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The long-standing assumption that aboveground plant litter inputs have a substantial influence on soil organic carbon storage (SOC) and dynamics has been challenged by a new paradigm for SOC formation and persistence. We tested the importance of plant litter chemistry on SOC storage, distribution, composition, and age by comparing two highly contrasting ecosystems: an old-growth coast redwood (Sequoia sempervirens) forest, with highly aromatic litter, and an adjacent coastal prairie, with more easily decomposed litter. We hypothesized that if plant litter chemistry was the primary driver, redwood would store more and older SOC that was less microbially processed than prairie. Total soil carbon stocks to 110 cm depth were higher in prairie (35 kg C m-2) than redwood (28 kg C m-2). Radiocarbon values indicated shorter SOC residence times in redwood than prairie throughout the profile. Higher amounts of pyrogenic carbon and a higher degree of microbial processing of SOC appear to be instrumental for soil carbon storage and persistence in prairie, while differences in fine-root carbon inputs likely contribute to younger SOC in redwood. We conclude that at these sites fire residues, root inputs, and soil properties influence soil carbon dynamics to a greater degree than the properties of aboveground litter.

Understanding climate change impacts on the Cape Floristic Region requires improved knowledge of plant physiological responses to the environment. Studies examining physiological responses of mountain fynbos have consisted of campaign-based measurements, capturing snapshots of plant water relations and photosynthesis. We examine conclusions drawn from prior studies by tracking in situ physiological responses of three species, representing dominant growth forms (proteoid, ericoid, restioid), over 2 years using miniature continuous sap flow technology, long-term observations of leaf/culm water potential and gas exchange, and xylem vulnerability to embolism. We observed considerable inter-specific variation in the timing and extent of seasonal declines in productivity. Shallow-rooted Erica monsoniana exhibited steep within-season declines in sap flow and water potentials, and pronounced inter-annual variability in total daily sap flux (Js). Protea repens showed steady reductions in Js across both years, despite deeper roots and less negative water potentials. Cannomois congesta-a shallow-rooted restioid-was least negatively impacted. Following rehydrating rain at the end of summer, gas exchange recovery was lower in the drier year compared with the normal year, but did not differ between species. Loss of function in the drier year was partially accounted for by loss of xylem transport capacity in Erica and Cannomois, but not Protea. Hitherto unseen water use patterns, including inter-annual variability of gas exchange associated with contrasting water uptake properties, reveal that species use different mechanisms to cope with summer dry periods. Revealing physiological responses of key growth forms enhances predictions of plant function within mountain fynbos under future conditions.

Plants are critical mediators of terrestrial mass and energy fluxes, and their structural and functional traits have profound impacts on local and global climate, biogeochemistry, biodiversity, and hydrology. Yet, Earth System Models (ESMs), our most powerful tools for predicting the effects of humans on the coupled biosphere-atmosphere system, simplify the incredible diversity of land plants into a handful of coarse categories of "Plant Functional Types" (PFTs) that often fail to capture ecological dynamics such as biome distributions. The inclusion of more realistic functional diversity is a recognized goal for ESMs, yet there is currently no consistent, widely accepted way to add diversity to models, that is, to determine what new PFTs to add and with what data to constrain their parameters. We review approaches to representing plant diversity in ESMs and draw on recent ecological and evolutionary findings to present an evolution-based functional type approach for further disaggregating functional diversity. Specifically, the prevalence of niche conservatism, or the tendency of closely related taxa to retain similar ecological and functional attributes through evolutionary time, reveals that evolutionary relatedness is a powerful framework for summarizing functional similarities and differences among plant types. We advocate that Plant Functional Types based on dominant evolutionary lineages ("Lineage Functional Types") will provide an ecologically defensible, tractable, and scalable framework for representing plant diversity in next-generation ESMs, with the potential to improve parameterization, process representation, and model benchmarking. We highlight how the importance of evolutionary history for plant function can unify the work of disparate fields to improve predictive modeling of the Earth system.

Shaping global water and carbon cycles, plants lift water from roots to leaves through xylem conduits. The importance of xylem water conduction makes it crucial to understand how natural selection deploys conduit diameters within and across plants. Wider conduits transport more water but are likely more vulnerable to conduction-blocking gas embolisms and cost more for a plant to build, a tension necessarily shaping xylem conduit diameters along plant stems. We build on this expectation to present the Widened Pipe Model (WPM) of plant hydraulic evolution, testing it against a global dataset. The WPM predicts that xylem conduits should be narrowest at the stem tips, widening quickly before plateauing toward the stem base. This universal profile emerges from Pareto modeling of a trade-off between just two competing vectors of natural selection: one favoring rapid widening of conduits tip to base, minimizing hydraulic resistance, and another favoring slow widening of conduits, minimizing carbon cost and embolism risk. Our data spanning terrestrial plant orders, life forms, habitats, and sizes conform closely to WPM predictions. The WPM highlights carbon economy as a powerful vector of natural selection shaping plant function. It further implies that factors that cause resistance in plant conductive systems, such as conduit pit membrane resistance, should scale in exact harmony with tip-to-base conduit widening. Furthermore, the WPM implies that alterations in the environments of individual plants should lead to changes in plant height, for example, shedding terminal branches and resprouting at lower height under drier climates, thus achieving narrower and potentially more embolism-resistant conduits.

Cross-slope climate differences in the midlatitudes are ecologically important, and impact vegetation-mediated water balance between the earth surface and the atmosphere. We made high-resolution in situ observations of air temperature, relative humidity, soil moisture, insolation, and sap velocity observations on 14 Pacific madrone trees (Arbutus menziesii) spanning adjacent north and south slopes at the University of California's Angelo Coast Range Reserve. To understand the cross-slope response of sap velocity, a proxy for transpiration, to microclimate, we modeled the sap velocity on each slope using a transpiration model driven by ambient environment and parameterized with a Markov Chain Monte Carlo parameter estimation process. The results show that trees on opposing slopes do not follow a shared pattern of physiological response to transpiration drivers. This means that the observed sap velocity differences are not due entirely to observed microclimate differences, but also due to population-level physiological differences, which indicates acclimation to inhabited microclimate. While our present data set and analytical tools do not identify mechanisms of acclimation, we speculate that differing proportions of sun-adapted and shade-adapted leaves, differences in stomatal regulation, and cross-slope root zone moisture differences could explain some of the observed and modeled differences.

Modern human behavioral innovations from the Middle Stone Age (MSA) include the earliest indicators of full coastal adaptation evidenced by shell middens, yet many MSA middens remain poorly dated. We apply 230Th/U burial dating to ostrich eggshells (OES) from Ysterfontein 1 (YFT1, Western Cape, South Africa), a stratified MSA shell midden. 230Th/U burial ages of YFT1 OES are relatively precise (median ± 2.7%), consistent with other age constraints, and preserve stratigraphic principles. Bayesian age-depth modeling indicates YFT1 was deposited between 119.9 to 113.1 thousand years ago (ka) (95% CI of model ages), and the entire 3.8 m thick midden may have accumulated within ∼2,300 y. Stable carbon, nitrogen, and oxygen isotopes of OES indicate that during occupation the local environment was dominated by C3 vegetation and was initially significantly wetter than at present but became drier and cooler with time. Integrating archaeological evidence with OES 230Th/U ages and stable isotopes shows the following: 1) YFT1 is the oldest shell midden known, providing minimum constraints on full coastal adaptation by ∼120 ka; 2) despite rapid sea-level drop and other climatic changes during occupation, relative shellfish proportions and sizes remain similar, suggesting adaptive foraging along a changing coastline; 3) the YFT1 lithic technocomplex is similar to other west coast assemblages but distinct from potentially synchronous industries along the southern African coast, suggesting human populations were fragmented between seasonal rainfall zones; and 4) accumulation rates (up to 1.8 m/ka) are much higher than previously observed for dated, stratified MSA middens, implying more intense site occupation akin to Later Stone Age middens.

Quantitative knowledge of xylem physical tolerance limits to dehydration is essential to understanding plant drought tolerance but is lacking in many long-vessel angiosperms. We examine the hypothesis that a fundamental association between sustained xylem water transport and downstream tissue function should select for xylem that avoids embolism in long-vessel trees by quantifying xylem capacity to withstand air entry of western North American oaks (Quercus spp.). Optical visualization showed that 50% of embolism occurs at water potentials below -2.7 MPa in all 19 species, and -6.6 MPa in the most resistant species. By mapping the evolution of xylem vulnerability to embolism onto a fossil-dated phylogeny of the western North American oaks, we found large differences between clades (sections) while closely related species within each clade vary little in their capacity to withstand air entry. Phylogenetic conservatism in xylem physical tolerance, together with a significant correlation between species distributions along rainfall gradients and their dehydration tolerance, suggests that closely related species occupy similar climatic niches and that species' geographic ranges may have shifted along aridity gradients in accordance with their physical tolerance. Such trends, coupled with evolutionary associations between capacity to withstand xylem embolism and other hydraulic-related traits, yield wide margins of safety against embolism in oaks from diverse habitats. Evolved responses of the vascular system to aridity support the embolism avoidance hypothesis and reveal the importance of quantifying plant capacity to withstand xylem embolism for understanding function and biogeography of some of the Northern Hemisphere's most ecologically and economically important plants.

In nature-based treatment systems, such as constructed wetlands, plant uptake of nutrients can be a significant removal pathway. Current methods for quantifying plant uptake of nitrogen in constructed wetlands, which often involve harvesting biomass and assuming that all nitrogen stored in plants was derived from wastewater, are inappropriate in pilot- and full-scale systems where other sources of nitrogen are available. To improve our understanding of nitrogen cycling in constructed wetlands, we developed a new method to quantify plant uptake of nitrogen by using stable isotopes and a mixing model to distinguish between nitrogen sources. We applied this new method to a pilot-scale horizontal levee system (i.e., a subsurface constructed wetland) over a two-year monitoring period, during which 14% of nitrogen in plants was wastewater-derived on average and the remaining plant nitrogen was obtained from the soil. Analysis of nitrogen isotopes indicated substantial spatial variability in the wetland: 82% of nitrogen in plants within the first 2 m of the slope came from wastewater while less than 12% of plant nitrogen in the remainder of the wetland originated from wastewater. By combining these source contributions with remote-sensing derived total biomass measurements, we calculated that 150 kg N (95% CI = 50 kg N, 330 kg N) was taken up and retained by plants during the two-year monitoring period, which corresponded to approximately 8% of nitrogen removed in the wetland. Nitrogen uptake followed seasonal trends, increased as plants matured, and varied based on design parameters (e.g., plant types), suggesting that design decisions can impact this removal pathway. This new method can help inform efforts to understand nitrogen cycling and optimize the design of nature-based nutrient control systems.

Amazonian droughts are increasing in frequency and severity. However, little is known about how this may influence species-specific vulnerability to drought across different ecosystem types. We measured 16 functional traits for 16 congeneric species from six families and eight genera restricted to floodplain, swamp, white-sand or plateau forests of Central Amazonia. We investigated whether habitat distributions can be explained by species hydraulic strategies, and if habitat specialists differ in their vulnerability to embolism that would make water transport difficult during drought periods. We found strong functional differences among species. Nonflooded species had higher wood specific gravity and lower stomatal density, whereas flooded species had wider vessels, and higher leaf and xylem hydraulic conductivity. The P50 values (water potential at 50% loss of hydraulic conductivity) of nonflooded species were significantly more negative than flooded species. However, we found no differences in hydraulic safety margin among species, suggesting that all trees may be equally likely to experience hydraulic failure during severe droughts. Water availability imposes a strong selection leading to differentiation of plant hydraulic strategies among species and may underlie patterns of adaptive radiation in many tropical tree genera. Our results have important implications for modeling species distribution and resilience under future climate scenarios.

Relatively mesic environments within arid regions may be important conservation targets as 'climate change refugia' for species persistence in the face of worsening drought conditions. Semi-arid southern California and the relatively mesic environments of California's Channel Islands provide a model system for examining drought responses of plants in potential climate change refugia. Most methods for detecting refugia are focused on 'exposure' of organisms to certain abiotic conditions, which fail to assess how local adaptation or acclimation of plant traits (i.e. 'sensitivity') contribute to or offset the benefits of reduced exposure. Here, we use a comparative plant hydraulics approach to characterize the vulnerability of plants to drought, providing a framework for identifying the locations and trait patterns that underlie functioning climate change refugia. Seasonal water relations, xylem hydraulic traits and remotely sensed vegetation indices of matched island and mainland field sites were used to compare the response of native plants from contrasting island and mainland sites to hotter droughts in the early 21st century. Island plants experienced more favorable water relations and resilience to recent drought. However, island plants displayed low plasticity/adaptation of hydraulic traits to local conditions, which indicates that relatively conserved traits of island plants underlie greater hydraulic safety and localized buffering from regional drought conditions. Our results provide an explanation for how California's Channel Islands function as a regional climate refugia during past and current climate change and demonstrate a physiology-based approach for detecting potential climate change refugia in other systems.

The large-scale patterns of movement for the Sharp-shinned Hawk (Accipiter striatus), a small forest hawk found throughout western North America, are largely unknown. However, based on field observations we set out to test the hypothesis that juvenile migratory A. striatus caught along two distinct migration routes on opposite sides of the Sierra Nevada Mountains of North America (Pacific Coast and Intermountain Migratory Flyways) come from geographically different natal populations. We applied stable isotope analysis of hydrogen (H) and oxygen (O) of feathers, and large scale models of spatial isotopic variation (isoscapes) to formulate spatially explicit predictions of the origin of the migrant birds. Novel relationships were assessed between the measured hydrogen and oxygen isotope values of feathers from A. striatus museum specimens of known origin and the isoscape modeled hydrogen and oxygen isotope values of precipitation at those known locations. We used these relationships to predict the origin regions for birds migrating along the two flyways from the measured isotope values of migrant's feathers and the associated hydrogen and oxygen isotopic composition of precipitation where these feathers were formed. The birds from the two migration routes had overlap in their natal/breeding origins and did not differentiate into fully separate migratory populations, with birds from the Pacific Coast Migratory Flyway showing broader natal geographic origins than those from the Intermountain Flyway. The methodology based on oxygen isotopes had, in general, less predictive power than the one based on hydrogen. There was broad agreement between the two isotope approaches in the geographic assignment of the origins of birds migrating along the Pacific Coast Flyway, but not for those migrating along the Intermountain Migratory Flyway. These results are discussed in terms of their implications for conservation efforts of A. striatus in western North America, and the use of combined hydrogen and oxygen stable isotope analysis to track the movement of birds of prey on continental scales.

Vulnerability to embolism varies between con-generic species distributed along aridity gradients, yet little is known about intraspecific variation and its drivers. Even less is known about intraspecific variation in tissues other than stems, despite results suggesting that roots, stems and leaves can differ in vulnerability. We hypothesized that intraspecific variation in vulnerability in leaves and stems is adaptive and driven by aridity. We quantified leaf and stem vulnerability of Quercus douglasii using the optical technique. To assess contributions of genetic variation and phenotypic plasticity to within-species variation, we quantified the vulnerability of individuals growing in a common garden, but originating from populations along an aridity gradient, as well as individuals from the same wild populations. Intraspecific variation in water potential at which 50% of total embolism in a tissue is observed (P50 ) was explained mostly by differences between individuals (>66% of total variance) and tissues (16%). There was little between-population variation in leaf/stem P50 in the garden, which was not related to site of origin aridity. Unexpectedly, we observed a positive relationship between wild individual stem P50 and aridity. Although there is no local adaptation and only minor phenotypic plasticity in leaf/stem vulnerability in Q. douglasii, high levels of potentially heritable variation within populations or strong environmental selection could contribute to adaptive responses under future climate change.

Despite the appeal of the iso/anisohydric framework for classifying plant drought responses, recent studies have shown that such classifications can be strongly affected by a plant's environment. Here, we present measured in situ drought responses to demonstrate that apparent isohydricity can be conflated with environmental conditions that vary over space and time. In particular, we (a) use data from an oak species (Quercus douglasii) during the 2012-2015 extreme drought in California to demonstrate how temporal and spatial variability in the environment can influence plant water potential dynamics, masking the role of traits; (b) explain how these environmental variations might arise from climatic, topographic, and edaphic variability; (c) illustrate, through a "common garden" thought experiment, how existing trait-based or response-based isohydricity metrics can be confounded by these environmental variations, leading to Type-1 (false positive) and Type-2 (false negative) errors; and (d) advocate for the use of model-based approaches for formulating alternate classification schemes. Building on recent insights from greenhouse and vineyard studies, we offer additional evidence across multiple field sites to demonstrate the importance of spatial and temporal drivers of plants' apparent isohydricity. This evidence challenges the use of isohydricity indices, per se, to characterize plant water relations at the global scale.

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

Current climate change scenarios indicate warmer temperatures and the potential for more extreme droughts in the tropics, such that a mechanistic understanding of the water cycle from individual trees to landscapes is needed to adequately predict future changes in forest structure and function. In this study, we contrasted physiological responses of tropical trees during a normal dry season with the extreme dry season due to the 2015-2016 El Niño-Southern Oscillation (ENSO) event. We quantified high resolution temporal dynamics of sap velocity (Vs), stomatal conductance (gs) and leaf water potential (ΨL) of multiple canopy trees, and their correlations with leaf temperature (Tleaf) and environmental conditions [direct solar radiation, air temperature (Tair) and vapor pressure deficit (VPD)]. The experiment leveraged canopy access towers to measure adjacent trees at the ZF2 and Tapajós tropical forest research (near the cities of Manaus and Santarém). The temporal difference between the peak of gs (late morning) and the peak of VPD (early afternoon) is one of the major regulators of sap velocity hysteresis patterns. Sap velocity displayed species-specific diurnal hysteresis patterns reflected by changes in Tleaf. In the morning, Tleaf and sap velocity displayed a sigmoidal relationship. In the afternoon, stomatal conductance declined as Tleaf approached a daily peak, allowing ΨL to begin recovery, while sap velocity declined with an exponential relationship with Tleaf. In Manaus, hysteresis indices of the variables Tleaf-Tair and ΨL-Tleaf were calculated for different species and a significant difference (p < 0.01, α = 0.05) was observed when the 2015 dry season (ENSO period) was compared with the 2017 dry season ("control scenario"). In some days during the 2015 ENSO event, Tleaf approached 40°C for all studied species and the differences between Tleaf and Tair reached as high at 8°C (average difference: 1.65 ± 1.07°C). Generally, Tleaf was higher than Tair during the middle morning to early afternoon, and lower than Tair during the early morning, late afternoon and night. Our results support the hypothesis that partial stomatal closure allows for a recovery in ΨL during the afternoon period giving an observed counterclockwise hysteresis pattern between ΨL and Tleaf.

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. [ABSTRACT FROM AUTHOR]

Terrestrial water fluxes are substantially mediated by vegetation, while the distribution, growth, health, and mortality of plants are strongly influenced by the availability of water. These interactions, playing out across multiple spatial and temporal scales, link the disciplines of plant ecophysiology and ecohydrology. Despite this connection, the disciplines have provided complementary, but largely independent, perspectives on the soil‐plant‐atmosphere continuum since their crystallization as modern scientific disciplines in the late 20th century. This review traces the development of the two disciplines, from their respective origins in engineering and ecology, their largely independent growth and maturation, and the eventual development of common conceptual and quantitative frameworks. This common ground has allowed explicit coupling of the disciplines to better understand plant function. Case studies both illuminate the limitations of the disciplines working in isolation, and reveal the exciting possibilities created by consilience between the disciplines. The histories of the two disciplines suggest opportunities for new advances will arise from sharing methodologies, working across multiple levels of complexity, and leveraging new observational technologies. Practically, these exchanges can be supported by creating shared scientific spaces. This review argues that consilience and collaboration are essential for robust and evidence‐based predictions and policy responses under global change. Summary statement: Ecophysiology and ecohydrology share an interest in plant–water interactions but have largely developed and operated along distinct paths. We review the development of these fields and difficulties that emerged and argue how greater consilience can address pressing challenges under global change. [ABSTRACT FROM AUTHOR]

How plants respond physiologically to leaf warming and low water availability may determine how they will perform under future climate change. In 2015-2016, an unprecedented drought occurred across Amazonia with record-breaking high temperatures and low soil moisture, offering a unique opportunity to evaluate the performances of Amazonian trees to a severe climatic event. We quantified the responses of leaf water potential, sap velocity, whole-tree hydraulic conductance (Kwt), turgor loss and xylem embolism, during and after the 2015-2016 El Niño for five canopy-tree species. Leaf/xylem safety margins (SMs), sap velocity and Kwt showed a sharp drop during warm periods. SMs were negatively correlated with vapour pressure deficit, but had no significant relationship with soil water storage. Based on our calculations of canopy stomatal and xylem resistances, the decrease in sap velocity and Kwt was due to a combination of xylem cavitation and stomatal closure. Our results suggest that warm droughts greatly amplify the degree of trees' physiological stress and can lead to mortality. Given the extreme nature of the 2015-2016 El Niño and that temperatures are predicted to increase, this work can serve as a case study of the possible impact climate warming can have on tropical trees.This article is part of a discussion meeting issue 'The impact of the 2015/2016 El Niño on the terrestrial tropical carbon cycle: patterns, mechanisms and implications'.

Many recent studies on drought-induced vegetation mortality have explored how plant functional traits, and classifications of such traits along axes of, for example, isohydry-anisohydry, might contribute to predicting drought survival and recovery. As these studies proliferate, the consistency and predictive value of such classifications need to be carefully examined. Here, we outline the basis for a systematic classification of plant drought responses that accounts for both environmental conditions and functional traits. We use non-dimensional analysis to integrate plant traits and metrics of environmental variation into groups that can be associated with alternative drought stress pathways (hydraulic failure and carbon limitation), and demonstrate that these groupings predict physiological drought outcomes using both synthetic and measured data. In doing so, we aim to untangle some confounding effects of environment and trait variations that undermine current classification schemes, advocate for more careful treatment of the environmental context within which plants experience and respond to drought, and outline a pathway towards a general classification of drought vulnerability.

Although recent findings suggest that xylem embolism represents a significant, drought-induced damaging process in land plants, substantial debate surrounds the capacity of long-vesseled, ring-porous species to resist embolism. We investigated whether recent methodological developments could help resolve this controversy within Quercus, a long-vesseled, ring-porous temperate angiosperm genus, and shed further light on the importance of xylem vulnerability to embolism as an indicator of drought tolerance. We used the optical technique to quantify leaf and stem xylem vulnerability to embolism of eight Quercus species from the Mediterranean-type climate region of California to examine absolute measures of resistance to embolism as well as any potential hydraulic segmentation between tissue types. We demonstrated that our optical assessment reflected flow impairment for a subset of our sample species by quantifying changes in leaf hydraulic conductance in dehydrating branches. Air-entry water potential varied 2-fold in leaves, ranging from -1.7 ± 0.25 MPa to -3.74 ± 0.23 MPa, and 4-fold in stems, ranging from -1.17 ± 0.04 MPa to -4.91 ± 0.3 MPa. Embolism occurred earlier in leaves than in stems in only one out of eight sample species, and plants always lost turgor before experiencing stem embolism. Our results show that long-vesseled North American Quercus species are more resistant to embolism than previously thought and support the hypothesis that avoiding stem embolism is a critical component of drought tolerance in woody trees. Accurately quantifying xylem vulnerability to embolism is essential for understanding species distributions along aridity gradients and predicting plant mortality during drought.

Cross-boundary transfers of nutrients can profoundly shape the ecology of recipient systems. The common hippopotamus, Hippopotamus amphibius, is a significant vector of such subsidies from terrestrial to river ecosystems. We compared river pools with high and low densities of H. amphibius to determine how H. amphibius subsidies shape the chemistry and ecology of aquatic communities. Our study watershed, like many in sub-Saharan Africa, has been severely impacted by anthropogenic water abstraction reducing dry-season flow to zero. We conducted observations for multiple years over wet and dry seasons to identify how hydrological variability influences the impacts of H. amphibius During the wet season, when the river was flowing, we detected no differences in water chemistry and nutrient parameters between pools with high and low densities of H. amphibius Likewise, the diversity and abundance of fish and aquatic insect communities were indistinguishable. During the dry season, however, high-density H. amphibius pools differed drastically in almost all measured attributes of water chemistry and exhibited depressed fish and insect diversity and fish abundance compared with low-density H. amphibius pools. Scaled up to the entire watershed, we estimate that H. amphibius in this hydrologically altered watershed reduces dry-season fish abundance and indices of gamma-level diversity by 41% and 16%, respectively, but appears to promote aquatic invertebrate diversity. Widespread human-driven shifts in hydrology appear to redefine the role of H. amphibius, altering their influence on ecosystem diversity and functioning in a fashion that may be more severe than presently appreciated.

The Hawaiian Islands offer a unique opportunity to test how changes in the properties of an isolated ecosystem are propagated through the organisms that occur within that ecosystem. The age-structured arrangement of volcanic-derived substrates follows a regular progression over space and, by inference, time. We test how well documented successional changes in soil chemistry and associated vegetation are reflected in organisms at higher trophic levels-specifically, predatory arthropods (spiders)-across a range of functional groups. We focus on three separate spider lineages: one that builds capture webs, one that hunts actively, and one that specializes on eating other spiders. We analyze spiders from three sites across the Hawaiian chronosequence with substrate ages ranging from 200 to 20,000 years. To measure the extent to which chemical signatures of terrestrial substrates are propagated through higher trophic levels, we use standard stable isotope analyses of nitrogen and carbon, with plant leaves included as a baseline. The target taxa show the expected shift in isotope ratios of δ15N with trophic level, from plants to cursorial spiders to web-builders to spider eaters. Remarkably, organisms at all trophic levels also precisely reflect the successional changes in the soil stoichiometry of the island chronosequence, demonstrating how the biogeochemistry of the entire food web is determined by ecosystem succession of the substrates on which the organisms have evolved.

Current models used for predicting vegetation responses to climate change are often guided by the dichotomous needs to resolve either (i) internal plant water status as a proxy for physiological vulnerability or (ii) external water and carbon fluxes and atmospheric feedbacks. Yet, accurate representation of fluxes does not always equate to accurate predictions of vulnerability. We resolve this discrepancy using a hydrodynamic framework that simultaneously tracks plant water status and water uptake. We couple a minimalist plant hydraulics model with a soil moisture model and, for the first time, translate rainfall variability at multiple timescales - with explicit descriptions at daily, seasonal, and interannual timescales - into a physiologically meaningful metric for the risk of hydraulic failure. The model, parameterized with measured traits from chaparral species native to Southern California, shows that apparently similar transpiration patterns throughout the dry season can emerge from disparate plant water potential trajectories, and vice versa. The parsimonious set of parameters that captures the role of many traits across the soil-plant-atmosphere continuum is then used to establish differences in species sensitivities to shifts in seasonal rainfall statistics, showing that co-occurring species may diverge in their risk of hydraulic failure despite minimal changes to their seasonal water use. The results suggest potential shifts in species composition in this region due to species-specific changes in hydraulic risk. Our process-based approach offers a quantitative framework for understanding species sensitivity across multiple timescales of rainfall variability and provides a promising avenue toward incorporating interactions of temporal variability and physiological mechanisms into drought response models.

Climate, physical landscapes, and biota interact to generate heterogeneous hydrologic conditions in space and over time, which are reflected in spatial patterns of species distributions. As these species distributions respond to rapid climate change, microrefugia may support local species persistence in the face of deteriorating climatic suitability. Recent focus on temperature as a determinant of microrefugia insufficiently accounts for the importance of hydrologic processes and changing water availability with changing climate. Where water scarcity is a major limitation now or under future climates, hydrologic microrefugia are likely to prove essential for species persistence, particularly for sessile species and plants. Zones of high relative water availability - mesic microenvironments - are generated by a wide array of hydrologic processes, and may be loosely coupled to climatic processes and therefore buffered from climate change. Here, we review the mechanisms that generate mesic microenvironments and their likely robustness in the face of climate change. We argue that mesic microenvironments will act as species-specific refugia only if the nature and space/time variability in water availability are compatible with the ecological requirements of a target species. We illustrate this argument with case studies drawn from California oak woodland ecosystems. We posit that identification of hydrologic refugia could form a cornerstone of climate-cognizant conservation strategies, but that this would require improved understanding of climate change effects on key hydrologic processes, including frequently cryptic processes such as groundwater flow.

Predicting how individual taxa will respond to climatic change is challenging, in part because the impacts of environmental conditions can vary markedly, even among closely related species. Studies of chipmunks (Tamias spp.) in Yosemite National Park provide an important opportunity to explore the reasons for this variation in response. While the alpine chipmunk (T. alpinus) has undergone a significant elevational range contraction over the past century, the congeneric and partially sympatric lodgepole chipmunk (T. speciosus) has not experienced an elevational range shift during this period. As a first step toward identifying the factors underlying this difference in response, we examined evidence for dietary changes and changes in cranial morphology in these species over the past century. Stable isotope analyses of fur samples from modern and historical museum specimens of these species collected at the same localities indicated that signatures of dietary change were more pronounced in T. alpinus, although diet breadth did not differ consistently between the study species. Morphometric analyses of crania from these specimens revealed significant changes in cranial shape for T. alpinus, with less pronounced changes in shape for T. speciosus; evidence of selection on skull morphology was detected for T. alpinus, but not for T. speciosus. These results are consistent with growing evidence that T. alpinus is generally more responsive to environmental change than T. speciosus, but emphasize the complex and often geographically variable nature of such responses. Accordingly, future studies that make use of the taxonomically and spatially integrative approach employed here may prove particularly informative regarding relationships between environmental conditions, range changes, and patterns of phenotypic variation.

Use of the stable isotopes of water (δD, δ18O) to determine vegetative water sources, runoff paths, and residence times generally assumes that, other than shallow evaporative enrichment, the isotopic composition of precipitation is conserved as it travels through the subsurface to the stream channel. Here we follow rainfall through a thick (up to 25m) vadose zone of soil, saprolite, and weathered bedrock mostly composed of argillite, and underlying a steep (32°) forested hillslope. We discover a persistent structured heterogeneity of water isotopes inside the hillslope. Summer dry season causes evaporative enrichment of the soil, but not in the saprolite and weathered bedrock. In winter, the mobile water, generated by successive rainstorms with widely varying isotopic composition, mixes in the vadose zone, elevating soil and rock moisture content, and eventually recharging the groundwater with isotopically invariant water similar to the seasonally averaged rainfall. Yet throughout the winter the less mobile winter soil and rock moisture remains relatively light, and water extracted from the interior of argillite lies well to the left of the local meteoric water line. This persistently light composition of soil and rock moisture and the deviation from average meteoric values suggest that subsurface fractionation, or the inheritance of paleo-meteoric rock moisture associated with rock uplift may lead to large enduring isotopic differences between high and low mobility water. These differences suggest that the use of water isotopes as tracers must consider the possibility of subsurface isotopic evolution and the influence of exchange with more tightly held water.

A defining feature of the redwood forest in coastal California is the presence of fog in the summer months, a time when there is typically little rainfall. Our goal was to determine the role of summer fog in canopy transformation of nitrogen, nitrogen uptake by trees and photosynthesis within a coastal redwood forest ecosystem. We measured horizontal and vertical inputs of nitrogen, the isotopic composition of nitrogen in a variety of atmospheric sources (summer fog, winter rain and throughfall throughout the year), nitrogen pools (soil solution) and plant tissue (roots and foliage), as well as rates of photosynthesis and nitrogen uptake by trees. Throughfall nitrogen fluxes were greater at the forest edge compared to the interior both within the canopy (sampled 10m above-ground) and onto the forest floor (sampled 1m above-ground; P0.05). Nitrification in the forest floor, rather than the canopy, is the primary source of NO3- in these soils throughout the year. Synthesis. Summer fog provides nitrogen directly and indirectly to redwood trees, especially those at the forest edge, and affects the physiologic function of redwood trees. Summer fog provides nitrogen directly and indirectly to redwood trees, especially those at the forest edge, and affects the physiologic function of redwood trees.

Attempts to understand mechanisms underlying plant mortality during drought have led to the emergence of a hydraulic framework describing distinct hydraulic strategies among coexisting species. This framework distinguishes species that rapidly decrease stomatal conductance (gs), thereby maintaining high water potential (Px; isohydric), from those species that maintain relatively high gs at low Px, thereby maintaining carbon assimilation, albeit at the cost of loss of hydraulic conductivity (anisohydric). This framework is yet to be tested in biodiverse communities, potentially due to a lack of standardized reference values upon which hydraulic strategies can be defined. We developed a system of quantifying hydraulic strategy using indices from vulnerability curves and stomatal dehydration response curves and tested it in a speciose community from South Africa's Cape Floristic Region. Degree of stomatal regulation over cavitation was defined as the margin between Px at stomatal closure (Pg12) and Px at 50% loss of conductivity. To assess relationships between hydraulic strategy and mortality mechanisms, we developed proxies for carbon limitation and hydraulic failure using time since Pg12 and loss of conductivity at minimum seasonal Px, respectively. Our approach captured continuous variation along an isohydry/anisohydry axis and showed that this variation was linearly related to xylem safety margin. Degree of isohydry/anisohydry was associated with contrasting predictions for mortality during drought. Merging stomatal regulation strategies that represent an index of water use behavior with xylem vulnerability facilitates a more comprehensive framework with which to characterize plant response to drought, thus opening up an avenue for predicting the response of diverse communities to future droughts.

The California Mediterranean savanna has harsh summer conditions with minimal soil moisture, high temperature, high incoming solar radiation and little or no precipitation. Deciduous blue oaks, Quercus douglasii Hook. and Arn., are winter-deciduous obligate phreatophytes, transpiring mostly groundwater throughout the summer drought. The objective of this work is to fully characterize the seasonal trends of photosynthesis in blue oaks as well as the mechanistic relationships between leaf structure and function. We estimate radiative load of the leaves via the FLiES model and perform in situ measurements of leaf water potential, leaf nitrogen content, an index of chlorophyll content (SPAD readings), photosynthetic and electron transport capacity, and instantaneous rates of CO2 assimilation and electron transport. We measured multiple trees over 3 years providing data from a range of conditions. Our study included one individual that demonstrated strong drought stress as indicated by changes in SPAD readings, leaf nitrogen and all measures of leaf functioning. In the year following severe environmental stress, one individual altered foliation patterns on the crown but did not die. In all other individuals, we found that net carbon assimilation and photosynthetic capacity decreased during the summer drought. SPAD values, electron transport rate (ETR) and quantum yield of photosystem II (PSII) did not show a strong decrease during the summer drought. In most individuals, PSII activity and SPAD readings did not indicate leaf structural or functional damage throughout the season. While net carbon assimilation was tightly coupled to stomatal conductance, the coupling was not as tight with ETR possibly due to contributions from photorespiration or other protective processes. Our work demonstrates that the blue oaks avoid structural damage by maintaining the capacity to convert and dissipate incoming solar radiation during the hot summer drought and are effective at fixing carbon by maximizing rates during the mild spring conditions.

Stable isotope analysis has provided insights into the trophic ecology of a wide diversity of animals. Knowledge about isotopic incorporation rates and isotopic discrimination between the consumer and its diet for different tissue types is essential for interpreting stable isotope data, but these parameters remain understudied in many animal taxa and particularly in aquatic invertebrates. We performed a 292-day diet shift experiment on 92 individuals of the predatory mantis shrimp, Neogonodactylus bredini, to quantify carbon and nitrogen incorporation rates and isotope discrimination factors in muscle and hemolymph tissues. Average isotopic discrimination factors between mantis shrimp muscle and the new diet were 3.0 ± 0.6 ‰ and 0.9 ± 0.3 ‰ for carbon and nitrogen, respectively, which is contrary to what is seen in many other animals (e.g. C and N discrimination is generally 0–1 ‰ and 3–4 ‰, respectively). Surprisingly, the average residence time of nitrogen in hemolymph (28.9 ± 8.3 days) was over 8 times longer than that of carbon (3.4 ± 1.4 days). In muscle, the average residence times of carbon and nitrogen were of the same magnitude (89.3 ± 44.4 and 72.8 ± 18.8 days, respectively). We compared the mantis shrimps’ incorporation rates, along with rates from four other invertebrate taxa from the literature, to those predicted by an allometric equation relating carbon incorporation rate to body mass that was developed for teleost fishes and sharks. The rate of carbon incorporation into muscle was consistent with rates predicted by this equation. Our findings provide new insight into isotopic discrimination factors and incorporation rates in invertebrates with the former showing a different trend than what is commonly observed in other animals.

© 2015 McCauley et al. The common hippopotamus, Hippopotamus amphibius, transports millions of tons of organic matter annually from its terrestrial feeding grounds into aquatic habitats. We evaluated whether carbon stable isotopes (δ13C) can be used as tracers for determining whether H. amphibius-vectored allochthonous material is utilized by aquatic consumers. Two approaches were employed to make this determination: (1) lab-based feeding trials where omnivorous river fish were fed a H. amphibius dung diet and (2) field sampling of fish and aquatic insects in pools with and without H. amphibius. Lab trials revealed that fish fed exclusively H. amphibius dung exhibited significantly more positive δ13C values than fish not fed dung. Fish and aquatic insects sampled in a river pool used for decades by H. amphibius also exhibited more positive δ13C values at the end of the dry season than fish and insects sampled from an upstream H. amphibius-free reference pool. Fish sampled in these same pools at the end of the wet season (high flow) showed no significant differences in δ13C values, suggesting that higher flows reduced retention and use of H. amphibius subsidies. These data provide preliminary evidence that δ13C values may be useful, in certain contexts, for quantifying the importance H. amphibius organic matter.

Leaf hydraulic conductance (k leaf) is a central element in the regulation of leaf water balance but the properties of k leaf remain uncertain. Here, the evidence for the following two models for k leaf in well-hydrated plants is evaluated: (i) k leaf is constant or (ii) k leaf increases as transpiration rate (E) increases. The difference between stem and leaf water potential (ΔΨstem-leaf), stomatal conductance (g s), k leaf, and E over a diurnal cycle for three angiosperm and gymnosperm tree species growing in a common garden, and for Helianthus annuus plants grown under sub-ambient, ambient, and elevated atmospheric CO₂ concentration were evaluated. Results show that for well-watered plants k leaf is positively dependent on E. Here, this property is termed the dynamic conductance, k leaf(E), which incorporates the inherent k leaf at zero E, which is distinguished as the static conductance, k leaf(0). Growth under different CO₂ concentrations maintained the same relationship between k leaf and E, resulting in similar k leaf(0), while operating along different regions of the curve owing to the influence of CO₂ on g s. The positive relationship between k leaf and E minimized variation in ΔΨstem-leaf. This enables leaves to minimize variation in Ψleaf and maximize g s and CO₂ assimilation rate over the diurnal course of evaporative demand.

Obtaining the d-excess parameter from oxygen and hydrogen stable isotope composition of meteoric waters has the potential power to reconstruct changes in atmospheric water pools (e,g. sources, origins and overall balance) and the climatic conditions that prevail during surface evaporation. Recently, plant and ecosystem scientists turned their attention using d-excess information to inform questions at these scales. Here, we use the d-excess parameter to evaluate the influence of forest canopies on atmospheric humidity within a mixed evergreen forest in coastal California. We found that during the day, when transpiration was at a maximum, the d-excess of atmospheric water vapour exceeded model predictions for the background atmosphere into which the ecosystem evapotranspiration mixes. At night when transpiration was minor, the d-excess of atmospheric water vapour was on average less than model predictions for an ocean derived water vapour source. The observed diurnal fluctuations around the d-excess of the modelled background water vapour provided a strong evidence that during the day as the land surface warms and the boundary layer grows plants alter the isotope composition of atmospheric humidity via non-steady state isotope effects. In contrast, at night equilibrium isotope effects dominate as the atmosphere stabilizes. These day and nighttime fluctuations around the d-excess of ocean derived water vapour highlight the importance of plant transpiration for the isotope hydrology of near surface humidity and subsequently for the isotope composition of condensate like dew, an important water input to this ecosystem. © 2013 John Wiley & Sons, Ltd.

We investigated photosynthesis of five plant species growing in the understory at three sites (1,170-, 1,600- and 2,100-mm annual moisture inputs), along the geographical range of coastal California redwood forest, to determine whether greater inputs of rain and fog at northern sites enhance photosynthetic utilization of fluctuating light. Measurements of understory light environment and gas exchange were carried out to determine steady state and dynamic photosynthetic responses to light. Leaf area index ranged from 4.84 at the 2,100-mm site to 5.98 at the 1,170-mm site. Maximum rates of net photosynthesis and stomatal conductance (g) did not vary appreciably within species across sites. Photosynthetic induction after a change from low to high light was significantly greater in plants growing in lower light conditions regardless of site. Photosynthetic induction also increased with the rate of g in diffuse light, prior to the increase to saturating light levels. Post-illumination CO2 assimilation was the largest factor contributing to variation in C gain during simulated lightflecks. The duration of post-illumination photosynthetic activity, total CO2 assimilation per light received, and light use efficiency during simulated lightflecks increased significantly with moisture inputs in four out of five species. Increasing leaf N concentration with increasing moisture inputs in three out of five species, coupled with changes in leaf N isotopic composition with the onset of the summer fog season suggest that natural N deposition increases with rain and fog inputs and contributes to greater utilization of fluctuating light availability in coastal California redwood forests.