i.e. conservation physiology) has become commonplace and confers an ability to understand mechanistic processes, develop predictive models and identify cause-and-effect relationships. Conservation physiology is making contributions to conservation solutions; the number of 'success stories' is growing, but there remain unexplored opportunities for which conservation physiology shows immense promise and has the potential to contribute to major advances in protecting and restoring biodiversity. Here, we consider how conservation physiology has evolved with a focus on reframing the discipline to be more inclusive and integrative. Using a 'horizon scan', we further explore ways in which conservation physiology can be more relevant to pressing conservation issues of today (e.g. addressing the Sustainable Development Goals; delivering science to support the UN Decade on Ecosystem Restoration), as well as more forward-looking to inform emerging issues and policies for tomorrow. Our horizon scan provides evidence that, as the discipline of conservation physiology continues to mature, it provides a wealth of opportunities to promote integration, inclusivity and forward-thinking goals that contribute to achieving conservation gains. To advance environmental management and ecosystem restoration, we need to ensure that the underlying science (such as that generated by conservation physiology) is relevant with accompanying messaging that is straightforward and accessible to end users., (© The Author(s) 2020. Published by Oxford University Press and the Society for Experimental Biology.)

Plant species are characterized along a spectrum of isohydry to anisohydry depending on their regulation of water potential (Ψ), but the plasticity of hydraulic strategies is largely unknown. The role of environmental drivers was evaluated in the hydraulic behavior of Larrea tridentata, a drought-tolerant desert shrub that withstands a wide range of environmental conditions. With a 1.5 yr time-series of 2324 in situ measurements of daily predawn and midday Ψ, the temporal variability of hydraulic behavior was explored in relation to soil water supply, atmospheric demand and temperature. Hydraulic behavior in Larrea was highly dynamic, ranging from partial isohydry to extreme anisohydry. Larrea exhibited extreme anisohydry under wet soil conditions corresponding to periods of high productivity, whereas partial isohydry was exhibited after prolonged dry or cold conditions, when productivity was low. Environmental conditions can strongly influence plant hydraulic behavior at relatively fast timescales, which enhances our understanding of plant drought responses. Although species may exhibit a dominant hydraulic behavior, variable environmental conditions can prompt plasticity in Ψ regulation, particularly for species in seasonally dry climates., (© 2019 The Authors. New Phytologist © 2019 New Phytologist Trust.)

The plant family Cactaceae is considered among the most threatened groups of organisms on the planet. The threatened status of the cacti family has created a renewed interest in the highly evolved physiological and morphological traits that underpin their persistence in some of the harshest subtropical environments in the Americas. Among the most important anatomical features of cacti is the modification of leaves into spines, and previous work has shown that the stable isotope chemistry of cacti spines records potential variations in stem water balance, stress, and Crassulacean acid metabolism (CAM). We review the opportunities, challenges, and pitfalls in measuring δ 13C, δ 2H, and δ 18O ratios captured in spine tissues that potentially reflect temporal and spatial patterns of stomatal conductance, internal to atmospheric CO2 partial pressures, and subsequent patterns of photosynthetic gas exchange. We then evaluate the challenges in stable isotope analysis in spine tissues related to variation in CAM expression, stem water compartmentalization, and spine whole-tissue composition among other factors. Finally, we describe how the analysis of all three isotopes can be used in combination to provide potentially robust analysis of photosynthetic function in cacti, and other succulent-stemmed taxa across broad spatio-temporal environmental gradients., (© The Author(s) 2019. Published by Oxford University Press on behalf of the Society for Experimental Biology. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

Tree mortality is a key driver of forest dynamics and its occurrence is projected to increase in the future due to climate change. Despite recent advances in our understanding of the physiological mechanisms leading to death, we still lack robust indicators of mortality risk that could be applied at the individual tree scale. Here, we build on a previous contribution exploring the differences in growth level between trees that died and survived a given mortality event to assess whether changes in temporal autocorrelation, variance, and synchrony in time-series of annual radial growth data can be used as early warning signals of mortality risk. Taking advantage of a unique global ring-width database of 3065 dead trees and 4389 living trees growing together at 198 sites (belonging to 36 gymnosperm and angiosperm species), we analyzed temporal changes in autocorrelation, variance, and synchrony before tree death (diachronic analysis), and also compared these metrics between trees that died and trees that survived a given mortality event (synchronic analysis). Changes in autocorrelation were a poor indicator of mortality risk. However, we found a gradual increase in inter-annual growth variability and a decrease in growth synchrony in the last ∼20 years before mortality of gymnosperms, irrespective of the cause of mortality. These changes could be associated with drought-induced alterations in carbon economy and allocation patterns. In angiosperms, we did not find any consistent changes in any metric. Such lack of any signal might be explained by the relatively high capacity of angiosperms to recover after a stress-induced growth decline. Our analysis provides a robust method for estimating early-warning signals of tree mortality based on annual growth data. In addition to the frequently reported decrease in growth rates, an increase in inter-annual growth variability and a decrease in growth synchrony may be powerful predictors of gymnosperm mortality risk, but not necessarily so for angiosperms.

Premise of the Study: Giant cacti species possess long cylindrical stems that store massive amounts of water and other resources to draw on for photosynthesis, growth, and reproduction during hot and dry conditions. Across all giant cacti taxa, stem photosynthetic surface area to volume ratio (S:V) varies by several fold. This broad morphological diversity leads to the hypothesis that giant cacti function along a predictable resource use continuum from a "safe" strategy reflected in low S:V, low relative growth rates (RGR), and low net assimilation rates (A net ) to a high-risk strategy that is reflected in high S:V, RGR, and A net ., Methods: To test this hypothesis, whole-plant gas exchange, chlorophyll fluorescence, and whole-spine-tissue carbon isotope ratios (δ 13 C) were measured in four giant cacti species varying in stem morphology and RGR. Measurements were conducted on five well-watered, potted plants per species., Key Results: Under conditions of mild diel temperatures and low atmospheric vapor pressure deficit, A net , transpiration (E), and stomatal conductance (G s ) were significantly higher, and water-use efficiency (A net : G s ) was lower in fast-growing, multi-stemmed species compared to the slower growing, single-stemmed species. However, under warmer, less optimal conditions, gas exchange converged between stem types, and neither δ 13 C nor chlorophyll fluorescence varied among species., Conclusions: The results add to a growing body of evidence that succulent-stemmed plants function along a similar economic spectrum as leaf-bearing plants such that functional traits including stem RGR, longevity, morphology, and gas exchange are correlated across species with varying life-history strategies., (© 2018 Botanical Society of America.)

Environmental changes have resulted in significant declines in native riparian forests that are comprised largely of dioecious tree taxa, including boxelder and iconic cottonwood/willow gallery forests. Dioecious species may be especially vulnerable to the effects of climate change given that they often exhibit skewed sex ratios that are reinforced by physiological and morphological specialization of each sex to specific microhabitats. A comprehensive data synthesis suggests that male individuals of boxelder and cottonwood taxa have a higher representation on dry microhabitats than females and are less physiologically sensitive to increased aridity than co-occurring females. Consequently, extreme male-biased sex ratios are possible under future climate conditions that could reduce population fitness below a sustainable threshold. Riparian willows, on the other hand, generally do not express obvious sexual dimorphism in habitat preference or physiological sensitivity to aridity. Thus, it is unclear whether climate change will impact population structure of willows in ways that parallel other dioecious riparian tree taxa. Future riparian tree restoration programs should aim to maintain future sex ratio balance that maximizes population fitness under projected hydro-climatological conditions. Recent advances in genomics will likely provide the critical tools for early sex determination in pre-reproductive trees across riparian tree species such that sex ratio balance could be targeted during initial stages of restoration, along with adaptations for drought tolerance and other key traits that are essential for survival under future conditions.

For over a century, physiological tools and techniques have been allowing researchers to characterize how organisms respond to changes in their natural environment and how they interact with human activities or infrastructure. Over time, many of these techniques have become part of the conservation physiology toolbox, which is used to monitor, predict, conserve, and restore plant and animal populations under threat. Here, we provide a summary of the tools that currently comprise the conservation physiology toolbox. By assessing patterns in articles that have been published in 'Conservation Physiology' over the past 5 years that focus on introducing, refining and validating tools, we provide an overview of where researchers are placing emphasis in terms of taxa and physiological sub-disciplines. Although there is certainly diversity across the toolbox, metrics of stress physiology (particularly glucocorticoids) and studies focusing on mammals have garnered the greatest attention, with both comprising the majority of publications (>45%). We also summarize the types of validations that are actively being completed, including those related to logistics (sample collection, storage and processing), interpretation of variation in physiological traits and relevance for conservation science. Finally, we provide recommendations for future tool refinement, with suggestions for: (i) improving our understanding of the applicability of glucocorticoid physiology; (ii) linking multiple physiological and non-physiological tools; (iii) establishing a framework for plant conservation physiology; (iv) assessing links between environmental disturbance, physiology and fitness; (v) appreciating opportunities for validations in under-represented taxa; and (vi) emphasizing tool validation as a core component of research programmes. Overall, we are confident that conservation physiology will continue to increase its applicability to more taxa, develop more non-invasive techniques, delineate where limitations exist, and identify the contexts necessary for interpretation in captivity and the wild.

Tree mortality is a key factor influencing forest functions and dynamics, but our understanding of the mechanisms leading to mortality and the associated changes in tree growth rates are still limited. We compiled a new pan-continental tree-ring width database from sites where both dead and living trees were sampled (2970 dead and 4224 living trees from 190 sites, including 36 species), and compared early and recent growth rates between trees that died and those that survived a given mortality event. We observed a decrease in radial growth before death in ca. 84% of the mortality events. The extent and duration of these reductions were highly variable (1-100 years in 96% of events) due to the complex interactions among study species and the source(s) of mortality. Strong and long-lasting declines were found for gymnosperms, shade- and drought-tolerant species, and trees that died from competition. Angiosperms and trees that died due to biotic attacks (especially bark-beetles) typically showed relatively small and short-term growth reductions. Our analysis did not highlight any universal trade-off between early growth and tree longevity within a species, although this result may also reflect high variability in sampling design among sites. The intersite and interspecific variability in growth patterns before mortality provides valuable information on the nature of the mortality process, which is consistent with our understanding of the physiological mechanisms leading to mortality. Abrupt changes in growth immediately before death can be associated with generalized hydraulic failure and/or bark-beetle attack, while long-term decrease in growth may be associated with a gradual decline in hydraulic performance coupled with depletion in carbon reserves. Our results imply that growth-based mortality algorithms may be a powerful tool for predicting gymnosperm mortality induced by chronic stress, but not necessarily so for angiosperms and in case of intense drought or bark-beetle outbreaks., (© 2016 John Wiley & Sons Ltd.)

Patterns of woody-plant mortality have been linked to global-scale environmental changes, such as extreme drought, heat stress, more frequent and intense fires, and episodic outbreaks of insects and pathogens. Although many studies have focussed on survival and mortality in response to specific physiological stresses, little attention has been paid to the role of genetic heritability of traits and local adaptation in influencing patterns of plant mortality, especially in non-native species. Tamarix spp. is a dominant, non-native riparian tree in western North America that is experiencing dieback in some areas of its range due to episodic herbivory by the recently introduced northern tamarisk leaf beetle ( Diorhabda carinulata ). We propose that genotype × environment interactions largely underpin current and future patterns of Tamarix mortality. We anticipate that (i) despite its recent introduction, and the potential for significant gene flow, Tamarix in western North America is generally adapted to local environmental conditions across its current range in part due to hybridization of two species; (ii) local adaptation to specific climate, soil and resource availability will yield predictable responses to episodic herbivory; and (iii) the ability to cope with a combination of episodic herbivory and increased aridity associated with climate change will be largely based on functional tradeoffs in resource allocation. This review focusses on the potential heritability of plant carbon allocation patterns in Tamarix , focussing on the relative contribution of acquired carbon to non-structural carbohydrate (NSC) pools versus other sinks as the basis for surviving episodic disturbance. Where high aridity and/or poor edaphic position lead to chronic stress, NSC pools may fall below a minimum threshold because of an imbalance between the supply of carbon and its demand by various sinks. Identifying patterns of local adaptation of traits related to resource allocation will improve forecasting of Tamarix population susceptibility to episodic herbivory.

It has been proposed that we are now living in a new geological epoch known as the Anthropocene, which is specifically defined by the impacts that humans are having on the Earth's biological diversity and geology. Although the proposal of this term was borne out of an acknowledgement of the negative changes we are imparting on the globe (e.g. climate change, pollution, coastal erosion, species extinctions), there has recently been action amongst a variety of disciplines aimed at achieving a 'good Anthropocene' that strives to balance societal needs and the preservation of the natural world. Here, we outline ways that the discipline of conservation physiology can help to delineate a hopeful, progressive and productive path for conservation in the Anthropocene and, specifically, achieve that vision. We focus on four primary ways that conservation physiology can contribute, as follows: (i) building a proactive approach to conservation; (ii) encouraging a pragmatic perspective; (iii) establishing an appreciation for environmental resilience; and (iv) informing and engaging the public and political arenas. As a collection of passionate individuals combining theory, technological advances, public engagement and a dedication to achieving conservation success, conservation physiologists are poised to make meaningful contributions to the productive, motivational and positive way forward that is necessary to curb and reverse negative human impact on the environment.

Volume-to-surface area ratio (V:S) across stem succulent taxa varies by almost two orders of magnitude. The broad range in V:S of cacti and other succulent species likely has considerable importance for adaptation since stem volume determines the storage capacity of water, carbon and nutrients and stem surface area is directly related to whole-stem photosynthetic capacity. We examined the intrinsic physiological tradeoffs across diverse stem morphologies in three divergent evolutionary groups where stem succulence is common: Cactoideae, Opuntioideae (Cactaceae) and Euphorbiaceae. We predicted that variation in physiological response to environmental conditions would be (1) constrained by stem V:S, and (2) detectable in the stable isotope ratios of plant tissues. Stable isotope ratios were measured in the spines/prickles of 62 stem-succulent species occurring in a common garden setting in Phoenix, AZ, USA. Biomass δ(13)C, δ(2)H and δ(18)O increased with V:S in Cactoideae only, possibly reflecting various levels of Crassulacean acid metabolism (CAM) strength in the other lineages. Within Cactoideae-group with the highest CAM strength and largest range in V:S-δ(13)C and δ(18)O increased 2.2 and 11.5 ‰, respectively, with a 22-fold increase in V:S. Both δ(13)C and V:S decreased with species climate-niche estimates of precipitation, indicating that stem morphology and physiology in Cactoideae may be constrained by available moisture. Taken together, these data suggest that physiological tradeoffs associated with stem V:S are detectable across broad evolutionary groups despite differences in CAM strength.

Climate change, particularly increased aridity, poses a significant threat to plants and the biotic communities they support. Dioecious species may be especially vulnerable to climate change given that they often exhibit spatial segregation of the sexes, reinforced by physiological and morphological specialization of each sex to different microhabitats. In dimorphic species, the overexpression of a trait by one gender versus the other may become suppressed in future climates. Data suggest that males will generally be less sensitive to increased aridity than co-occurring females and, consequently, extreme male-biased sex ratios are possible in a significant number of populations. The effects of male-biased sex ratios are likely to cascade to dependent community members, especially those that are specialized on one sex.

The potential benefits of physiology for conservation are well established and include greater specificity of management techniques, determination of cause-effect relationships, increased sensitivity of health and disturbance monitoring and greater capacity for predicting future change. While descriptions of the specific avenues in which conservation and physiology can be integrated are readily available and important to the continuing expansion of the discipline of 'conservation physiology', to date there has been no assessment of how the field has specifically contributed to conservation success. However, the goal of conservation physiology is to foster conservation solutions and it is therefore important to assess whether physiological approaches contribute to downstream conservation outcomes and management decisions. Here, we present eight areas of conservation concern, ranging from chemical contamination to invasive species to ecotourism, where physiological approaches have led to beneficial changes in human behaviour, management or policy. We also discuss the shared characteristics of these successes, identifying emerging themes in the discipline. Specifically, we conclude that conservation physiology: (i) goes beyond documenting change to provide solutions; (ii) offers a diversity of physiological metrics beyond glucocorticoids (stress hormones); (iii) includes approaches that are transferable among species, locations and times; (iv) simultaneously allows for human use and benefits to wildlife; and (v) is characterized by successes that can be difficult to find in the primary literature. Overall, we submit that the field of conservation physiology has a strong foundation of achievements characterized by a diversity of conservation issues, taxa, physiological traits, ecosystem types and spatial scales. We hope that these concrete successes will encourage the continued evolution and use of physiological tools within conservation-based research and management plans.

Desert riparian ecosystems of North America are hotspots of biodiversity that support many sensitive species, and are in a region experiencing some of the highest rates of climatic alteration in North America. Fremont cottonwood, Populus fremontii, is a foundation tree species of this critical habitat, but it is threatened by global warming and regional drying, and by a non-native tree/shrub, Tamarix spp., all of which can disrupt the mutualism between P. fremontii and its beneficial mycorrhizal fungal communities. Specialist herbivorous leaf beetles (Diorhabda spp.) introduced for biocontrol of Tamarix are altering the relationship between this shrub and its environment. Repeated episodic feeding on Tamarix foliage by Diorhabda results in varying rates of dieback and mortality, depending on genetic variation in allocation of resources, growing conditions, and phenological synchrony between herbivore and host plant. In this article, we review the complex interaction between climatic change and species introductions and their combined impacts on P. fremontii and their associated communities. We anticipate that (1) certain genotypes of P. fremontii will respond more favorably to the presence of Tamarix and to climatic change due to varying selection pressures to cope with competition and stress; (2) the ongoing evolution of Diorhabda's life cycle timing will continue to facilitate its expansion in North America, and will over time enhance herbivore impact to Tamarix; (3) defoliation by Diorhabda will reduce the negative impact of Tamarix on P. fremontii associations with mycorrhizal fungi; and (4) spatial variability in climate and climatic change will modify the capacity for Tamarix to survive episodic defoliation by Diorhabda, thereby altering the relationship between Tamarix and P. fremontii, and its associated mycorrhizal fungal communities. Given the complex biotic/abiotic interactions outlined in this review, conservation biologists and riparian ecosystem managers should strive to identify and conserve the phenotypic traits that underpin tolerance and resistance to stressors such as climate change and species invasion. Such efforts will greatly enhance conservation restoration efficacy for protecting P. fremontii forests and their associated communities., (© The Author 2015. Published by Oxford University Press on behalf of the Society for Integrative and Comparative Biology. All rights reserved. For permissions please email: journals.permissions@oup.com.)

Columnar cacti occur naturally in many habitats and environments in the Americas but are conspicuously dominant in very dry desert regions. These majestic plants are widely regarded for their cultural, economic, and ecological value and, in many ecosystems, support highly diverse communities of pollinators, seed dispersers, and frugivores. Massive amounts of water and other resources stored in the succulent photosynthetic stems of these species confer a remarkable ability to grow and reproduce during intensely hot and dry periods. Yet many columnar cacti are potentially under severe threat from environmental global changes, including climate change and loss of habitat. Stems in columnar cacti and other cylindrical-stemmed cacti are morphologically diverse; stem volume-to-surface area ratio (V:S) across these taxa varies by almost two orders of magnitude. Intrinsic functional trade-offs are examined here across a broad range of V:S in species of columnar cacti. It is proposed that variation in photosynthetic gas exchange, growth, and response to stress is highly constrained by stem V:S, establishing a mechanistic framework for understanding the sensitivity of columnar cacti to climate change and drought. Specifically, species that develop stems with low V:S, and thus have little storage capacity, are expected to express high mass specific photosynthesis and growth rates under favourable conditions compared with species with high V:S. But the trade-off of having little storage capacity is that low V:S species are likely to be less tolerant of intense or long-duration drought compared with high V:S species. The application of stable isotope measurements of cactus spines as recorders of growth, water relations, and metabolic responses to the environment across species of columnar cacti that vary in V:S is also reviewed. Taken together, our approach provides a coherent theory and required set of observations needed for predicting the responses of columnar cacti to climate change., (© The Author 2014. Published by Oxford University Press on behalf of the Society for Experimental Biology. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

Air temperatures in the arid western United States are predicted to increase over the next century. These increases will likely impact the distribution of plant species, particularly dioecious species that show a spatial segregation of the sexes across broad resource gradients. On the basis of spatial segregation patterns, we hypothesized that temperature increases will have a greater negative impact on female plants compared with co-occurring male plants of dioecious species. This hypothesis was tested by examining the whole-plant carbon and water relations of 10-year-old female (n = 18) and male (n = 13) Acer negundo Sarg. trees grown in a common garden in Salt Lake City, UT. The trees were established from cuttings collected where the growing season temperature averaged about 6.5 °C cooler than at the common garden. During May and June, stem sap flux (Js ) was similar between genders, but averaged 25% higher in males during the warmer months of July and August. Daytime canopy stomatal conductance (gs ) per unit leaf area was 12% higher in females in May : June, but was 11% higher in males in July : August. We combined measurements of sap flux-scaled transpiration with measurements of tree allometry and δ(13) C of leaf soluble sugars to estimate whole-tree carbon assimilation (Atree ) and water use efficiency (WUE) (Atree  : Etree ). Atree was similar between genders until late August when Atree was 32% higher in male trees. Atree  : Etree was on average 7% higher in females than in males during the growing season. Patterns of Js , gs , Atree and Atree  : Etree in the present study were in contrast to those previously reported for A. negundo genders under native growing season temperatures. Results suggest that the spatial segregation of the sexes could shift under global warming such that female plants lose their dominance in high-resource habitats, and males increase their dominance in relatively lower-resource habitats., (© 2013 John Wiley & Sons Ltd.)

Background and Aims: Under equal conditions, plants that allocate a larger proportion of resources to growth must do so at the expense of investing fewer resources to storage. The critical balance between growth and storage leads to the hypothesis that in high-resource environments, plants that express high growth rates are more susceptible to episodic disturbance than plants that express lower growth rates., Methods: This hypothesis was tested by measuring the radial growth, basal area increment (BAI) and carbon isotope ratios (δ(13)C) in tree-ring α-cellulose of 62 mature tamarisk trees (Tamarix spp.) occurring at three sites in the western USA (n = 31 live and 31 killed trees across all sites, respectively). All of the trees had been subjected to periods of complete foliage loss by episodic herbivory over three or more consecutive growing seasons by the tamarisk leaf beetle (Diorhabda carinulata), resulting in approx. 50 % mortality at each site., Key Results: Mean annual BAI (measured from annual ring widths) in the 10 years prior to the onset of herbivory was on average 45 % higher in killed trees compared with live trees (P < 0·0001). Killed trees that had higher growth rates also expressed higher (less negative) δ(13)C ratios compared with live trees. In fact, at one site near Moab, UT, the mean annual BAI was 100 % higher in killed trees despite having about a 0·5 ‰ higher δ(13)C relative to live trees (P = 0·0008). Patterns of δ(13)C suggest that the intrinsic water-use efficiency was higher in killed than surviving trees, possibly as a consequence of lower whole-canopy stomatal conductance relative to live trees., Conclusions: The results show that a likely trade-off occurs between radial growth and survival from foliage herbivory in Tamarix spp. that currently dominates riparian areas throughout the western USA and northern Mexico. Thus, herbivory by D. carinulata may reduce the overall net primary productivity of surviving Tamarix trees and may result in a reduction in genetic variability in this dominant invasive tree species if these allocation patterns are adaptive.

Thermal dissipation probes (the Granier method) are routinely used in forest ecology and water balance studies to estimate whole-tree transpiration. This method utilizes an empirically derived equation to measure sap flux density, which has been reported as independent of wood characteristics. However, errors in calculated sap flux density may occur when large gradients in sap velocity occur along the sensor length or when sensors are inserted into non-conducting wood. These may be conditions routinely associated with ring-porous species, yet there are few cases in which the original calibration has been validated for ring-porous species. We report results from laboratory calibration measurements conducted on excised stems of four ring-porous species and two diffuse-porous species. Our calibration results for ring-porous species were considerably different compared with the original calibration equation. Calibration equation coefficients obtained in this study differed by as much as two to almost three orders of magnitude when compared with the original equation of Granier. Coefficients also differed between ring-porous species across all pressure gradient conditions considered; however, no differences between calibration slopes were observed for data collected within the range of expected in situ pressure gradients. In addition, dye perfusions showed that in three of the four ring-porous species considered, active sapwood was limited to the outermost growth ring. In contrast, our calibration results for diffuse-porous species showed generally good agreement with the empirically derived Granier calibration, and dye perfusions showed that active sapwood was associated with many annual growth rings. Our results suggest that the original calibration of Granier is not universally applicable to all species and xylem types and that previous estimates of absolute rates of water use for ring-porous species obtained using the original calibration coefficients may be associated with substantial error.

Riparian cottonwood/willow forest assemblages are highly valued in the southwestern United States for their wildlife habitat, biodiversity, and watershed protection. Yet these forests are under considerable threat from climate change impacts on water resources and land-use activities to support human enterprise. Stream diversions, groundwater pumping, and extended drought have resulted in the decline of cottonwood/willow forests along many riparian corridors in the Southwest and, in many cases, the replacement of these forests with less desirable invasive shrubs and trees. Nevertheless, ecophysiological responses of cottonwood and willow, along with associated ecohydrological feedbacks of soil water depletion, are not well understood. Ecophysiological processes of mature Fremont cottonwood and coyote willow stands were examined over four consecutive growing seasons (2004-2007) near Salt Lake City, Utah, USA. The tree stands occurred near the inlet of a reservoir that was drained in the spring of 2005 and remained empty until mid-summer of 2006, effectively removing the primary water source for most of two growing seasons. Stem sap flux density (Js) in cottonwood was highly correlated with volumetric soil moisture (theta) in the upper 60 cm and decreased sevenfold as soil moisture dropped from 12% to 7% after the reservoir was drained. Conversely, Js in willow was marginally correlated with 0 and decreased by only 25% during the same period. Opposite patterns emerged during the following growing season: willow had a lower whole-plant conductance (kt) in June and higher leaf carbon isotope ratios (delta13C) than cottonwood in August, whereas k(t) and delta13C were otherwise similar between species. Water relations in both species recovered quickly from soil water depletion, with the exception that sapwood area to stem area (As:Ast) was significantly lower in both species after the 2007 growing season compared to 2004. Results suggest that cottonwood has a greater sensitivity to interannual reductions in water availability, while willow is more sensitive to longer periods of soil water depletion. These data shed light on the linkage between soil water deficits and ecophysiological processes of threatened riparian forests given potential land-use and long-term drought impacts on freshwater resources.

Acer negundo Sarg. (box elder) is a dioecious tree species that dominates riparian systems at mid elevations throughout the southwest and Intermountain West of the United States. Previous studies have shown that female A. negundo trees occur at higher frequencies along stream margins, whereas males occur at higher frequencies in drier microsites. To better understand the adaptive significance of sex ratio biases and their impact on the ecohydrology of riparian ecosystems, we examined whole-plant water relations and hydraulic properties of mature male and female A. negundo trees occurring within 1 m of a perennial stream channel. We hypothesized that (1) females would have significantly greater canopy water fluxes than males (particularly during periods of seed production: May-June), and (2) xylem in females is more hydraulically efficient but more vulnerable to cavitation than xylem in males. Mean sap flux density (J(s)) during the early growing season (May and June) was 43% higher in female trees than in male trees (n = 6 and 7 trees respectively, P < 0.0001). Mean J(s) in July and August remained 17% higher in females than in males (P = 0.0009). Mean canopy stomatal conductance per unit leaf area (g(s,leaf)) in May and June was on average 140% higher in females than in males (P < 0.0001). Mean g(s,leaf) in July and August remained 69% higher in female trees than in male trees (P < 0.0001). Canopy stomatal conductance scaled to basal area was 90 and 31% higher in females relative to males during May-June and July-August, respectively (P < 0.0001 during both periods). Conversely, there were no apparent differences in either branch hydraulic conductance or branch xylem cavitation vulnerability between genders. These results improve our capacity to describe the adaptive forces that shape the spatial distribution of male and female trees in dioecious species, and their consequences for ecohydrological processes in riparian ecosystems.

Anthropogenic climate change is likely to alter the patterns of moisture availability globally. The consequences of these changes on species distributions and ecosystem function are largely unknown, but possibly predictable based on key ecophysiological differences among currently coexisting species. In this study, we examined the environmental and biological controls on transpiration from a piñon-juniper (Pinus edulis-Juniperus osteosperma) woodland in southern Utah, USA. The potential for climate-change-associated shifts in moisture inputs could play a critical role in influencing the relative vulnerabilities of piñons and junipers to drought and affecting management decisions regarding the persistence of this dominant landscape type in the Intermountain West. We aimed to assess the sensitivity of this woodland to seasonal variations in moisture and to mechanistically explain the hydraulic strategies of P. edulis and J. osteosperma through the use of a hydraulic transport model. Transpiration from the woodland was highly sensitive to variations in seasonal moisture inputs. There were two distinct seasonal pulses of transpiration: a reliable spring pulse supplied by winter-derived precipitation, and a highly variable summer pulse supplied by monsoonal precipitation. Transpiration of P. edulis and J. osteosperma was well predicted by a mechanistic hydraulic transport model (R2 = 0.83 and 0.92, respectively). Our hydraulic model indicated that isohydric regulation of water potential in P. edulis minimized xylem cavitation during drought, which facilitated drought recovery (94% of pre-drought water uptake) but came at the cost of cessation of gas exchange for potentially extended periods. In contrast, the anisohydric J. osteosperma was able to maintain gas exchange at lower water potentials than P. edulis but experienced greater cavitation over the drought and showed a lesser degree of post-drought recovery (55% of pre-drought uptake). As a result, these species should be differentially affected by shifts in the frequency, duration, and intensity of drought. Our results highlight the sensitivity of this woodland type to potential climate-change-associated shifts in seasonal moisture patterns and demonstrate the utility of mechanistic hydraulic models in explaining differential responses of coexisting species to drought.

Plant transpiration is strongly constrained by hydraulic architecture, which determines the critical threshold for cavitation. Because species vary greatly in vulnerability to cavitation, hydraulic limits to transpiration and stomatal conductance have not generally been incorporated into ecological and climate models. We measured sap flow, leaf transpiration, and vulnerability to cavitation of a variety of tree species in a well-irrigated but semi-arid urban environment in order to evaluate the generality of stomatal responses to high atmospheric vapor pressure deficit (D). We found evidence of broad patterns of stomatal responses to humidity based on systematic differences in vulnerability to cavitation. Ring-porous taxa consistently had vulnerable xylem and showed strong regulation of transpiration in response to D, while diffuse-porous taxa were less vulnerable and transpiration increased nearly linearly with D. These results correspond well to patterns in the distribution of the taxa, such as the prevalence of diffuse-porous species in riparian ecosystems, and also provide a means of representing maximum transpiration rates at varying D in broad categories of trees.

Previous studies of pinyon-juniper woodlands show that Pinus edulis Engelm. makes better use of soil water from summer precipitation pulses than does co-occurring Juniperus osteosperma (Torr.) Little. To investigate the basis of this difference, we examined seasonal variation in cavitation and hydraulic conductance. Pinus edulis remained isohydric over the growing season. Minimum water potentials never fell below -2.3 MPa, and the extent of xylem cavitation remained near constant during the dry season. In contrast, J. osteosperma was anisohydric, reaching water potentials as low as -6.9 MPa, and experiencing progressively greater xylem cavitation as the dry season progressed despite having more cavitation-resistant xylem than P. edulis. We conducted an irrigation experiment to observe the responses of the study species to a summer pulse of water. Although sap flow increased in both species in response to the 25-mm irrigation pulse, only J. osteosperma responded to the 10-mm pulse. This was inconsistent with the response of P. edulis to light rain events and may have been due to a difference in the distribution of irrigation water and rain water between the under- and between-canopy areas. Whole-plant conductance increased following the 25-mm irrigation in P. edulis but remained constant in J. osteosperma. We hypothesized that this difference was caused, in part, by differential refilling of embolized xylem. Area specific hydraulic conductivity was 66% higher in roots of irrigated P. edulis trees relative to roots of control trees 3 days after the 25-mm irrigation (t = 2.14, P = 0.02, df = 16). There was no change in hydraulic conductivity of the roots of J. osteosperma or in the stems of either species. Our results indicate that the response to an irrigation pulse in P. edulis depended on cavitation avoidance in stems and the reversal of cavitation in roots, resulting in increased whole-plant conductance and water uptake. In contrast, J. osteosperma failed to exploit light summer rain events but was able to extract deep soil water at low water potentials.

The global water cycle is intimately linked to vegetation structure and function. Nowhere is this more apparent than in the arid west where riparian forests serve as ribbons of productivity in otherwise mostly unproductive landscapes. Dioecy is common among tree species that make up western North American riparian forests. There are intrinsic physiological differences between male and female dioecious riparian trees that may influence population structure (i.e., the ratio of male to female trees) and impact ecohydrology at large scales. In this paper, we review the current literature on sex ratio patterns and physiology of dioecious riparian tree species. Then develop a conceptual framework of the mechanisms that underlie population structure of dominant riparian tree species. Finally, we identify linkages between population structure and ecohydrological processes such as evapotranspiration and streamflow. A more thorough understanding of the mechanisms that underlie population structure of dominant riparian tree species will enable us to better predict global change impacts on vegetation and water cycling at multiple scales.

In water-limited environments of the intermountain region of North America, summer precipitation may play a role in the structure and function of aridland communities and ecosystems. This study examined the potential reliance on summer precipitation of two widespread, coexisting woody species in the southwestern United States, Pinus edulis Englmn. (Colorado piñon) and Juniperus osteosperma (Torr) Little (Utah juniper). The current distributions of P. edulis and J. osteosperma are highly suggestive of different dependencies on summer rainfall. We hypothesized that P. edulis was dependent on summer precipitation, utilizing summer precipitation even during extremely dry summers, whereas J. osteosperma was not dependent, using summer precipitation only when amounts were above some minimum threshold. Using sap flux and stable isotopic methods to assess seasonal water sources and water use efficiency, we examined the response of these two species to seasonal variations in moisture at a site located near the northern limits of the North American monsoon. Both sap flux and isotopic results indicated that P. edulis was responsive to summer rain, while J. osteosperma was not. Following summer rain events, sap flux density increased in P. edulis for several days, but not in J. osteosperma. Isotopic evidence indicated that P. edulis took up summer-derived moisture to a greater extent than J. osteosperma. Values of the natural abundance stable isotope ratio of carbon of leaf soluble carbohydrates increased over the summer for P. edulis, indicative of assimilation at higher water use efficiency, but were invariant for J. osteosperma. Our results supported the hypothesis that P. edulis and J. osteosperma are differentially sensitive to summer precipitation and are discussed in the light of potential changes in the seasonality of precipitation associated with climate change.